Topographically controlled soil moisture is the primary driver

of local vegetation patterns across a lowland region

JESPER E. MOESLUND,1,2,3,5,  LARS ARGE,2 PEDER K. BøCHER,1 TOMMY DALGAARD,3 METTE V. ODGAARD,1,3
BETTINA NYGAARD,4 AND JENS-CHRISTIAN SVENNING1
1
Ecoinformatics & Biodiversity Group, Department of Bioscience, Aarhus University,
Ny Munkegade 114, DK-8000 Aarhus C, Denmark
2
Center for Massive Data Algorithmics (MADALGO), Department of Computer Science, Aarhus University,
IT-parken, Åbogade 34, DK-8200 Aarhus N, Denmark
3
Agricultural Systems and Sustainability, Department of Agroecology, Aarhus University,
Blichers Allé 20, DK-8830 Tjele, Denmark
4
Wildlife Ecology and Biodiversity, Department of Bioscience, Aarhus University, Grenåvej 14, DK-8410 Rønde, Denmark

Citation: Moeslund, J. E., L. Arge, P. K. Bøcher, T. Dalgaard, M. V. Odgaard, B. Nygaard, and J.-C. Svenning. 2013.
Topographically controlled soil moisture is the primary driver of local vegetation patterns across a lowland region.
Ecosphere 4(7):91. http://dx.doi.org/10.1890/ES13-00134.1

Abstract. Topography is recognized as an important factor in controlling plant distribution and

diversity patterns, but its scale dependence and the underlying mechanisms by which it operates are not
well understood. Here, we used novel high-resolution (2-m scale) topographic data from more than 30500
vegetation plots to assess the importance of topography for local plant diversity and distribution patterns
across Denmark, a 43000 km2 lowland region. The vegetation data came from 901 nature conservation sites
(mean size ¼ 0.16 km2) distributed throughout Denmark, each having an average of 34 plots (five-meter
radius) per site. We employed a variety of statistical measures and techniques to investigate scale
dependence and mechanistic drivers operating within the study region. Ordinary Least Squares (OLS)
multiple regression modeling scaled at different spatial resolutions (2 3 2, 10 3 10, 50 3 50, 100 3 100 and
250 3 250 m) was used to identify the horizontal resolution yielding the strongest vegetation–topography
relationships. Using data scaled at this resolution, we quantified local (within-site) and regional (among
sites) relationships between elevation, mechanistic topographic factors (slope, heat index, potential solar
radiation, wind exposure, wetness index) and 10 vegetation measures representing species composition,
richness and functional composition (average plant preferences along key environmental niche axes). We
also investigated how overall site-level environmental characteristics affect the strength of these local
relationships. Topography exerted the strongest effects at the 10 3 10 m horizontal resolution scale.
Elevation exerted the strongest influence on vegetation, followed by slope and wetness. Topography
generally affected all vegetation measures and exhibited the strongest local relationships with the main
species-compositional gradient, the main functional gradient and the plant’s average soil moisture
preference. The strength of these relationships was strongly influenced by habitat and site-level average
moisture conditions, with the strongest relationships found in wet habitats. Our findings show that fine-
scale topography can strongly influence local vegetation patterns across a wide range of habitat types even
in low-relief lowland regions. Notably, topography exhibited a consistently strong relationship with the
main local floristic and functional compositional gradients. While a plurality of underlying mechanisms
may contribute to the relationship between topography and vegetation patterns, topographically
controlled soil moisture exerts primary control on the relationship.

Key words: Ellenberg indicator values; Europe; NATURA 2000; plant community assembly; plant diversity; solar
radiation; spatial scale; species richness; topographic wetness index; wind.

Received 12 April 2013; revised 31 May 2013; accepted 3 June 2013; published 31 July 2013. Corresponding Editor: D. P.
C. Peters.

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Copyright: Ó 2013 Moeslund et al. This is an open-access article distributed under the terms of the Creative Commons
Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the
original author and source are credited. http://creativecommons.org/licenses/by/3.0/
5
Present address: Wildlife Ecology and Biodiversity, Department of Bioscience, Aarhus University, Grenåvej 14, DK-8410
Rønde, Denmark.
  E-mail: jesper.moeslund@biology.au.dk
INTRODUCTION
Among the many theories on what determines
local species diversity and distribution patterns
(Zobel 1997, Grace 1999, Leibold et al. 2004),
niche theory serves as a key explanatory frame-
work (Shmida and Wilson 1985). Topography
tends to influence the characteristics of the local
niche hyperspace for a given plant community
due to its pervasive effects on hydrology (Vivian-
Smith 1997, Silvertown et al. 1999, Økland et al.
2008), temperature and light availability (Boyko
1947, Oke 1987, Bennie et al. 2008, Lenoir et al.
2013), edaphic factors (Chen et al. 1997, Gould
and Walker 1997, Fu et al. 2004) and wind
exposure (Knapp 1985, Chen et al. 1997, Lortie
and Cushman 2007, Li et al. 2009). Most studies
on the ecological influence of topography have
addressed these factors in mountainous areas
(Sanders and Rahbek 2012) where elevation
exerts a strong and direct influence on temper-
ature (Calvert 1990), creating pronounced gradi-
ents in plant diversity and distribution patterns
(Grytnes 2003). While previous studies have
shown that topography affects local vegetation
patterns in lowland regions (e.g., Svenning et al.
2009), the exact role of topography in lowland
ecology is not well understood.
Elevation is itself an indirect environmental
factor with no direct effects on plants; it affects
vegetation patterns through other more direct
factors (Guisan and Zimmermann 2000). Most
studies addressing the local effects of elevation
on plant communities in lowland areas have
focused on wet habitats (Vivian-Smith 1997,
Silvertown et al. 1999, Økland et al. 2008,
Raulings et al. 2010, Moeslund et al. 2011). In
these settings, even cm-scale topographic hetero-
geneity can cause major differences in water
availability, salinity and flooding frequency that
may in turn impact species richness and compo-
sition (Vivian-Smith 1997, Silvertown et al. 1999,
Økland et al. 2008, Moeslund et al. 2011). Terrain
slope and aspect may also affect local plant
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MOESLUND ET AL.
communities (Pausas and Carreras 1995, Olivero
and Hix 1998). North- and south-facing slopes for
example may experience temperature-driven
variation in nutrient and moisture conditions
even in lowland areas (Boyko 1947, Olivero and
Hix 1998, Bennie et al. 2008). Wind exposure is
also a topographically dependent factor that may
influence local vegetation patterns in lowlands
(Chen et al. 1997, Lortie and Cushman 2007, Li et
al. 2009). This factor may exert contingent effects
on soil grain size characteristics (Barbour et al.
1974) and on local temperature and moisture
conditions (Knapp 1985). Although, many stud-
ies have emphasized the importance of topo-
graphically controlled soil moisture in wet
habitats only a few have considered this issue
in dry habitats. In spite of their limited number,
these have found compelling evidence of the role
of soil moisture and terrain-determined surface
run-off patterns in structuring local vegetation
patterns also in dry areas (Augustine 2003, Araya
et al. 2011). Most studies addressing topogra-
phy’s influence on lowland vegetation patterns
have included only one or a few local study sites.
The general role of topography in determining
vegetation patterns across large, non-mountain-
ous regions is therefore still poorly understood.
Recent advances in Light Detection and Rang-
ing (LiDAR) remote-sensing technology have
enabled scientists to develop highly accurate
(10–15 cm vertical resolution) digital elevation
models (DEM) over large areas at high spatial
resolution (1–2 m; Vierling et al. 2008). In spite of
coverage gaps spanning large areas of the Earth’s
surface, LiDAR availability is rapidly expanding
and has now begun to offer a comprehensive
view of the role played by topography in
determining local vegetation patterns. In this
study, we combined novel, high-resolution topo-
graphic information with an unprecedented
volume of vegetation data to investigate rela-
tionships between topography and local plant
species diversity over the large lowland region of
Denmark (;43000 km2). We specifically assessed
2 July 2013 v Volume 4(7) v Article 91

the scale dependence of topographic variables
and the underlying mechanisms by which they
operate. Our specific study objectives were to
determine (1) the spatial scale at which relation-
ships between topography and local plant
diversity and distributions operate, (2) the extent
to which topography controls local plant species
diversity and distribution patterns in natural and
semi-natural areas across the lowland study area,
(3) the mechanisms through which topography
influences vegetation patterns, and (4) site-level
environmental factors that may affect the
strength and nature of local (within-site) vegeta-
tion–topography relationships.
METHODS
Study sites
This study included 901 sites (0.16 km2 on
average). A majority of the sites are located
within special areas of conservation protected
under the European Union’s Habitats Directive
(Council of the European Communities 1992).
Each site covered one or more of the natural and/
or semi-natural NATURA 2000 habitat types
(Commission of the European communities
2002) ranging from dry dunes and grassland to
wet flushes and mires. The sites are distributed
throughout Denmark (Fig. 1A) and their plant
species compositions were recorded by the
Danish National Monitoring and Assessment
Programme for the Aquatic and Terrestrial
Environment (NOVANA; Svendsen and Norup
2005; data available via www.naturdata.dk).
Table A1 (Appendix A) gives detailed descrip-
tions of the habitat types included in this study.
Vegetation and topographic data
The data used in this study consisted of
vegetation records from randomly selected sites
that included 20 to 60 circular sample plots (5-m
radius) georeferenced at 2–5 m precision. Vege-
tation sampling procedures included recording
the type and abundance of all vascular plant
species rooted within the 0.25 m2 quadrat at the
center of each plot (Svendsen and Norup 2005).
The vascular plant species data used in this study
was collected from 2004 to 2009 from 37422 plots.
Following initial extraction of the data from the
NOVANA database, we removed duplicates and
entries having incomplete or erroneous values
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MOESLUND ET AL.
(e.g., obvious identification errors). The final
dataset contained data from 30544 plots distrib-
uted among the 901 monitoring sites (Fig. 1A),
and included a total of 1262 vascular plant
species, subspecies and varieties.
The DK-DEM (‘‘Danmarks højdemodel’’) ele-
vation model provided LiDAR based elevation
data of the study region at 1.6 m horizontal
resolution and 0.10–0.15 m vertical precision
(Knudsen et al. 2008). In addition to the first
order horizontal resolution of 1.6 m, we aggre-
gated (mean) the model to horizontal resolution
scales of 9.6, 49.6, 100.8 and 249.6 m. For
simplicity, these resolutions are referred to
according to respective orders of magnitude as
the 2, 10, 50, 100 and 250 m resolutions in the text
that follows. Terrain slope, a heat index, potential
solar radiation, wind exposure and a topograph-
ical wetness index (TWI) were calculated at each
horizontal resolution scale in order to evaluate
the impact of both direct and indirect topograph-
ic factors. These were used as explanatory
variables in the statistical analyses conducted in
this study. The text below describes each variable
except the elevation variable (also see Appendix
B).
Slope, aspect and potential solar radiation
layers were computed using the ‘Slope’, ‘Aspect’
and ‘Area Solar Radiation’ tools in the ArcGIS 10
Spatial Analyst module (Environmental Systems
Research Institute, Redlands, California, USA).
The default settings were used for each tool
except in the case of the Area Solar Radiation. For
this tool we selected the time configuration
‘whole year (2009) with monthly interval’. As
circular variables do not fit into the statistical
analyses conducted here and to represent heat
balance, the aspect variable was converted into a
heat index using the formula (1  cos(h  45))/2,
where h represents the aspect in degrees
(McCune and Keon 2002).
The ‘Hillshade’ tool (Spatial Analyst, ArcGIS
10, Environmental Systems Research Institute,
Redlands, California, USA) was used to represent
topographically controlled wind exposure ac-
cording to methods described by Mikita and
Klimánek (2010). See Appendix B for details.
To represent topographically controlled effects
on soil moisture, we calculated a topographic
wetness index (TWI) as a proportional measure
of water retention for a given area (Hengl and
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 MOESLUND ET AL.

Fig. 1. The distribution of the 901 natural and semi-natural study sites located throughout Denmark underlain
by a digital elevation model used in this study. Panel (A) shows the general strength of the vegetation–
topography relationships (average Spearman r 2 values for all vegetation–topography correlations) at each site.
Panel (B) and (C) show the average Ellenberg indicator values (Ellenberg et al. 2001) for soil moisture and
temperature, respectively.

Reuter 2009). Appendix B explains the TWI
calculation procedures in detail.
Data and analysis
Local within-site relationships: preparation of
data.—We selected 10 measures of plant diversity
and composition to assess topographic effects on
local vegetation patterns among sample plots
within each study site (i.e., these measures were
used as response variables). These measures
included species richness, a measure of species
composition, seven measures of functional com-
position (i.e., species preferences along key
environmental niche axes), and a synthetic
measure of functional composition based on the
previous measures of functional composition.
The species richness measure represented the
number of taxa per plot counting all species,
subspecies and varieties consistently identified
within the plot for all years of observation. The
far majority of the observations used were at the
level of species. Plot scores on the first axis of a
Detrended Correspondence Analysis (DCA) of
the species composition in the plots within a site
provided a measure of the plots’ position on the
main local floristic gradient (cf. Ejrnæs and
Bruun 2000).
Ellenberg indicator values (EIV; Ellenberg et al.
2001) were used to indicate plant species’ pre-
ferred position along key environmental niche
axes. Using these indicator values to identify
relationships between vegetation and environ-
ment is a well-known and widely used method
in vegetation ecology (Diekmann 2003). Here, we
calculated weighted average (by abundance)
Ellenberg indicator values per plot and used these
as measures of functional composition along
individual key environmental niche axes. We
included values for light (EIVlight), temperature
(EIVtemp), continentality (EIVcont), soil moisture
(EIVmoist), soil reaction (EIVreac), soil nitrogen
content (EIVnitr) and soil salinity (EIVsalt). When-
ever possible, we used the EIV values from Hill et
al. (1999), since these offered a better fit for the
Danish flora. Ellenberg et al. (2001) for example
assign the grasses Leymus arenarius (L.) Hochst.
and Festuca arundinacea Schreb. an EIVsalt value of
1 (non-saline to periodically slightly saline)
whereas Hill et al. (1999) assigned the same
species an EIVsalt value of 3 (coastal, slightly
saline soils). These grass species are found almost
exclusively within coastal areas in Denmark
where soil salinity is undoubtedly higher than
that suggested by an EIVsalt value of 1 (i.e., the
original values given in Ellenberg et al. 2001). The
wetland species Silene flos-cuculi (L.) Greuter &

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Burdet serves as an additional example of the
higher specificity of the Hill et al. (1999) indicator
values. Ellenberg et al. (2001) assigned this species
an EIVmoist value of 5 (dampness indicator
species) whereas Hill et al. (1999) assigned it a
value of 7 (wet-site indicator species). The latter
value more accurately reflects this species’ prefer-
ence for wetlands in Denmark (see Hill et al. 1999
for further details on indicator values). Values for
taxa not found in Hill et al. (1999) were taken from
Ellenberg et al. (2001), or if unavailable therein,
from Landolt et al. (2010). Values taken from
Landolt et al. (2010) were adjusted to match
Ellenberg indicator value ranges (i.e., maximum
Landolt indicator values correspond to maximum
EIVs and the same for minimum and intermediate
values). For taxa not listed in the aforementioned
sources, we estimated indicator values primarily
using distribution information from Danish floris-
tic handbooks (Appendix A: Table A3). A
correlation analysis showed that this approach
did not introduce bias to our data: the mean plot
EIVs used here correlates highly with mean plot
EIVs based solely on data from the literature
mentioned above (Pearson’s r . 0.99 for almost all
EIVs, Appendix A: Table A4). Subspecies and
varieties were treated as unique taxa given that
they typically exhibit divergent environmental
preferences. If a subspecies or variety did not
consistently appear each year in a given plot,
records for these organisms were combined with
those belonging to the taxonomic rank immedi-
ately above. Together, these methods allowed us
to assemble the most accurate measures of
ecological preference (average indicator values)
for the taxa present in each plot.
Plot scores on the first axis of a Principal
Component Analysis (PCA) calculated on the
seven average Ellenberg indicator values for all
plots within a site represented the plots’ position
on the main local functional gradient, and were
used as a synthetic measure of functional
composition.
Local within-site relationships: analysis proce-
dure.—We initially used ordinary least square
(OLS) multiple regression modeling to explore
the relative effects of various horizontal resolu-
tion scales exerted on vegetation–topography
relationships. This analysis included 50 different
models for each of the 901 study sites, namely
regressions of the ten response variables against
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MOESLUND ET AL.
all topographic variables at the five different
horizontal resolution scales (45050 models total
for all sites). The 10-m horizontal resolution scale
revealed the strongest links between topography
and vegetation (see Results). All further statistical
analyses were therefore conducted using this
scale of horizontal resolution.
We implemented both parametric and non-
parametric models to assess local relationships
between vegetation and topography. Parametric
tests included multiple OLS regressions of each
response variable against the explanatory vari-
ables for each study site (9010 local models in
total). Non-parametric tests included Spearman
correlation statistics calculated between each pair
of response and explanatory variable (60 pairs,
54060 correlation coefficients total). The results
identified elevation as one of the most important
topographic variables influencing plant diversity
and distribution patterns (see Results). Due to the
influence of elevation on all the other topograph-
ic variables, we removed the elevation variable
and re-calculated local models to determine
relationships between vegetation and the remain-
ing, more mechanistic variables.
Cross-regional relationships (all plots).—In order
to examine regional vegetation–topography rela-
tionships, we performed a multiple OLS regres-
sion of each response variable against all
explanatory variables and the primary habitat
type using the full set of plot data from all sites
combined (10 regional models). For these models,
the first axis of a DCA and a PCA including all
plots from the regional dataset represented the
main regional floristic and functional composi-
tion. To avoid bias in DCA results introduced by
rare species entries, we omitted plots having less
than five species. The regional models were also
run without habitat type included in order to
evaluate the effect of habitat type.
Calculations of Moran’s I for OLS model
residuals revealed a small degree of spatial
autocorrelation within regional models on rela-
tively short distance scales. To address this issue,
we calculated a series of analogue Linear Mixed
Effects (LME) models (Zuur et al. 2009) treating
site as a random effect. This procedure removed
the spatial autocorrelation giving Moran’s I
,j60.1j for all model residuals except for the
EIVnitr and EIVreac models, for which the Moran’s
I still exhibited low values of 0.3–0.45 in the first
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distance class (0–20 km). For the regional models,
we therefore report P-values from LME analyses
rather than from the OLS models noting that the
two approaches identified the same factors as
having the strongest influence on vegetation.
Site effects on local relationships.—We assembled
a number of measures of the overall environment
at each site. These included the main habitat type
according to NATURA 2000 habitat codes (Table
A1 and A2, Appendix A), as well as site means
and standard deviations for the seven average
Ellenberg indicator values and for all plot-level
topographic explanatory variables. Fig. 1b and 1c
show examples of the average conditions and
variability in conditions at sites throughout the
study region. To determine whether site-level
environmental factors affect the strength of local
(within-site) vegetation–topography relation-
ships, we regressed the squared Spearman
correlation coefficients (r 2; following Healey
2006) for five correlation pairs against all of the
site-level environmental descriptors using OLS
multiple regression models. The five correlation
pairs included: (1) the strongest correlation pair
at each site, (2) the DCA first axis plot scores’
correlation with elevation, (3) the PCA first axis
plot scores’ correlation with elevation, (4) species
richness’ correlation with elevation, and (5) the
EIVmoist-TWI correlation. The last four pairs
cover the most important vegetation measures
and topographic variables.
Prior to these analyses, all explanatory vari-
ables (except for habitat type) were checked for
multicollinearity. Those having a tolerance
(Quinn and Keough 2002) below 0.1 were not
included in the analysis. Hence, the following
measures (where l indicates a mean and r
indicates a standard deviation) were used as
explanatory variables: EIVlight (l, r), EIVtemp (l,
r), EIVcont (l, r), EIVmoist (l, r), EIVreac (r),
EIVnitr (r), EIVsalt (r), potential solar radiation
(l), wind exposure (l), elevation (l), heat index
(l, r) and TWI (l ,r). Moran’s I calculations
using model residuals (Legendre and Legendre
1998) demonstrated the absence of spatial auto-
correlation for all 60 models.
Statistical issues and software.—We applied log-
or square-root transformations as needed to
fulfill assumptions regarding the normal distri-
bution of model residuals and the validity of
associated P-values. All statistical analysis pro-
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MOESLUND ET AL.
cedures were performed using the R (ver. 2.13.2
for 64-bit operating systems) statistical program-
ming environment (R Development Core Team
2011).
RESULTS
The strength of the vegetation–topography
relationships varied with the scale of horizontal
spatial resolution, with the highest adjusted
coefficients of determination (R 2adj) at the 10 3
10 m resolution and the lowest R 2adj at the 250 3
250 m resolution (Fig. 2).
Topography explained a significant proportion
of variation in all measures of plant diversity and
distribution within sites (local models) as well as
across Denmark (regional models). All the 60
within-site correlation pairs between plant and
topography measures were statistically signifi-
cant in .5% of the sites (i.e., at frequencies above
those expected to randomly occur, Table 1). The
average strength (r 2) of the strongest Spearman
correlation in each site was 38% (616% standard
deviation; see Fig. C1, Appendix C for examples),
indicating a generally moderately strong topo-
graphic effect on the local vegetation. Seventy
percent of the vegetation–topography relation-
ships were significantly weaker across the whole
study region (regional models) than within sites
(local models, Fig. 3).
Within sites, the relationship between soil
moisture preference (EIVmoist) and topography
was always among the strongest (Table 1, Fig. 3).
In addition, strong and consistent local relation-
ships between topography and the most impor-
tant local floristic and functional gradients (DCA
and PCA first axis plot scores, respectively) were
also evident (Table 1, Fig. 3). While all topo-
graphic variables had statistically significant
relations to the plant measures, elevation and
slope had the strongest effects (Table 1). Similar
to the local relationships, EIVmoist also exhibited
the strongest relationship with topography
across the whole study region (Fig. 3). In
contrast, relationships between the main regional
floristic and functional gradients (DCA and PCA
first axis plot score) and topography were much
weaker than their local relationships (Fig. 3).
The habitat-specific patterns in the local
vegetation–topography relationships were large-
ly consistent with the overall local patterns just
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 MOESLUND ET AL.

Fig. 2. A summary of the R 2adj values for the local (within-site) OLS multiple regression models linking each of
the ten response variables to all topographic variables at the five resolutions. Upper extreme, upper quartile,
median, lower quartile and lower extreme are represented for each box. Circles mark the means. Different letters
denote significantly different average R 2adj values (Student’s t-test, P , 0.05). The number of statistically
significant models (out of 9010) for each resolution scale is shown next to each bar.

Table 1. Average squared Spearman’s correlation coefficients (r 2) for topography and vegetation within each of
the 901 natural and semi-natural sites distributed throughout Denmark.

Vegetation Potential Topographic

measure Elevation Heat index Slope solar radiation wetness index Wind index
EIVcont 0.098(6)A,DE 0.046(6)D,C 0.061(6)B,D 0.062(6)BC,C 0.061(6)B,D 0.051(6)C,CD
EIVlight 0.101()A,E 0.050(6)D,C 0.071()B,BCD 0.058(6)C,C 0.073(þ)B,C 0.057(þ)C,CD
EIVmoist 0.147( )A,B 0.056(6)E,A 0.096(2 2)C,A 0.073(2)D,A 0.109(11)B,A 0.065(2)D,AB
EIVnitr 0.098()A,E 0.046(6)D,BC 0.076(6)B,BC 0.063(6)C,BC 0.069(6)B,C 0.056(6)C,CD
EIVreac 0.122( )A,C 0.048()E,ABC 0.075(6)B,B 0.057(6)DE,C 0.067(þ)C,C 0.056(6)D,C
EIVsalt 0.116()A,E 0.047(6)D,C 0.070(6)B,CD 0.056(6)C,C 0.076(6)B,C 0.053(6)C,CD
EIVtemp 0.084(6)A,F 0.046(6)E,BC 0.064(6)B,D 0.057(6)CD,C 0.060(6)BC,D 0.050(6)DE,D
DCA 0.166(6)A,A 0.054(6)D,A 0.098(6)B,A 0.072(6)C,A 0.101(6)B,B 0.072(6)C,A
PCA 0.147(6)A,AB 0.053(6)D,A 0.091(6)B,A 0.068(6)C,AB 0.091(6)B,B 0.064(6)C,AB
SpRich 0.118(6)A,CD 0.053(6)D,AB 0.092(11)B,A 0.067()C,AB 0.068()C,C 0.059(6)D,BC
Notes: The general direction of the relationships is given by plusses or minuses. Two minuses: negative relationships
outnumbered positive relationships by .100%, one minus: by .25%, plus/minus: no clear direction, one plus: positive
relationships outnumbered negative relationships by .25%, two plusses: by .100%. All relationships were statistically
significant in more than 5% of the cases (P , 0.05). Details about the general direction and the statistical significance of the
relationships is given in Table C1, Appendix C. Values were compared by row (superscripted letters before comma) and by
column (superscripted letters after comma) with different letters indicating significantly different relationships (Student’s t-test,
P , 0.05). Bold values indicate the best and second best topographic determinants of the vegetation measure in question
according to Student’s t-test results. Underlined values refer to the two vegetation measures that are the most strongly
controlled by the topographic variable in question (according to Student’s t-tests). SpRich: species richness. DCA: plot score on
the DCA first axis. PCA: plot score on PCA first axis. EIV: plot-level average Ellenberg indicator value for subscript variables.
The subscripts indicate variables as follows: salt, soil salinity; moist, soil moisture; nitr, soil nitrogen; cont, continentality; reac,
soil reaction; and temp, temperature.

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 MOESLUND ET AL.

Fig. 3. Coefficients of determination (R 2adj) for each vegetation measure from the regional-scale models
(combining plot data excluding habitat type from all 901 study sites across Denmark) and from the within-site
(local) models (including elevation as explanatory variable). For the local models 1r error bars are shown.
Abbreviations next to each bar identify statistically significant (P , 0.05) explanatory variables. Ele, Elevation;
Heat, Heat Index; Slo, Slope; Rad, Potential Solar Radiation; TWI, Topographic Wetness Index; Wind, Wind
Index. Asterisks indicate that the local model mean R 2adj was significantly greater (Student’s t-test, P , 0.05) than
that of the regional model. See Table 1 for the remaining abbreviations and Fig. C2 and C3 (Appendix C) for
results for the regional models including habitat as explanatory variable and for the local models excluding
elevation as explanatory factor. The local-model-R 2adj shown here (i.e., including elevation as explanatory
variable) were in all cases significantly different from models not including elevation (Student’s t-test, P , 0.05).

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 MOESLUND ET AL.

Fig. 4. Habitat-specific average Spearman correlation (vegetation–topography) r 2 values for (A) the two
strongest topographic determinants of vegetation measures and (B) the two vegetation measures exhibiting the
strongest topographic relationships. Panel (C) shows habitat specific r 2 values for pairs of variables exhibiting (1)
the highest overall correlation and (2) the highest correlation between functional variables (i.e., excluding
elevation and slope, given their mainly indirect effects). The color codes from panel (A) and (B) are used in (C) to
denote the correlation in question (e.g., blue-blue ¼ TWI-EIVmoist). A plus denotes a positive relationship, a minus
denotes a negative one. See Table 1 for abbreviations.

described, but with some variability at least
partially related to specific environmental condi-
tions in certain habitats, e.g., the uniquely strong
relationship between salt tolerance (EIVsalt) and
topography in salt meadows (Fig. 4). When
considering only the more functional variables
(i.e., not elevation and slope), the strongest local
relationships in all habitats always included at
least one of the hydrology-related variables, TWI
or EIVmoist, and often both (Fig. 4C).
Environmental site characteristics such as
habitat type affected the strength of the local
vegetation–topography relationships, with the
strongest relationships in wet habitats (Table 2).
Correspondingly, the site-means of TWI and
secondarily EIVmoist had the clearest effects on
the local relationships (Table 3).
DISCUSSION
The presented cross-scale and cross-habitat
analysis of vegetation–topography relationships
based on .30500 plots and very high-resolution
topographic data provides the first geographi-

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 MOESLUND ET AL.

Table 2. An overview of the NATURA 2000 habitat slope and TWI having the strongest relation-
types as well as their frequencies observed from the ships. Hydrology-related variables exhibited the
901 natural and semi-natural study sites located strongest correlation coefficients among the more
throughout Denmark. functional topographic and plant-related vari-
ables across all habitat types. Topography,
No. sites
Habitat type included Average R 2adj furthermore, exerted the strongest influence on
Salt meadows 95 0.26A local vegetation–topography relations in wet
Xeric calcareous grasslands 6 0.20ABC habitats, although it was also important in dry
Humid dune slacks 41 0.20B habitats.
Acidic bogs 78 0.19B
Grasslands (unspecified) 31 0.19BC
Molinia meadows 58 0.17BC Scale effects on vegetation–topography
Calcareous fens 160 0.16BC
Wet heaths 39 0.16BC relationships
Calcareous grasslands 80 0.15BC Spatial scale affects ecological patterns and
Pioneer communities 11 0.15BCD
Forests 13 0.15BCD processes (Levin 1992). However, multiscale
Acidic grasslands 96 0.15C ecological studies are relatively rare (Dalgaard
Calcareous dunes 68 0.14C
Xeric heaths 89 0.10D
et al. 2003, Moser et al. 2007), probably due to the
Relatively dry acidic dunes 65 0.09D lack of high-resolution data covering large
Notes: See Table A1 and A2, Appendix A for further details. geographic areas. We found that the topographic
The habitat-specific average R 2adj values were calculated from influence on local plant diversity and distribution
OLS multiple regression models running each vegetation
measure against all the topography measures for local sites.
patterns became less evident at horizontal
The R 2adj values represent the general strength of the resolutions coarser than 10 m. Hence, our multi-
vegetation–topography relationships for a given habitat. scale modeling suggests that the mechanisms
Different letters indicate significantly different mean R 2adj
values (Tukey’s Honestly Significant Difference test, P , 0.05). underlying the local vegetation–topography re-
lationships presented here operate primarily at
the 10 m scale. The weaker vegetation–topogra-
cally comprehensive assessment of the impor- phy relationships of the 2 3 2 m models may,
tance of topography as a determinant of local however, reflect incomplete sampling: a 2 3 2 m
plant diversity and distribution patterns. Topo- area located in the center of the plot would not
graphic variables had the strongest explanatory necessarily capture the topographic variation
power in models scaled at 10 3 10 m horizontal affecting the vegetation in a larger vegetation
resolution and the weakest in models at 250 3 sample-plot like the ones used here. That local
250 m. All topographic variables were shown to (within-site) vegetation–topography relation-
influence vegetation patterns, with elevation, ships were generally stronger than cross-region

Table 3. Results from OLS multiple regression models linking the correlation strength of five selected correlation
pairs to site-level environmental variables for the 901 study sites located throughout Denmark.

Environmental variables
2
Response variable Full model R adj Mean TWI SD of EIVmoist Mean EIVtemp
Strongest correlation pair at each site 0.14 0.18 0.11 0.14
EIVmoist-TWI 0.10 0.06 0.31 0.10
DCA-elev 0.10 0.24 0.09 0.07
PCA-elev 0.09 0.14 0.07 0.07
SpRich-elev 0.06 0.24 0.07 0.15
Mean absolute regression coefficient 0.17 0.13 0.11
Notes: The five response variables are the strength (r 2) of (1) the strongest correlation pair (between a vegetation and a
topography variable) within each site, and (2–5) four within-site Spearman correlation pairs selected to cover the most
important vegetation measures and topographic variables. For each response variable, the R 2adj is given for the full (also
including environmental variables not shown here) multiple regression model along with standardized regression coefficients
for each of the three most important (see below) environmental variables. The mean absolute standardized regression
coefficient for the five response variables represents the general importance of each of the environmental variables in
determining the strength of the local (within-site) vegetation–topography relationships. TWI, topographic wetness index. See
Table 1 for the remaining abbreviations. For full results including all environmental variables used in this study, see Table C2,
Appendix C.

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relationships, probably reflects that other factors
such as climate become more important for
vegetation patterns at regional extents (Levin
1992).
Topographic control of vegetation patterns
Generally topography was important for local
plant diversity and distribution patterns. It
consistently influenced the plant’s average pref-
erence for moisture (EIVmoist), resembling previ-
ous studies, which have observed strong
relationships between local topography and the
soil moisture affinity of different plant species
(Vivian-Smith 1997, Silvertown et al. 1999, Øk-
land et al. 2008, Raulings et al. 2010). Topography
was also strongly and consistently linked to the
most important local floristic and functional
gradients (DCA and PCA), as expected from
niche-based species sorting (Leibold et al. 2004).
Other factors such as dispersal, history, compe-
tition, facilitation, and stochasticity have also
been suggested as important for local plant
diversity (Wilson 2011). These factors may
explain some of the variation that topography
could not explain. However, the aforementioned
clear link between topography and the main
floristic gradient suggests that topographically
controlled niche-based sorting generally is the
predominant driver of local plant community
assembly across habitat types in the lowland
study region.
Topography was also consistently, but less
strongly linked to local species richness patterns,
especially in salt meadows, unspecified grass-
lands, forests and xeric heaths. These results
demonstrate that topography structures local
plant species richness patterns not only in saline
(Moeslund et al. 2011) and freshwater wetland
plant communities (Silvertown et al. 1999, Øk-
land et al. 2008, Raulings et al. 2010), but also in
other habitat types. The present study is the first
to identify a link between local species richness
and topography in dry habitats generally. Be-
sides soil moisture, other topographically depen-
dent features such as soil pH and fertility may
explain this finding: in the current study,
topography influenced average plant preferences
for soil pH and nitrogen in a manner similar to
how it affects species richness in xeric heaths and
unspecified grasslands (not shown). These topo-
graphic effects on local richness patterns could
v www.esajournals.org
MOESLUND ET AL.
thus reflect productivity- and/or species-pool-
related mechanisms (Zobel 1997, Mittelbach et al.
2001).
Underlying mechanisms for vegetation–topography
relationships
In this study, elevation itself exerted the
strongest influence on local vegetation patterns,
followed by slope, TWI, potential solar radiation,
wind exposure and heat index. The role of
elevation in a low relief region (Denmark’s
highest point: 171 m above sea level) is surprising
since altitude is mostly considered to influence
vegetation patterns through temperature effects
as seen in mountains (Grytnes 2003, but see
Lenoir et al. 2013). Broadly, lowland areas
experience only short temperature gradients
given standard temperature lapse rates (Calvert
1990, Sanders and Rahbek 2012). The relative
importance of elevation evident in the current
study probably reflects: (1) its effect on niche
gradients important to plant species, (2) that
these gradients and their role for plant growth
may vary among sites and (3) that the nature of
such gradients may be complex due to geo-
graphic factors. Elevation represents both slope
(Burbank 1992) and soil moisture (TWI; Wilson
and Gallant 2000). Furthermore, elevation corre-
lates with soil salinity in coastal settings (Moe-
slund et al. 2011) and with pH in areas underlain
by limestone, which is common in many areas of
Denmark (Thomsen 1995).
Studies of plant diversity especially those
conducted locally have treated slope as a
predictive variable (Gottfried et al. 1998, Jones
et al. 2011). This factor typically explains a
relatively high proportion of the observed vari-
ation in vegetation patterns (Luoto et al. 2002,
Bennie et al. 2006). The present research therefore
agrees with previous studies. Resembling eleva-
tion, slope is a non-functional variable (Guisan
and Zimmermann 2000). It may capture eleva-
tion variation within an area and thus relate to
the potential microhabitat variation occurring
here. Given that such variation and species
richness is often tightly linked (Vivian-Smith
1997), the positive correlation between slope and
species richness in this study supports this
interpretation. Slope also affects the amount of
solar radiation received (Gottfried et al. 1998,
Bennie et al. 2008), soil moisture (Wilson and
11 July 2013 v Volume 4(7) v Article 91

Gallant 2000) and parameters such as erosion
rates, litter accumulation and biological dynam-
ics (Troeh and Thompson 2005, Gomes de Freitas
et al. 2012). Together, these factors may explain
why slope was one of the most important factors
for local vegetation patterns.
TWI was the third strongest variable affecting
plant diversity and distribution patterns and the
second strongest determinant of plant preferenc-
es for moist (EIVmoist). TWI has been shown to
provide fairly accurate local estimates of the
average EIV for moisture (Kopecký and Čı́žkova
2010) as well as of actual measurements of
groundwater and soil moisture levels (Sørensen
et al. 2006). Numerous studies have demonstrat-
ed the role of soil moisture in determining plant
species distributions and community structure
within single habitats and/or small areas (e.g.,
Økland et al. 2008, Raulings et al. 2010). Silver-
town et al. (1999) and Araya et al. (2011) for
example showed the influence of hydrologic
factors in 2–18 meadow sites in England and 8
fynbo sites in South Africa. To our knowledge, no
previous study has investigated the role of
topographically controlled soil moisture across
a wide range of habitats and sites in order to
determine its general importance for local vege-
tation patterns. Our results indeed show that
topographically controlled soil moisture exerts a
strong and ubiquitous influence on these pat-
terns. TWI was the most important functional
variable affecting local vegetation patterns across
all habitats. This suggests that local topography–
soil moisture dynamics serve as the primary
mechanism underlying topographic control of
local vegetation patterns across a number of
different habitat types, and over large areas.
Hence, the topographically affected niche axes
generally important for local plant community
assembly seems to be soil moisture related across
habitat types in the study region.
Results from this study indicate that potential
solar radiation does not strongly influence local
plant diversity and distributions. Nevertheless, it
did influence vegetation patterns in xeric calcar-
eous and acidic grasslands, indicating that as a
community-assembly mechanism, it primarily
operates in relatively dry semi-natural areas
(Ejrnæs and Bruun 2000). Hence, under certain
circumstances, solar radiation may contribute
(along with hydrologic factors) to topography’s
v www.esajournals.org
MOESLUND ET AL.
influence on local vegetation patterns. Potential
solar radiation exerted a stronger influence than
the heat index (aspect) whereas it remained less
than that of the slope variable, in spite of the fact
that both aspect and slope is considered in the
calculation of potential solar radiation. Given the
contributions of these factors, potential solar
radiation should theoretically exert greater influ-
ence on vegetation patterns than that exerted by
slope and aspect individually. However, our
results controvert these expectations and instead
comply with studies showing that simple topo-
graphic parameters reflect local thermal climate
better than potential solar radiation in humid
temperate regions due to the effects of cloud
cover in obscuring solar radiation estimates
(Reger et al. 2011).
We observed only weak associations between
aspect-determined heat balance and vegetation
patterns indicating that the heat index exerted
very little influence relative to other topographic
variables. Additionally, the index exhibited neg-
ative correlation with the average preferences for
temperature and light contrary to expectations.
Aspect-determined heat therefore did not appear
to be substantial in determining local vegetation–
topography relationships within the study region
even though it does appear to influence these
relations quite a bit in other studies (Boyko 1947).
The complexity of interactions between topog-
raphy and wind has prevented successful mod-
eling and rigorous study of the relationships
between topographically controlled wind re-
gimes and local vegetation patterns (but see
Knapp 1985, Chen et al. 1997, Ennos 1997, Lortie
and Cushman 2007, Li et al. 2009). In the current
study, wind exposure exhibited only weak
correlation to these patterns. Nonetheless, the
correlations between wind exposure and average
soil moisture preference (EIVmoist) were among
the strongest observed for forests as well as xeric
and mesic calcareous grasslands in this study.
Elevated levels of evapotranspiration and soil
evaporation caused by intensified wind exposure
(Knapp 1985) provide a reasonable explanation
for these observations. Under such circumstanc-
es, we would expect a negative correlation
between wind exposure and EIVmoist. The afore-
mentioned habitats affirmed this expectation in
showing strong negative correlations between
wind and the EIVmoist variable (in contrast to its
12 July 2013 v Volume 4(7) v Article 91

positive and weak relationship observed in other
habitats). Wind-driven desiccation is recognized
as an important factor in determining local plant
species distributions in forests (Ellenberg 1988).
Importance of topography
in different environments
The results given here indicate that the degree
to which topography influences vegetation de-
pends on site-level environmental parameters
including habitat type and moisture conditions
(as indicated by the site-mean TWI). Notably, the
strongest relationships between topographic fac-
tors and vegetation were observed in wet
habitats, and particularly so in tidal salt mead-
ows, while the weakest relationships occurred in
dry, sandy habitats and forest. The strength of
species-environment relationships depends on
the length of the environmental gradients under
investigation (Austin 1985). As the hydrologic
gradients in wetlands is longer than in other
habitats (Adam 1990), this explains the stronger
relationships there, i.e., when considering the
evidence that topography’s control of soil mois-
ture is the primary mechanism underlying the
local vegetation–topography relationships.
Sandy soil types have a relatively low water
retention capacity (Troeh and Thompson 2005),
reducing the influence of topography on soil
moisture in ecosystems that depend on infiltra-
tion from precipitation. This can explain the
relatively weak vegetation–topography relations
observed in dry, sandy habitats. We note how-
ever, that in groundwater-fed ecosystems, topo-
graphically controlled hydrology may play a
major role for vegetation patterns even on sandy
soils (Araya et al. 2011).
Albeit the vegetation–topography relation-
ships were weaker within dry sites generally,
TWI was still the most important functional
variable in determining vegetation patterns even
in the relatively dry dunes and heathlands.
Hence, our results suggest that topographically
controlled soil moisture is an important structur-
ing component of local vegetation patterns in
both wet and dry habitats.
CONCLUSIONS
New high-resolution LiDAR-based digital ele-
vation models and a massive vegetation plot
v www.esajournals.org
MOESLUND ET AL.
inventory allowed us to comprehensively assess
the general role played by topography in
structuring plant communities across habitats
and scales in a large lowland region. Topograph-
ically controlled factors such as temperature are
well-known determinants of vegetation patterns
in mountainous regions. Surprisingly, our results
showed that topography also consistently influ-
ences vegetation patterns at local scales in natural
and semi-natural lowland habitats. Topography’s
control of soil moisture patterns rather than
temperature emerged as the primary mechanism
underlying the observed lowland vegetation–
topography relationships, even within dry hab-
itat sites. Overall, our findings suggest that niche-
based species sorting along topographically
controlled gradients in soil moisture are among
the primary local plant community assembly
mechanisms across large lowland regions.
ACKNOWLEDGMENTS
We gratefully acknowledge funding from the Aar-
hus University Research Foundation via the Center for
Interdisciplinary Geospatial Informatics Research (CI-
GIR), the Danish Strategic Research Council, Center for
Massive Data Algorithmics, a Center of the Danish
National Research Foundation, and the Oticon Foun-
dation (grant to JEM).
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SUPPLEMENTAL MATERIAL
APPENDIX A

Table A1. An overview of the habitat types included in this study (a subset of Annex 1 of the EU Habitats
Directive; Council of the European Communities 1992).

Habitat number Habitat name Typical plant species

Coastal and halophytic habitats
1300  Atlantic and continental salt
marshes and salt meadows
1330 Atlantic salt meadows (Glauco-
Puccinellietalia maritimae)
1340 Inland salt meadows
Coastal sand dunes and inland dunes
2100  Sea dunes of the Atlantic, North
Sea and Baltic coasts
2130 Fixed coastal dunes with
herbaceous vegetation (‘‘grey
dunes’’)
2140 Decalcified fixed dunes with
Empetrum nigrum
2190 Humid dune slacks
Juncus gerardi ssp. gerardi, Agrostis stolonifera, Festuca rubra, Puccinellia
maritima, Seriphidium maritimum ssp. maritimum, Elytrigia repens ssp.
repens, Elytrigia atherica, Armeria maritima s.l., Spergularia salina,
Blysmus rufus, Puccinellia capillaris, Atriplex prostrata, A. portulacoides,
A. pedunculata, A. littoralis, Tripolium vulgare, Beta vulgaris spp.
maritima, Suaeda maritima, Tripleurospermum maritimum ssp.
maritimum, Glaux maritima, Triglochin maritima, Plantago maritima
spp. maritima, Eleocharis sp., Carex extensa, Puccinellia distans
Same as 1330
Aira praecox, Bromus hordeaceus ssp. hordeaceus, Carex arenaria,
Cerastium sp., Corynephorus canescens, Erodium cicutarium, Galium
verum ssp. verum, Gentianella campestris s.l., Koeleria glauca, Myosotis
ramosissima, Ononis spinosa ssp. maritina, Phleum arenarium, Polygala
vulgaris, Silene otites, Viola tricolor spp. curtisii, Hieracium sekt.
hieracioides, Hypochoeris radicata, Viola canina, Jasione montana, Festuca
ovina, F. rubra, Thymus serpyllum ssp. serpyllum, Sedum acre,
Geranium sanguineum, Geranium molle, Lotus corniculatus,
Anthoxantum odoratum, Artemisia campestris ssp. campestris, Pimpinella
saxifrage ssp. saxifrage, Poa pratensis ssp. pratensis, Armeria maritima
ssp. maritima
Carex arenaria, Empetrum nigrum, Genista tinctoria, Pyrola rotundifolia
ssp. maritima, Calluna vulgaris, Erica tetralix, Polypodium vulgare,
Lotus corniculatus, Deschampsia flexuosa, Ammophila arenaria,
Hieracium sekt. hieracioides
Highly variable, but some of the following species are usually present:
Hippuris vulgaris, Potamogeton sp., Phragmites australis, Schoenoplectus
maritimus, Juncus buffonius, Centaurium sp., Baldellia ranunculoides,
Hydrocotyle vulgaris, Salix repens ssp. argentea, Salix repens ssp.
rosmarinifolia

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Table A1. Continued.
Habitat number Habitat name
2250 Coastal dunes with Juniperus spp. Juniperus communis ssp. communis
2300  Inland dunes, old and decalcified
2320 Dry sand heaths with Calluna
and Empetrum nigrum
Temperate heath and scrub
4000 
4010 Northern Atlantic wet heaths
with Erica tetralix
4030 European dry heaths
Sclerophyllous scrub (matorral)
5130 Juniperus communis formations on
heaths or calcareous grasslands
Natural and semi-natural grassland formations
6100 
6120 Xeric sand calcareous grasslands
6200  Semi-natural dry grasslands and
scrubland facies
6210 Semi-natural dry grasslands and
scrubland facies on calcareous
substrates (Festuco-Brometalia)
(* important orchid sites)
6230 Species-rich Nardus grasslands,
on silicious substrates in
mountain areas (and
submountain areas in
Continental Europe)
6400  Semi-natural tall-herb humid
meadows
6410 Molinia meadows on calcareous,
peaty or clayey-silt-laden soils
(Molinion caeruleae)
6430 Hydrophilous tall herb fringe
communities of plains and of
the montane to alpine levels
Raised bogs, mires and fens
7000 
7100  Sphagnum acid bogs
7110 Active raised bogs
7120 Degraded raised bogs still
capable of natural regeneration
v www.esajournals.org
MOESLUND ET AL.
Typical plant species
Calluna vulgaris, Empetrum nigrum
Erica tetralix, Andromeda polifolia, Vaccinium uliginosum ssp. uliginosum,
V. oxycoccus, Calluna vulgaris
Calluna vulgaris, Empetrum nigrum, Vaccinium vitis-idaea ssp. vitis idaea,
V. uva-ursi, Genista anglica, G. pilosa, G. germanica
Juniperus communis, Crataegus laevigata, Rosa sp., Prunus spinosa
Allium schoenoprasum var. schoenoprasum, Cardaminopsis arenosa, Carex
ligerica, Dianthus deltoides, Helichrysum arenarium, Herniaria glabra,
Koeleria glauca, Petrorhagia prolifera, Cerastium semidecandrum, Vicia
lathyroides, Phleum arenarium, Aira caryophyllea, Trifolium striatum,
Erophila verna, Myosotis stricta, Silene conica, S. otites, Festuca polesica
Anthyllis vulneraria s.l., Arabis hirsuta var. hirsuta, Brachypodium
pinnatum, Campanula glomerata ssp. glomerata, Carex caryophyllea,
Carlina vulgaris ssp. vulgaris, Centaurea scabiosa, Koeleria pyramidata,
Leontodon hispidus, Medicago sativa ssp. falcata, Orchis mascula, O.
purpurea, Anacamptis morio, Neotinea ustulata, Primula veris,
Sanguisorba minor ssp. minor, Scabiosa columbaria, Bromopsis erecta,
Dactylis glomerata ssp. glomerata, Carex flacca, Prunella vulgaris,
Leucanthemum vulgare, Cirsium acaule, Plantago media, Linum
catharticum, Juniperus communis
Antennaria dioica, Arnica montana, Carex ericetorum, C. pallescens, C.
panicea, C. pilulifera, Festuca ovina, F. rubra, Galium saxatile, Gentiana
pneumonanthe, Hypericum perforatum, Hypochoeris maculata, Lathyrus
linifolius, Pseudorchis albida, Nardus stricta, Pedicularis sylvatica ssp.
sylvatica, Platanthera bifolia ssp. bifolia, Polygala vulgaris, Potentilla
erecta, Veronica officinalis, Viola canina, Deschampsia flexuosa, Danthonia
decumbens, Anthoxantum odoratum, Poa pratensis ssp. angustifolia,
Calamagrostis epigejos
Molinea caerulea, Carex flacca, C. nigra, C. pallescens, Inula salicina,
Dianthus superbus, Juncus subnodulosus, Primula farinosa, Selinum
carvifolia, Serratula tinctoria, Tetragonolobus maritimus, Viola palustris,
V. persicifolia, Galium uliginosum, Juncus conglomerates, J. acutiflorus,
Luzula multiflora ssp. multiflora, Ophioglossum vulgatum, Crepis
paludosa, Potentilla erecta, P. anglica, Lotus pedunculatus var.
pedunculatus, Dianthus deltoides
Glechoma hederacea, Epilobium hirsutum, Filipendula ulmaria, Angelica
archangelica ssp. litoralis, Petasites hybridus, Cirsium oleraceum,
Aegopodium podagraria, Alliaria petiolata, Geranium robertianum var.
robertianum, G. sylvaticum, Silene dioica, Lamium album, Lysimachia
punctata, Lythrum salicaria, Crepis paludosa, Trollius europaeus,
Calamagrostis arundinacea, Cirsium helenioides
Andromeda polifolia, Carex pauciflora, C. nigra, C. limosa, Drosera
rotundifolia, D. anglica, D. intermedia, Eriophorum vaginatum, E. gracile,
Vaccinium oxycoccus, Calluna vulgaris, Empetrum nigrum, Rhynchospora
alba, R. fusca, Scheuchzeria palustris, Utricularia intermedia, U. minor,
U. ochroleuca
Same as 7110
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 MOESLUND ET AL.

Table A1. Continued.

Habitat number Habitat name Typical plant species

7140 Transition mires and quaking Rhynchospora alba, R. fusca, Carex rostrata, C. lasiocarpa, C. limosa, C.
bogs chordorrhiza, C. diandra, Scheuchzeria palustris, Comarum palustre,
Menyanthes trifoliata, Epilobium palustre, Eriophorum gracile,
Hammarbya paludosa, Liparis loeselii, Pedicularis palustris ssp. palustris
7150 Depressions on peat substrates of Rhynchospora alba, R. fusca, Drosera intermedia, D. rotundifolia,
the Rhynchosporion Lycopodiella inundata
7200  Calcareous fens
7210 Calcareous fens with Cladium Cladium mariscus
mariscus and species of the
Caricion davallianae
7220 Petrifying springs with tufa Pinguicula vulgaris, Carex appropinquata, C. lepidocarpa ssp. lepidocarpa,
formation (Cratoneurion) Equisetum telmateia, Cardamine amara, Berula erecta, Chrysosplenium
sp., Hypericum tetrapterum
7230 Alkaline fens Schoenus nigricans, S. ferrugineus, Carex sp., Eriophorum latifolium,
Juncus subnodulosus, Dactylorhiza incarnata var. incarnata, D.
purpurella ssp. purpurella, Liparis loeselii, Herminium monorchis,
Epipactis palustris, Pinguicula vulgaris, Primula farinosa, Blysmus
compressus, Eleocharis quinqueflora, Parnassia palustris, Peucedanum
palustre, Eupatorium cannabinum, Calamagrostis canescens, Phragmites
australis
Forests
9100  Forests of Temperate Europe
  Main group number covering all subgroups beginning with the first two digits of this number (e.g., 1300 covers both 1330
and 1340).

Table A2. An overview of how the NATURA 2000

habitat types were grouped for this study.

Habitat type NATURA 2000 habitat codes

Salt meadows 1300 þ 1330 þ 1340
Xeric calcareous grasslands 6120
Humid dune slacks 2190
Acidic bogs 7100 þ 7110 þ 7120 þ 7140
Grasslands (unspecified) 6200
Molinia meadows 6400 þ 6410
Calcareous fens 7200 þ 7210 þ 7220 þ 7230
Wet heaths 4010
Calcareous grasslands 6210 þ 5130
Pioneer communities 7150
Forests 9xxx
Acidic grasslands 6230
Calcareous dunes 2130
Xeric heaths 4030
Relatively dry acidic dunes 2140 þ 2250 þ 2320
Notes: See Table A1 for further details about each NATURA
2000 habitat type. An ‘x’ denotes a single character wildcard.

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 MOESLUND ET AL.

Table A3. An overview of plants species that were assigned at least one Ellenberg indicator value (Ellenberg et al.
2001) by the authors of this study (bold numbers).

Species L T K F R N S Ecological information derived from

Abies grandis 3 5 4 5 6 4 0 Mossberg and Stenberg (2005) þ conifers.org
Abies nordmanniana 3 5 4 5 6 4 0 Mossberg and Stenberg (2005) þ conifers.org
Abies procera 3 5 4 5 6 4 0 Mossberg and Stenberg (2005) þ conifers.org
Alchemilla baltica 7 4 5 5 7 6 0 Mossberg and Stenberg (2005)
Alchemilla filicaulis var. vestita 6 6 3 5 7 4 0 Mossberg and Stenberg (2005)
Ammophila arenaria 3 8 6 3 4 7 2 1 Mossberg and Stenberg (2005)
Calamagrostis epigeios
Arabis hirsuta var. glaberrima 7 5 3 4 9 3 0 Mossberg and Stenberg (2005)
Arenaria serpyllifolia ssp. lloydii 7 5 3 3 7 4 2 Mossberg and Stenberg (2005)
Aronia arbutifolia 6 5 4 7 7 6 0 plant-encyclopedia.net
Artemisia maritima ssp. maritima 9 6 2 5 8 7 5 Mossberg and Stenberg (2005)
Aster tripolium 9 6 2 8 7 6 5 Mossberg and Stenberg (2005)
Atriplex littoralis 9 6 2 6 7 6 4 Mossberg and Stenberg (2005)
Atriplex calotheca 9 7 2 5 7 7 4 Christiansen and Anthon (1970)
Atriplex longipes ssp. longipes 9 6 2 6 7 8 5 Mossberg and Stenberg (2005)
Atriplex pedunculata 9 6 1 8 7 8 7 Mossberg and Stenberg (2005)
Atriplex portulacoides 9 6 1 7 7 7 8 Mossberg and Stenberg (2005)
Avena sterilis L. 7 6 6 4 7 7 0 Schou et al. (2009)
Betula pendula 3 pubescens 7 5 5 6 3 3 0 Mossberg and Stenberg (2005)
Blysmus rufus 8 6 2 8 7 4 5 Mossberg and Stenberg (2005)
Bromus xpseudothominii 8 6 2 4 7 4 0 Schou et al. (2009)
Cakile maritima ssp. baltica 9 6 2 6 7 8 3 Mossberg and Stenberg (2005)
Cakile maritima ssp. integrifolia 9 6 2 6 7 8 3 Mossberg and Stenberg (2005)
Carex divulsa ssp. leersii 5 5 5 5 4 6 0 Schou (2006)
Carex nigra var. recta 7 4 3 8 4 2 0 Schou (2006)
Carex ovalis 7 5 3 7 3 3 0 Schou (2006)
Carex salina 7 2 2 9 7 3 3 Mossberg and Stenberg (2005)
Carex trinervis 9 6 1 9 3 2 1 Schou (2006)
Chenopodium desiccatum var. 8 6 5 3 8 3 0 Crawford (1975)
leptophylloides (Murr) Wahl
Cotoneaster acutifolius var. lucidus 7 4 5 5 7 5 1 hort.uconn.edu/Plants/ 
(Schltdl.) L.T. Lu
Crambe maritima 9 6 2 5 8 7 3 Mossberg and Stenberg (2005)
Crataegus xmedia 5 5 3 5 7 5 0 Mossberg and Stenberg (2005)
Dactylorhiza incarnata var. lobelii 8 6 2 6 9 2 0 Pedersen and Faurholdt (2010)
Dactylorhiza maculata subsp. 8 5 4 3 3 2 0 thewesternisles.co.ukà
ericetorum (E.F.Linton) P.F.Hunt
& Summerh.
Dactylorhiza praetermissa 8 6 3 7 5 4 0 Pedersen and Faurholdt (2010)
Dactylorhiza purpurella ssp. 8 4 1 7 9 2 0 Pedersen and Faurholdt (2010)
purpurella
Dactylorhiza purpurella var. 8 4 1 7 9 2 0 Pedersen and Faurholdt (2010)
cambrensis (R.H.Roberts)
R.M.Bateman & Denholm
Deschampsia flexuosa ssp. flexuosa 6 4 2 5 2 3 0 Schou et al. (2009)
Eleocharis uniglumis ssp. uniglumis 7 5 2 10 7 5 5 Schou (2006)
Elytrigia atherica 3 E. juncea 9 6 2 4 7 3 2 Schou et al. (2009)
Elytrigia juncea 3 Leymus 9 6 2 5 7 6 3 Schou et al. (2009)
arenarius
Elytrigia juncea 3 E. repens 8 6 3 5 7 7 3 Schou et al. (2009)
Elytrigia repens ssp. repens 7 6 7 4 7 7 0 Schou et al. (2009)
Elytrigia repens 3 Hordeum 7 6 2 5 6 6 2 Schou et al. (2009)
secalinum
Epilobium glandulosum 7 4 3 5 7 7 0 Mossberg and Stenberg (2005)
Epipactis helleborine var. 7 6 2 4 8 2 0 Pedersen and Faurholdt (2010)
neerlandica
Euphrasia arctica ssp. minor 8 4 1 6 7 3 2 Mossberg and Stenberg (2005)
Euphrasia dunensis 8 5 1 4 9 2 0 Mossberg and Stenberg (2005)
Euphrasia micrantha 6 4 2 3 2 1 0 Mossberg and Stenberg (2005)
Fargesia nitida (Mitford) Keng f. 5 7 9 5 7 8 0 www.fargesiaheaven.com
ex T.P.Yi
Festuca arundinacea 8 5 6 6 7 6 1 Schou et al. (2009)
Festuca gigantea 3 Lolium perenne 6 5 3 5 6 6 0 Schou et al. (2009)
Festuca pratensis 3 Lolium perenne 7 5 3 6 6 6 0 Schou et al. (2009)
Festuca rubra ssp. commutata 8 6 5 3 6 4 9 Schou et al. (2009)
Galeopsis speciosa 7 5 6 5 7 7 0 Mossberg and Stenberg (2005)

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 MOESLUND ET AL.

Table A3. Continued.

Species L T K F R N S Ecological information derived from

Galium mollugo 3 G. verum 7 6 3 4 6 3 0 Mossberg and Stenberg (2005)
Gentianella uliginosa 8 6 3 8 7 2 0 Mossberg and Stenberg (2005)
Geranium robertianum var. 8 6 1 4 7 3 1 Mossberg and Stenberg (2005)
rubricaule
Glaux maritima 8 6 1 7 7 5 4 Mossberg and Stenberg (2005)
Glyceria maxima 7 5 4 10 7 8 0 Mossberg and Stenberg (2005)
Hieracium sect. Hieracium 5 6 4 5 7 6 0 Schou (2001)
Hieracium sect. Sabauda 9 7 7 4 4 3 0 Schou (2001)
Hieracium sect. Tridentata 8 7 4 4 3 2 0 Schou (2001)
Hieracium sect. Vulgata 7 6 5 5 5 6 0 Schou (2001)
Honckenya peploides ssp. peploides 9 6 1 6 7 7 5 Mossberg and Stenberg (2005)
Juncus alpinoarticulatus ssp. 8 3 4 6 4 2 0 Mossberg and Stenberg (2005)
nodulosus
Juncus gerardii ssp. gerardii 8 6 2 7 7 4 7 Mossberg and Stenberg (2005)
Juncus ranarius 9 6 2 8 4 3 4 Mossberg and Stenberg (2005)
Larix kaempferi 7 7 4 6 5 3 0 Mossberg and Stenberg (2005)
Larix xmarschlinsii 7 6 5 5 6 3 0 Mossberg and Stenberg (2005)
Lathyrus japonicus ssp. acutifolius 8 6 4 4 7 3 1 Mossberg and Stenberg (2005)
Leymus arenarius 9 6 2 5 7 6 3 Schou et al. (2009)
Ligusticum scoticum 8 6 1 6 7 5 3 Mossberg and Stenberg (2005)
Limonium humile 9 6 1 8 7 5 6 Mossberg and Stenberg (2005)
Lithospermum arvense var. arvense 8 6 5 5 8 3 0 Mossberg and Stenberg (2005)
Lotus pedunculatus var. villosus 8 6 1 6 8 2 9 Mossberg and Stenberg (2005)
Malus toringo 7 6 4 5 6 6 0 Mossberg and Stenberg (2005)
Tripleurospermum perforatum 7 6 3 5 6 6 0 Mossberg and Stenberg (2005)
Odontites littoralis ssp. littoralis 8 6 3 7 7 4 4 Mossberg and Stenberg (2005)
Ononis spinosa ssp. maritima 7 6 4 4 7 3 0 Mossberg and Stenberg (2005)
Picea glauca 5 5 5 6 6 6 0 Mossberg and Stenberg (2005) þ conifers.org
Picea omorika (Pancic) Purk. 5 5 6 5 - 4 0 wikipedia.org
Picea pungens Engelm. 5 4 6 6 7 3 0 wikipedia.org
Picea sitchensis 7 5 3 7 2 2 0 Mossberg and Stenberg (2005) þ conifers.org
Pinus contorta 7 4 4 5 5 2 0 Mossberg and Stenberg (2005) þ conifers.org
Plantago major ssp. winteri 8 6 3 5 5 4 2 Mossberg and Stenberg (2005)
Plantago maritima ssp. maritima 8 6 2 7 6 4 3 Mossberg and Stenberg (2005)
Polygonum aviculare ssp. 7 6 4 4 8 5 1 Mossberg and Stenberg (2005)
rurivagum
Populus balsamifera 6 5 3 5 7 7 0 borealforest.org§
Populus xcanadensis 5 7 5 8 7 7 0 Mossberg and Stenberg (2005)
Potentilla tabernaemontani 8 5 2 3 8 2 0 Mossberg and Stenberg (2005)
Potentilla xsubarenaria 9 6 3 3 7 2 0 Mossberg and Stenberg (2005)
Puccinellia capillaris 9 6 1 8 7 4 7 Mossberg and Stenberg (2005)
Pyrola rotundifolia ssp. maritima 5 5 1 7 9 2 0 Mossberg and Stenberg (2005)
Ranunculus peltatus ssp. baudotii 8 6 2 10 9 7 6 Mossberg and Stenberg (2005)
Rhinanthus serotinus ssp. vernalis 7 5 3 5 7 2 0 Mossberg and Stenberg (2005)
Rumex acetosa ssp. acetosa var. 8 6 5 9 5 6 0 Mossberg and Stenberg (2005)
hydrophilus
Sagina maritima 9 6 3 7 7 4 4 Mossberg and Stenberg (2005)
Salicornia europaea 9 5 2 8 8 6 9 Mossberg and Stenberg (2005)
Salix aurita 3 S. repens 7 5 3 7 3 3 0 Mossberg and Stenberg (2005)
Salix xmeyeriana 7 5 2 8 7 5 0 Mossberg and Stenberg (2005)
Salix xrubens 7 6 5 5 6 6 1 Mossberg and Stenberg (2005)
Salsola kali 9 7 3 4 7 8 6 Mossberg and Stenberg (2005)
Satureja acinos 9 7 6 3 8 6 0 Christiansen and Anthon (1970)
Schistophyllidium bifurcum 8 6 8 5 7 6 0 Mossberg and Stenberg (2005)
Scirpus maritimus 8 6 3 10 8 7 2 Mossberg and Stenberg (2005)
Silene latifolia ssp. alba 8 6 4 4 7 7 0 Christiansen and Anthon (1970)
Silene uniflora 8 6 2 6 6 4 3 Christiansen and Anthon (1970)
Sonchus arvensis var. maritimus 8 5 3 6 5 4 2 Mossberg and Stenberg (2005)
Spartina xtownsendii 9 5 2 9 8 6 7 Schou et al. (2009)
Spergula morisonii 9 5 4 3 5 2 0 Mossberg and Stenberg (2005)
Stachys palustris 3 S. sylvatica 6 5 4 7 7 7 0 Mossberg and Stenberg (2005)
Stellaria crassifolia 9 5 7 9 8 3 0 Mossberg and Stenberg (2005)
Suaeda maritima 9 6 2 8 8 6 7 Mossberg and Stenberg (2005)
Tilia xeuropaea 5 7 5 5 6 4 0 Mossberg and Stenberg (2005)
Trifolium ornithopodioides (L.) Sm. 9 6 3 6 5 3 0 nlbif.eti.uva.nl/bis/flora.php}
Triglochin maritima 8 6 3 7 7 5 8 Christiansen and Anthon (1970)
Typha angustifolia 3 T. latifolia 8 6 5 10 7 7 0 Mossberg and Stenberg (2005)

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 MOESLUND ET AL.

Table A3. Continued.

Species L T K F R N S Ecological information derived from

Viola uliginosa 3 5 4 7 4 7 0 Mossberg and Stenberg (2005)
Notes: The values assigned by the authors of this paper are bolded. Plain numbers represent values taken from the indicator-
value references in the priority mentioned in the main text. Nomenclature follows Mossberg and Stenberg (2005) unless the
author is given. L, Ellenberg indicator value (EIV) for light; T, for temperature; K, for continentality; F, for soil moisture; R, for
soil reaction; N, for soil nitrogen content; S, for soil salinity.
  University of Connecticut plant database.
à Western Isles Wildflowers.
§ Faculty of Natural, Resources Management, Lakehead University.
} Interactive Flora of NW Europe.

Table A4. A comparison (Pearson correlation coeffi-

cients, r) of mean plot indicator values with and
without author assigned values (Table A3).

Indicator value r
EIVlight 0.999
EIVtemp 0.987
EIVcont 0.767
EIVmoist 0.998
EIVreac .0.999
EIVnitr 0.998
EIVsalt .0.999
Notes: The table is based on the plot mean indicator values
for the first 100 sites in the dataset embracing 3878 plots
equally covering all habitat types included in the study.

APPENDIX B
Detailed descriptions of the calculation of the wind
exposure and the topographic wetness index
Wind exposure.—The ‘‘Hillshade’’ tool in the Spa-
tial Analyst module in ArcGIS 10 creates a
shaded relief from a DEM by considering the
azimuth (compass direction) and altitudinal
angles of an artificial illumination source. By
using an altitude angle of 58 hillshade addition-
ally represents the potential wind exposure from
a given direction (Boose et al. 1994, Mikita and
Klimánek 2010). Using this tool and wind speed
and direction data for Denmark we created a
map of potential wind exposure for all Denmark.
Initially, one hillshade raster was created for each
of the following compass directions: 08, 308, 608,
908, 1208, 1508, 1808, 2108, 2408, 2708, 3008 and
3308. In Denmark wind is not equally distributed
among the compass directions (Cappelen and
Jørgensen 1999). To account for this we incorpo-
rated data on general wind speed and direction:
for each compass direction a map of average
wind speed and a map of frequency for that
particular wind direction were created by inter-
polating (Inverse Distance Weighting, IDW) data
from 62 weather stations distributed throughout
Denmark (Cappelen and Jørgensen 1999). The
IDW calculations were performed using the
‘‘IDW’’ tool in the ‘‘Geostatistical Analyst’’
module in ArcGIS 10 with default settings
(default power ¼ 2) except that we used an
output cell size of 250 m since the general wind
speed and direction is unlikely to change notably
over such short distances. These two maps were
used for weighting (equal weights) the potential
impact by wind as represented by the corre-
sponding hillshade layers, allowing us to com-
pute a map of weighted-average potential wind
exposure for all of Denmark.
Topographic wetness index.—The TWI of a DEM
Cell (i ) is given by ln(As/tan(h i )), where As is the
specific catchment area associated with i (de-
tailed below) and hi is the slope angle of i
(Wilson and Gallant 2000). As is defined as Au/Li,
where Au is the area of the upslope cells draining
into i and Li is the dimension of i measured as
the width of the cell in the direction orthogonal

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to the water flow direction. The calculations of
Au and Li are typically based on outputs from
flow accumulation and flow direction computer
algorithms. Here, we used the output of a single-
flow direction algorithm (D8) (Wilson and
Gallant 2000) for the subsequent run of a flow
accumulation algorithm. The SCALGO Hydrol-
Fig. B1. Flowchart of the procedure used for calculation of the topographic wetness index (TWI).
v www.esajournals.org
MOESLUND ET AL.
ogy software package version 1.0.5 (SCALGO,
Aarhus, Denmark) was used for these opera-
tions. Based on these the TWI was computed in
the ArcGIS 10 environment using the above
formula. For a flowchart showing all TWI
calculation details, see Fig. B1.
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 MOESLUND ET AL.

(continuation of Fig. B1 legend)

Below, the steps requiring further explanation are detailed. Steps initiated by the word SCALGO was
performed using the SCALGO Hydrology software package version 1.0.5 (SCALGO, Aarhus, Denmark). The
remaining steps were performed using ArcGIS 10 (Environmental Systems Research Institute, Redlands,
California, USA). Blue boxes are input data, yellow ovals represent calculation procedures, green boxes are
output data sets, and transparent boxes are output data sets not used here.
SCALGO SFD flow directions: This module models a water flow regime where the water is allowed to flow in
only one direction, namely to the lowest lying downslope neighbor cell relative to the initial cell (O’Callaghan
and Mark 1984).
SCALGO flow accumulation: This module constructs a flow accumulation raster where each cell is assigned a
value proportional to the amount of water that potentially reaches it from all cells draining into it according to the
given flow direction method (cf. above).
3 3 3 focal minimum: For each cell the lowest value of the 8 neighboring cells is allocated. This is taken to be the
elevation value of the cell to which the flow is directed (as calculated by SFD-flow). The focal statistics tool in
ArcGIS was used for this operation.
Calculate DZ: The elevation difference between each raster cell and the cell to which it flows (under an SFD
regime) is computed by subtracting the output of ‘‘3 3 3 focal minimum’’ from the DEM.
Calculate cell dimension (L): For the cardinal directions the value is equal to the length of the cell edge; for the
diagonal directions the value is equal to the diagonal length of the cell.
Calculate catchment size (number of cells): This is equal to the flow accumulation value plus 1 (the cell itself ).
Calculate catchment area (Au): The catchment area is equal to catchment size multiplied by the cell area.
Calculate Tan(h): Tan(h) is equal to ‘‘rise’’/’’run’’, i.e., DZ/L. h is the slope between the cell in question and the
lowest lying downslope neighbor cell.
Calculate specific catchment area (As): The As is equal to the Au divided by tan(h ).
Shift 0 with 0.000001: since tan(h ) is used as denominator in a division (and it may take the value zero), all zero
entrances in the raster is assigned the value 0.000001.
Calculate TWI: calculates ln(As/ tan(h i )), the topographic wetness index.

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 MOESLUND ET AL.

APPENDIX C
Implementation examples and detailed results

Fig. C1. Vegetation–topography relationships at two sites (both alkaline fens). Panel (A) illustrates one of the
strongest (r ¼0.68) correlations between the topographical wetness index (TWI) and the average preference for
moisture (EIVmoist). Site location: 55.978 N, 12.358 E. TWI is superimposed upon the elevation model for the area.
White means high TWI (22.5) and black low (2.3). Elevation ranges 9.2–21.4 meters above sea-level. Panel (B)
shows the same as (A), but for a strong elevation-DCA first axis correlation (r ¼ 0.94). Site location: 55.478 N, 9.798
E. Here, the white-black gradient refers to terrain elevation, 0.3–7.9 meters above sea-level.

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 MOESLUND ET AL.

Fig. C2. Coefficients of determination (R 2adj) for each vegetation measure from the regional-scale models
combining plot data (including habitat type) from all 901 study sites across Denmark. Abbreviations next to each
bar identify statistically significant (P , 0.05) explanatory variables. Hab, Habitat type; Ele, Elevation; Heat, Heat
Index; Slo, Slope; Rad, Potential Solar Radiation; TWI, Topographic Wetness Index; Wind, Wind Index. See Table
1 for abbreviations.

Fig. C3. Mean coefficients of determination (R 2adj) for each vegetation measure from the within-site (local)
models with elevation excluded as explanatory variable. See Table 1 for abbreviations.

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 MOESLUND ET AL.

Table C1. Average squared Spearman’s correlation coefficients (r 2) for topography and vegetation within each of
the 901 natural and semi-natural sites distributed throughout Denmark.

Vegetation Potential Topographic

measure Elevation Heat index Slope solar radiation wetness index Wind index
EIVcont 0.098(22)(3)A,DE 0.046(þ4)(1)D,C 0.061(17)(2)B,D 0.062(4)(2)BC,C 0.061(þ8)(2)B,D 0.051(4)(2)C,CD
EIVlight 0.101(56)(3)A,E 0.050(24)(2)D,C 0.071(56)(2)B,BCD 0.058(þ17)(2)C,C 0.073(þ70)(2)B,C 0.057(þ33)(2)C,CD
EIVmoist 0.147(200)(4)A,B 0.056(113)(2)E,A 0.096(2181)(3)C,A 0.073(244)(3)D,A 0.109(1326)(4)B,A 0.065(238)(2)D,AB
(3)A,E
EIVnitr 0.098(50) 0.046(4) (1)D,BC
0.076(0)(3)B,BC 0.063(8)(2)C,BC 0.069(þ17)(2)B,C 0.056(22)(2)C,CD
EIVreac 0.122(103)(4)A,C 0.048(27)(1)E,ABC 0.075(17)(3)B,B 0.057(þ8)(2)DE,C 0.067(þ38)(2)C,C 0.056(þ13)(2)D,C
EIVsalt 0.116(51)(3)A,E 0.047(19)(1)D,C 0.070(2)(2)B,CD 0.056(0)(2)C,C 0.076(þ11)(2)B,C 0.053(þ19)(2)C,CD
EIVtemp 0.084(13)(3)A,F 0.046(17)(1)E,BC 0.064(þ17)(2)B,D 0.057(þ8)(2)CD,C 0.060(17)(2)BC,D 0.050(17)(2)DE,D
DCA 0.166(225)(4)A,A 0.054(14)(2)D,A 0.098(14)(3)B,A 0.072(22)(2)C,A 0.101(16)(3)B,B 0.072(220)(2)C,A
(4)A,AB (2)D,A (3)B,A (2)C,AB
PCA 0.147(213) 0.053(117) 0.091(113) 0.068(17) 0.091(24)(3)B,B 0.064(213)(2)C,AB
(4)A,CD (2)D,AB (3)B,A (2)C,AB
SpRich 0.118(þ4) 0.053(4) 0.092(1113) 0.067(50) 0.068(50)(2)C,C 0.059(4)(2)D,BC
Notes: The balance between positive (þ) and negative () relationships is given as percentages in parentheses (e.g., 22 means
that negative relationships outnumbered positive relationships by 22%). Superscript parentheses: (1): .5%, (2): .10%, (3):
.20%, (4): .30% significant relationships (P , 0.05). Values were compared by row (superscripted letters before comma) and
by column (superscripted letters after comma) with different letters indicating significantly different relationships (Student’s t-
test, P , 0.05). Bold values indicate the best and second best topographic determinants of the vegetation measure in question
according to Student’s t-test results. Underlined values refer to the two vegetation measures that are the most strongly
controlled by the topographic variable in question (according to Student’s t-tests). EIV: plot-level average Ellenberg indicator
value for subscript variables. See Table 1 for abbreviations.

Table C2. Results from OLS multiple regression models linking the correlation strength of five selected correlation
pairs to 8 of the 18 site-level environmental variables for the 901 study sites located throughout Denmark.

Full Mean SD of Mean SD of SD of Mean SD of SD of

Response variable model R 2adj TWI EIVmoist EIVtemp EIVtemp EIVsalt wind EIVcont EIVlight
Strongest correlation pair 0.14 0.18 0.11 0.14 0.11 0.02 0.14 0.10 0.06
at each site
EIVmoist-TWI 0.10 0.06 0.31 0.10 0.11 0.12 0.13 0.03 0.04
DCA-elev 0.10 0.24 0.09 0.07 0.10 0.09 0.08 0.07 0.04
PCA-elev 0.09 0.14 0.07 0.07 0.10 0.08 0.02 0.07 0.07
SpRich-elev 0.06 0.24 0.07 0.15 0.02 0.07 0.02 0.06 0.02
Mean absolute regression 0.17 0.13 0.11 0.09 0.08 0.08 0.07 0.05
coefficient
Notes: The five response variables are the strength (r 2) of (1) the strongest correlation pair (between a vegetation and a
topography variable) within each site, and (2–5) four within-site Spearman correlation pairs selected to cover the most
important vegetation measures and topographic variables. For each response variable, the R 2adj is given for the full multiple
regression model along with standardized regression coefficients for each of the environmental variables. The mean absolute
standardized regression coefficient for the five response variables represents the general importance of each of the
environmental variables in determining the strength of the local (within-site) vegetation–topography relationships. Elev,
elevation; Heat, heat index; Rad, potential solar radiation; TWI, topographic wetness index; Wind, wind exposure. See Table 1
for the remaining abbreviations.

Table C3. Results from OLS multiple regression models linking the correlation strength of five selected correlation
pairs to the remaining 10 of the 18 site-level environmental variables for the 901 study sites located throughout
Denmark.

Mean Mean SD of SD of SD of Mean SD of Mean Mean Mean

Response variable EIVcont EIVmoist EIVreac Heat TWI EIVlight EIVnitro Rad Elev Heat
Strongest correlation pair 0.05 0.11 0.02 0.09 0.05 0.04 0.04 0.07 0.01 0.02
at each site
EIVmoist-TWI 0.04 0.04 0.12 0.06 0.12 0.03 0.08 0.03 0.00 0.07
DCA-elev 0.10 0.02 0.00 0.02 0.02 0.02 0.01 0.01 0.04 0.05
PCA-elev 0.02 0.03 0.07 0.04 0.03 0.06 0.01 0.03 0.07 0.00
SpRich-elev 0.04 0.03 0.06 0.03 0.05 0.05 0.02 0.02 0.02 0.00
Mean absolute regression 0.05 0.05 0.05 0.05 0.05 0.04 0.03 0.03 0.03 0.03
coefficient

Note: See Table C2 for details.

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