Biodivers Conserv (2012) 21:3487–3506

DOI 10.1007/s10531-012-0376-1

ORIGINAL PAPER

Inclusion of explicit measures of geodiversity improve

biodiversity models in a boreal landscape

Jan Hjort • Risto K. Heikkinen • Miska Luoto

Received: 19 April 2012 / Accepted: 20 September 2012 / Published online: 2 October 2012
Ó Springer Science+Business Media Dordrecht 2012

Abstract Measures of geodiversity may provide a potentially useful surrogate for bio-
diversity patterns in insufficiently surveyed areas. However, their reliability in modelling
the spatial variation in species richness is inadequately understood. We investigated
whether the explanatory and predictive power of species richness models can be improved
by considering explicit measures of geodiversity (variability of earth surface materials,
forms and processes) in addition to climate and topography variables. Vascular plant
species richness was modelled in two study areas in Northern Europe, Finland at the
resolution of 500 or 1000 m, and as a function of three geodiversity (geological, geo-
morphological and hydrological diversity) variables, and six climate and topography
variables. Variation partitioning was used to identify the independent and shared contri-
butions of the geodiversity, climate and topography variable groups in explaining the
spatial patterns of species richness. Generalized additive models were used to explore the
ability of the different explanatory variables in predicting plant species richness within and
between the study areas. In both the study areas, the inclusion of measures of geodiversity
improved the explanatory power, predictive ability and robustness of the plant species
richness models. In conclusion, the explicit measures of geodiversity appear to be prom-
ising surrogates of biodiversity, which reflect important abiotic resource factors, and may
thus provide an equally, or even more reliable basis for transferring biodiversity models to
new areas than models based on climate and topography variables.

Electronic supplementary material The online version of this article (doi:10.1007/s10531-012-0376-1)

contains supplementary material, which is available to authorized users.

J. Hjort (&)
Department of Geography, University of Oulu, P.O. Box 3000, 90014 Oulu, Finland
e-mail: jan.hjort@oulu.fi

R. K. Heikkinen
Natural Environment Centre, Finnish Environment Institute, P.O. Box 140, 00251 Helsinki, Finland

M. Luoto
Department of Geosciences and Geography, University of Helsinki, P.O. Box 64, 00014 Helsinki,
Finland

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Keywords Biodiversity  Boreal vegetation zone  Conservation  Generalized additive

model  Geodiversity  Plant species richness  Variation partitioning

Introduction

The preservation of biodiversity is of paramount importance under current global change

(Sala et al. 2000; Brooks et al. 2006; Barnosky et al. 2011). However, effective conservation
planning is often difficult because spatial data on biodiversity patterns are sparse in many
areas and expensive to acquire. Thus, a long-term challenge in conserving biologically
valuable areas is the development of an efficient, but yet at the same time comprehensive
approach for identification of biologically diverse landscapes, and thereby establishing robust
methods for targeting of conservation actions (Whittaker et al. 2005). The most straight-
forward approach to assessing conservation priorities would be the direct mapping of all
biological features of the target area. As this is rarely feasible, numerous indirect surrogates of
biodiversity, especially for species richness, have been developed (e.g. Ferrier 2002; Sarkar
et al. 2006; Grantham et al. 2010). Desirable attributes for robust and cost-effective surrogates
include rapidity and ease of data gathering, sufficient data coverage in the study area, and
ecologically plausible links between the species richness patterns and the given surrogate(s).
Biological diversity is determined by internal biotic factors, or extrinsic abiotic factors,
or both (Huston 1994). In recent years, the concept of geodiversity has been put forward as
a novel alternative and potentially useful means to assess and model spatial biodiversity
patterns (see Parks and Mulligan 2010, and the references therein). According to the
classical definition, geodiversity refers to the variability of geological, geomorphological
and soil features (Gray 2004). In contrast, in the broadest sense geodiversity includes
geology, geomorphology, topography, hydrology and climate (Benito-Calvo et al. 2009;
Parks and Mulligan 2010). These five components are intimately linked with key abiotic
drivers of biodiversity, such as energy, water and nutrients (Richerson and Lum 1980; Kerr
and Packer 1997). In other words, geodiversity captures features from the geosphere
(geology, geomorphology and topography), hydrosphere (e.g. streams and springs) and
atmosphere (e.g. temperature) and their spatial variation, and by these means displays the
heterogeneity of abiotic features of Earth surface. Thus, in cases in which biodiversity
patterns are mainly governed by abiotic factors, geodiversity may provide a useful sur-
rogate for various aspects of biodiversity (Whittaker et al. 2001; Willis and Whittaker
2002; Hjort and Luoto 2010; Parks and Mulligan 2010).
A better understanding of the relationship between geodiversity (in a broad sense) and
biodiversity has been one of the main aims of biogeographical research for many decades (e.g.
Harner and Harper 1976; Swanson et al. 1988; Nichols et al. 1998), but the topic is still
insufficiently understood. Firstly, this is because most of the previous studies have been
conducted on broad scales and covered only one or some aspects of geodiversity. The focal
point in such modelling studies has predominantly been on the impacts of factors such as
climate and topography (e.g. Richerson and Lum 1980; Owen 1989; Kerr and Packer 1997;
Kocher and Williams 2000; Ferrer-Castán and Vetaas 2005; Davidar et al. 2007). Thus, the
classical geodiversity attributes have largely been ignored and studies focussing on aspects of
hydrology or soils, for example, have clearly been in the minority (e.g. Burnett et al. 1998;
Lobo et al. 2001; Pausas et al. 2003; Titeux et al. 2009; Illán et al. 2010). Secondly, most of the
previous studies have used oversimplified (e.g. digital elevation model, DEM) measures of
geodiversity (Nichols et al. 1998; Mutke and Barthlott 2005; Barthlott et al. 2007; Randin

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et al. 2009). Thirdly, the relationship between species diversity and measures of geodiversity
has not been unequivocally described or explicitly tested (Parks and Mulligan 2010). As a
result of these shortcomings, there is a lack of studies in which the potential of geodiversity as
a surrogate of biodiversity has systematically been assessed using comprehensive species
richness data and unambiguous measures of geodiversity.
In this study, we specifically aimed at comparing the performance of explicitly measured
variables of geodiversity (i.e. measures of geological, geomorphological and hydrological
variability) and commonly used abiotic surrogates of biodiversity in modelling plant species
richness. This was done by developing species richness models for two study areas in the
boreal vegetation zone in Finland, Northern Europe, with comprehensive plant species dis-
tribution and environmental data. We investigated whether (i) the explanatory power and (ii)
predictive ability of the vascular plant species richness models was improved by including
measures of geodiversity as an explanatory variable to complement the set of variables
describing climatic and topographical conditions. The predictive ability of the models was
assessed with respect to within area (within-area prediction, alias model interpolation) and
between areas (between-area prediction, alias model extrapolation) model performance.

Study area

The study areas are located in Finland, Northern Europe (Fig. 1). The Kevo study area
(69°390 N, 26°460 E; 362 km2) lies north of the northern limit of continuous pine forests in
the transition zone between the northern boreal and hemiarctic zones (Ahti et al. 1968;
Oksanen and Virtanen 1995). The study area is within the Kevo Strict Nature Reserve. The
area is characterized mostly by open uplands with forests of subalpine mountain birch
Betula pubescens ssp. czerepanovii, shallow peats supporting mires, and gently sloping
hills and treeless fells (Heikkinen et al. 1998, Hjort and Luoto 2009). The Kevo area
includes most of the catchment area of the Kevojoki River which runs in a steep-sided
valley, the walls of which can be over 150 m. The climate is subarctic with a mean annual
air temperature (MAAT) of -1.7 °C (-14.8 °C in January, 13.0 °C in July) and mean
annual precipitation (MAP) of 414 mm (1971–2000; Drebs et al. 2002). The growing
season ([5 °C) in the area is 110–120 days.
The Oulanka study area (66°220 N, 29°190 E; *110 km2), part of Oulanka National Park,
is located in the northern boreal zone (Ahti et al. 1968; Fig. 1). The area is characterized by
forested hills, and a mosaic of river valleys, water bodies and open wetlands is typical
(Parviainen et al. 2009a). Despite the rather harsh climatic conditions, the vegetation is
relatively rich with Arctic, eastern, Siberian and southern species (Vasari et al. 1996), and
with Norway spruce (Picea abies), Scots pine (Pinus sylvestris) and birches (Betula spp.)
as the dominant tree species. The climate of the region is rather continental with a MAAT
of -0.7 °C (-15.0 °C in January, 14.7 °C in July) and MAP of 568 mm (1971–2000;
Drebs et al. 2002). The growing season in the area is ca. 120–130 days.

Materials and methods

Species richness data

A grid system at a mesoscale resolution was utilized as a spatial study design (Fig. 2). The
grid cell size was determined by the resolution of the plant species data. A grid cell size of

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Fig. 1 Locations of the study areas in Finland, Northern Europe: a Kevo (362 km2) and b Oulanka
(* 110 km2) (HB hemiboreal, SB southern boreal, MB middle boreal, NB northern boreal). The terrain
views were produced using the shaded relief surface model derived from digital elevation models (sun
angle = 45°, azimuth = 315°). The elevations ranged from 95 to 550 m above sea level (a.s.l.) in Kevo and
from 140 to 385 m a.s.l. in Oulanka

1,000 9 1,000 m was used in Kevo (n = 362). In Oulanka (n = 440) most of the grid
cells were 500 9 500 m, but the sizes of the marginal grid cells were often less than 25
hectares. The species data were obtained from Söyrinki and Saari (1980) and Heikkinen
and Kalliola (1990) for Oulanka and Kevo, respectively. Both of the two areas have been
intensively surveyed, as they have been subject to a number of academic theses conducted
by the Turku and Oulu university research stations, several hundred days of floristic
mapping activities, as well as vegetation mapping of the whole area. Moreover, a number
of successful earlier studies have been based on these two plant species data sets, including
Heikkinen and Neuvonen (1997), Heikkinen et al. (1998), and Parviainen et al. (2009b).
Due to such a high intensity of the surveys, we consider that our two study areas represent
some of the floristically best-known areas in their size category in the entire circumboreal
vegetation zone. A total of 313 vascular plant species were recorded in the Kevo area,
whereas 476 were recorded in Oulanka, respectively. In the subsequent analysis, the total
number of vascular plant species recorded in each grid cell was used as a measure of
biodiversity (Araújo et al. 2004; Fig. 2).

Explanatory variables

In this study, we compiled a total of 17 climate and topography-based variables to cover the
most commonly used abiotic resource factors affecting the vascular plant species richness at
the mesoscale resolution (Table 1; e.g. Whittaker et al. 2001; Willis and Whittaker 2002;
Field et al. 2009). In addition, we compiled three explicit (‘direct’) measures of geodiversity
(i.e. diversity of geology, geomorphology and hydrology) to be used in the comparative
modelling setting. In this study, we specifically focused (based on the stricter definition of
geodiversity) on directly measured geodiversity variables which are hereafter referred to as
measures of geodiversity, and the explanatory power of climate and topography variables

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Fig. 2 Vascular plant species richness and measures of geodiversity in Kevo (a–d) and Oulanka (e–h).
Spatial patterns of species richness are shown in (a) and (e), geological diversity in (b) and (f),
geomorphological diversity in (c) and (g) and hydrological diversity in (d) and (h). The grid cell size is
1,000 9 1,000 m in Kevo and 500 9 500 m in Oulanka

were assessed and treated separately. A starting hypothesis for our study was that the geology,
geomorphology and hydrology variables have the potential to reflect the variability of abiotic
conditions more intimately than climate and topography variables. Especially, plant species
richness was expected to have rather weak relationship with climate variables in our meso-
scale analysis.
We compiled measures of geodiversity following Hjort and Luoto (2010) by simply
summing the total number of different geological, geomorphological and hydrological
features in the study grid cells (Table 1; Fig. 2). For example, the geomorphological
diversity is the sum of different geomorphological feature types regardless of the number
and cover of the specific features in the study grid cells. We mapped the features of the

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Table 1 List of potential explanatory variables compiled for variation partitioning and generalized additive
modelling
Explanatory variable Unit Kevo (NB–HA) Oulanka (NB)
Md (min to max) Md (min to max)

Geodiversity
Geological diversity – 5 (2 to 7) 4 (2 to 7)
Geomorphological diversity – 10 (2 to 23) 5 (1 to 16)
Hydrological diversity – 2 (0 to 4) 1 (0 to 4)
Climate
Mean annual air temperature °C -2.5 (-3.6 to -1.3) -0.3 (-1.0 to 0.2)
Mean temperature of coldest month °C -15.1 (-16.3 to -13.9) -13.2 (-13.9 to -12.6)
Mean annual precipitation mm 466 (462 to 470) 598 (592 to 604)
Potential evapotranspiration mm year-1 182 (153 to 210) 236 (218 to 248)
Water balance mm year-1 293 (203 to 371) 358 (324 to 409)
Theoretical solar radiation (mean) Mj cm-2 year-1 0.41 (0.34 to 0.48) 0.50 (0.36 to 0.59)
Theoretical solar radiation (std) Mj cm-2 year-1 0.04 (\0.01 to 0.16) 0.03 (\0.01 to 0.11)
Theoretical solar radiation (range) Mj cm-2 year-1 0.24 (0.03 to 0.74) 0.19 (0.02 to 0.56)
Topography
Elevation (mean) m a.s.l. 335 (144 to 509) 226 (145 to 338)
Elevation (std) m 17.9 (2.7 to 72.4) 8.5 (\0.1 to 44.2)
Elevation (range) m 75 (13 to 206) 34 (1 to 173)
Slope angle (mean) ° 4.7 (0.7 to 17.8) 4.2 (0.3 to 15.0)
Slope angle (std) ° 3.4 (0.6 to 14.8) 3.0 (0.4 to 11.8)
Slope angle (range) ° 18.3 (2.6 to 62.3) 14.2 (1.3 to 37.2)
Topographical wetness index – 9.5 (8.1 to 13.0) 9.7 (7.0 to 18.5)
(mean)
Topographical wetness index (std) – 4.3 (3.2 to 5.5) 2.4 (0.9 to 4.5)
Topographical wetness index – 20.6 (12.9 to 28.4) 19.1 (9.7 to 29.1)
(range)

A total of nine explanatory variables (in bold) were selected for modelling based on correlation analysis (see the
text for further details)
NB northern boreal, HA hemiarctic, a.s.l. above sea level

geology, geomorphology and hydrology groups using a comprehensive survey of ortho-

rectified aerial photographs (taken 2003–2009; ground resolution *30 cm) and topo-
graphical and geological maps (1:20,000–1:400,000). The group of geological features
included rock types and surficial ground materials. We classified the features of geo-
morphology according to the national system of geomorphological mapping (Fogelberg
and Seppälä 1986), which is based on international recommendations (Demek 1972).
Geomorphological features included process units and landforms from different geomor-
phological process groups (aeolian, biogenic, cryogenic, fluvial, glacigenic, glaciofluvial,
littoral and marine, polygenetic bedrock, slope and mass-wasting and weathering). The
group of hydrological features included springs, streams, rivers, ponds and lakes. The sizes
of the smallest mapped features varied from 1 to 2 m for point features (e.g. tors and erratic
blocks) to ca. 5 m for areal features (e.g. sand bars and wind deflation sites) (Supple-
mentary material 1).

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We estimated that the costs of compiling geodiversity data were higher (ca. 50–100 %)
when compared to the costs of the other explanatory variables (assuming free access to the
source data sets). However, the data costs are highly dependent on the study area (e.g.
availability of aerial photographs, geological maps, topographical and climate data). Thus,
the costs for topography and climate data may also be several times higher than geodi-
versity data. However, the costs to compile geodiversity data were considered to be several
tens of times less expensive than direct mapping of plant species distributions of the target
study areas.
We derived the climate variables from data provided by the Finnish Meteorological
Institute (1961–1990) (Venäläinen and Heikinheimo 2002) and topographical data (DEM,
grid cell size = 25 9 25 m) from the National Land Survey of Finland (NLS 2009). We
used a trend surface based on multivariate linear regressions to relate climatic variables to
the latitude, longitude and elevation of each study grid cell, downscaling climate data from
10 9 10 km resolution to a 1 and 0.25 km2 resolution (for details on the method, see Vajda
and Venäläinen 2003). The trend surface consisted of the nine terms of cubic trend surface
analysis [centred geographical coordinates (x, y) and the higher and cross-product terms
(x2, xy, y2, x3, x2y, xy2 and y3)] and the mean elevation of a grid cell (m above sea level,
a.s.l.) calculated for each study square. Besides linear terms, adding the quadratic and
cubic terms of the coordinates and their interactions into the model permits the detection of
more complex spatial features, e.g. spatial clumping or clustering. The elevation variable
enabled us to take into account the effect of local relief on the climatic variables. Vajda and
Venäläinen (2003) have shown in a study based on Finnish conditions that latitude (y) and
elevation (m a.s.l.) have the most pronounced influence on annual temperature (correlation
coefficients = -0.86 and -0.56), whereas the influence of longitude (x) was minor
(correlation coefficient = -0.07).
The compiled temperature and moisture-related climate variables were MAAT, mean
temperature of the coldest month (MTCM), MAP, potential evapotranspiration (PET) and
water balance (WAB) (Skov and Svenning 2004). Water balance was calculated as the
monthly difference between precipitation and potential evapotranspiration, and then
summing the separate monthly differences. The potential evapotranspiration was calcu-
lated following Skov and Svenning (2004):
PET ¼ 58:93  Tabove 0C =12: ð1Þ
WAB is a simple but widely used index that requires only monthly average values of
temperature and precipitation [Skov and Svenning (2004) and the references therein].
Despite its simplicity, this measure of WAB has been found to provide a valuable variable
for the distribution modelling of different taxa [see Skov and Svenning (2004), Luoto and
Heikkinen (2008) and Heikkinen et al. (2010) for successful applications]. Moreover, we
computed mean, standard deviation and range of theoretical solar radiation (SRAD) in the
study grid cells using a computer model of clear sky insolation and exposure of different
slopes (McCune and Keon 2002). The DEM-based topographical variables were mean,
standard deviation and range of elevation, slope angle and topographical wetness index
(TWI). The TWI was calculated using pit-filled DEMs (i.e. sinks were removed to ensure a
continuous drainage network) and the formula:
TWI ¼ lnðAs = tan aÞ ð2Þ
where ln denotes the natural logarithm, As represents the upslope-contributing area and a
represents the slope angle (Beven and Kirkby 1979).

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We acknowledge here one basic difference between the geodiversity and climate and
topography variables. Namely, the geodiversity variables were measures of diversity (i.e.
they described how many different elements were present in each grid cell). In contrast, the
climate and topography based explanatory variables were not diversity measures (e.g. how
many topographically different conditions were present in a given grid cell) but descrip-
tions of which particular elements were present in each grid cell. Nevertheless, the per-
formances of these variable groups were compared here with each other in the modelling,
because we deliberately selected the climate and topography variables to represent tradi-
tionally employed explanatory variables in species richness analyses and models, whereas
the geodiversity variables represented a new, alternative approach to compile explanatory
data (Field et al. 2009; Parks and Mulligan 2010).

Statistical analysis

The relationship between species richness and explanatory variables was analysed using
two different approaches, partial generalized linear modelling (GLM; McCullagh and
Nelder 1989) analyses (i.e. variation partitioning, VP) and generalized additive modelling
(GAM). More precisely, VP was used to identify the independent and shared contributions
of the geodiversity (i.e. measures of geological, geomorphological and hydrological
diversity), climate and topography variable groups in explaining the spatial patterns of
species richness in the study areas (Borcard et al. 1992; Hawkins et al. 2003). GAMs were
used to explore the ability of the different explanatory variable combinations in predicting
the plant species richness patterns within and between the study areas.
In VP, we applied the approach of Hawkins et al. (2003) by which the coefficients of
determination [here D2 = (null deviance - residual deviance)/null deviance] were used to
determine the independent and shared contributions of the explanatory variable groups.
Consequently, we calibrated seven GLMs and computed D2 using (i) climate, (ii) topog-
raphy, (iii) geodiversity, (iv) climate and topography, (v) climate and geodiversity, (vi)
topography and geodiversity variables and (vii) variables from all three groups. Based on
D2 values extracted from these seven separate GLM runs, we calculated the independent
and shared fractions for the three explanatory variable groups [for more details on the
calculation process, see Hawkins et al. (2003) and Heikkinen et al. (2004)]. GLMs were
calibrated using the statistical package R version 2.11.1 with standard glm functions (R
Development Core Team 2011), a Poisson distribution with a log link function in the
model fitting, and quadratic terms in addition to the linear terms of the explanatory vari-
ables to capture potential curvilinear responses. The selection of the explanatory variables
for the analyses was made so that correlations (Spearman’s rank order correlation, Rs)
within the variable groups were \0.7 in order to minimize collinearity problems (e.g.
Zimmermann et al. 2007). If Rs between two explanatory variables was [0.7, the one
correlated higher with the species richness variable was selected. In cases where the
correlation analysis showed rather equal performance of two explanatory variables, an
ecologically more relevant variable was selected. The variables for the final GLMs were
selected using a step-wise approach and Akaike’s Information Criterion (AIC) (Akaike
1974).
GAMs are semi-parametric extensions of GLMs in which the linear explanatory vari-
able is substituted with a smoothing function that can take various forms (Hastie and
Tibshirani 1990). The high performance of GAMs in predictive modelling has been evident
in biogeography and ecology studies (e.g. Guisan et al. 2007). GAMs are parameterized
just like GLMs, except that some explanatory variables can be modelled non-

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parametrically in addition to linear and polynomial terms for other variables. GAMs relate
the expected value (l) to the explanatory variables (xj) by
X
p
 
gðlÞ ¼ a þ fj xj ð3Þ
j¼1

where g is a link function, a is the constant (i.e. intercept) and fj are unspecified smooth
functions. [In practice, the fj are estimated from the data by using techniques developed for
smoothing scatterplots (Hastie and Tibshirani 1990).]
In GAMs, Rs between all explanatory variables was determined to be\0.7. The selected
variables were fitted to the species richness using a smoothing spline with the degrees of
freedom (d.f.) permitted to vary between one (i.e. straight-line relationship) and three (i.e.
nonlinear relationship). Similarly to GLMs, GAMs were calibrated using a step-wise AIC
approach, with standard gam function (R Development Core Team 2011). The spatial
autocorrelation (SAC) in the residuals of GLMs and GAMs were explored using Moran’s I
(e.g. Dormann et al. 2007; Bini et al. 2009).
GAMs were calibrated and evaluated using interpolation (i.e. within-area) and extrap-
olation (i.e. between-area) assessment of model performance. First, a split-sample
approach was used to split the study areas into semi-independent calibration and evaluation
data sets. The study areas were split into four parts (geographical split of the study areas
into south, west, north and east parts; each area = 25 % of the samples). Thus, the cali-
brated models were evaluated with three data sets (within-area or interpolative evaluation).
Second, the calibrated models were evaluated with data from the other study area
(between-area or extrapolative evaluation). We utilized the original data sets in the
assessment of between-area predictive ability of the GAMs to keep the extrapolation
setting straightforward. Differences in the predictive performances of the GAMs with
different sets of explanatory variables were tested using Student’s paired t test (Sokal and
Rohlf 1995).

Results

A total of nine explanatory variables were selected for the VP and six for the GAM
analyses (Tables 1, 2; Fig. 3). The set of variables for VP included all three measures of
geodiversity and six variables describing the climatic and topographical conditions of the
modelling grid cells. The climate variables MAAT, MTCM, PET and WAB correlated
equally well with species richness, but were highly intercorrelated (Rs commonly [0.9).
Due to the importance of temperature conditions at high latitudes, the MAAT variable was
selected for further analyses. In the GAM model building, all these nine explanatory
variables were considered simultaneously and again here, the highly intercorrelated vari-
ables were identified and excluded from the actual modelling. Consequently, mean ele-
vation, standard deviation of the slope angle and range of SRAD variables were excluded
from the following GAM analysis because they correlated highly with the MAAT and
geomorphological diversity variables.
Overall, the general form of the relationship between measures of geodiversity and plant
species richness was rather constant, i.e. the shape of the response functions showed a
positive trend in both areas indicating increasing species richness with increasing geodi-
versity (Fig. 3). However, in some of the cases an asymptotic curvilinear relationship was
observed in the model outputs. According to the residual plots of the final GLMs and

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Fig. 3 Univariate generalized additive modelling-based response curves for vascular plant species richness c
and explanatory variables used in variation partitioning analysis in Kevo (a) and Oulanka (b). The
explanatory variables were fitted to species richness using a smoothing spline with the degrees of freedom
permitted to vary between one (i.e., straight-line relationship) and three (i.e. nonlinear relationship)
(y axis = fitted function with estimated degrees of freedom). Explained variation (%) also appears in this
figure. GeolDiv geological diversity, GeomDiv geomorphological diversity, HydroDiv hydrological
diversity, MAAT mean annual air temperature, MAP mean annual precipitation, SradRange range of
theoretical solar radiation, ElevMean mean of elevation, SlopeStd standard deviation of slope angle,
TwiRange range of topographical wetness index

GAMs, the assumption of Poisson errors was appropriate. Moreover, SAC in the residuals
of the final models was generally rather low (Supplementary material 2). In Kevo, the
Moran’s I values in the first distance class ranged from -0.01 to 0.14 and from -0.19 to
0.22 for GLMs and GAMs, respectively. In Oulanka, the comparable values were
0.18–0.33 for GLMs and -0.10–0.35 for GAMs.
The results of VP were surprisingly similar in the two study areas (Fig. 4). The GLMs
with explanatory variables from all the three groups explained 51.7 % (Kevo) and 53.4 %
(Oulanka) of the variation in the plant species richness data. The shared contribution of all
the three groups of variables was notably high, explaining 23.6 % (Oulanka) and 25.6 %
(Kevo) of the variation. Considering pure fractions, the group of geodiversity measures
showed the highest pure explanatory power for the species richness patterns (9.1 and
11.2 %), while the climate variables showed the lowest (Fig. 4).
GAMs calibrated with measures of geodiversity were on average more robust than
models with only climate and topography variables, especially GAMs that included only
measures of geodiversity (Table 3). In both evaluation settings, models with measures of
geodiversity were better (statistically, from nearly significant to significant differences) in
predicting the plant species richness than models with climate and topography variables
(‘GD-models’ vs. ‘CLIMTOPO-models’) (Tables 3, 4). The inclusion of geodiversity
variables into the set of explanatory variables improved the predictive abilities of the
GAMs significantly in the within-area but not significantly in the between-area evaluation
(CLIMTOPO vs. ALL). Interestingly, models including only measures of geodiversity
showed a better predictive performance than models including variables from all the

Table 2 Spearman’s rank cor-

Explanatory variable Kevo Oulanka
relation coefficients between
(NB–HA) (NB)
vascular plant species richness
and explanatory variables selec-
ted for variation partitioning and Geodiversity
generalized additive modelling Geological diversity 0.49*** 0.42***
Geomorphological diversity 0.52*** 0.53***
Hydrological diversity 0.46*** 0.50***
Climate
Mean annual air temperature 0.47*** 0.46***
Mean annual precipitation -0.22*** 0.13**
NB northern boreal, HA
hemiarctic Theoretical solar radiation (range)a 0.36*** 0.45***
*** p \ 0.001, ** p \ 0.01 Topography
a
Variable excluded from the Elevation (mean)a -0.46*** -0.45***
generalized additive modelling Slope angle (std)a 0.38*** 0.46***
due to the high bivariate Topographical wetness index (range) 0.50*** 0.50***
intercorrelation

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Fig. 4 Results of the variation partitioning (VP) analyses in the study areas, based on a series of partial
regression analyses with generalized linear models. The figures describe the amount of explained variation
(%) in the species richness data as independent and shared contributions (U undetermined variation). Note
that the shared variation may be negative due to suppressor variables (i.e., a explanatory variable having low
correlation with the response variable and a correlation with another explanatory variable, which in turn is
correlated with the response variables) or due to two strongly correlated predictors with strong effects on
response of opposite signs (one positive and the other negative) (e.g. Legendre and Legendre 1998). The
climate group included the following variables: mean annual air temperature, mean annual precipitation and
range of theoretical solar radiation. Topographical variables were mean elevation, standard deviation of
slope angle and range of topographical wetness index. The geodiversity group included the variables
geological diversity, geomorphological diversity and hydrological diversity

groups (GD vs. ALL) (Table 4). Moreover, the models with measures of geodiversity
seemed to be more accurate in intra-area prediction of the plant species richness when
assessed based on the optimal fit line between predicted and observed values (i.e. slopes
were closer to 1 and intercepts to 0) (Fig. 5).

Discussion

We have shown here that consideration of explicit measures of geodiversity—not just

DEM-based topographical or ‘‘geomorphological’’ variables—can improve species rich-
ness models in three fundamental ways at the mesoscale resolution. Firstly, the inclusion of
variables describing geological, geomorphological and hydrological variability increased
the explanatory power of our species richness models. Secondly, the predictive ability and
accuracy of the species richness models were higher when measures of geodiversity were
included in the set of explanatory variables in addition to the commonly used variables
describing abiotic resources. In fact, our results suggested that models with only measures
of geodiversity showed a better predictive performance than models with explanatory
variables from all three groups. Thirdly, the inclusion of measures of geodiversity
improved the robustness of the models.
The two key rationales for using features of geodiversity to model and predict the
spatial variation in species richness patterns are the following: (i) the potential that long-
term targets in biodiversity conservation may be better achieved by focusing conservation
actions on preserving heterogenic abiotic units rather than on biological communities,
which are more subject to changes under changing environmental conditions, and (ii) lower
survey costs in mapping the geodiversity features of the areas compared to running
comprehensive biological surveys. In other words, when geodiversity is the focal site-
selection criterion and when sites with high geodiversity are protected, the aim is not to
preserve certain types of static biotic communities. Rather, the focus is on protecting areas

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Table 3 Results of generalized additive modelling. The mean of Spearman’s rank correlation coefficients
(Rs) between observed and fitted/predicted plant species richness values in model calibration and evaluation
data sets
Mean of R2 Mean of Rs Mean of Rs Change in Rs Mean of Rs
(adj.) in in calibration in intra-area (calibration ? in inter-area
calibration evaluationa evaluationa) evaluationb

Kevo
GD 0.42 0.65 0.57 -0.08 0.59
CLIMTOPO 0.38 0.64 0.43 -0.21 0.33
ALL 0.52 0.75 0.51 -0.24 0.38
Oulanka
GD 0.52 0.70 0.61 -0.09 0.49
CLIMTOPO 0.51 0.74 0.46 -0.28 0.24
ALL 0.62 0.80 0.59 -0.21 0.31
GD explanatory variables from the geodiversity group
CLIMTOPO explanatory variables from the climate and topography groups
ALL explanatory variables from all the three groups, geodiversity, climate and topography
a
Model calibration and evaluation in the same study area, e.g. Kevo
b
Model calibration and evaluation in different study areas, e.g. Kevo models evaluated in Oulanka

Table 4 Results of the Student’s paired t test for the comparison of the generalized additive model per-
formances in model evaluation. The model evaluation was performed in two different settings: interpolation
(i.e. intra-area prediction) and extrapolation (i.e. inter-area prediction)
Compared models Rs between observed and predicted Rs between observed and predicted
values in intra-area evaluation dataa values in inter-area evaluation datab

GD vs. CLIMTOPO p = 1.0-5 (GD)c p = 0.011 (GD)

CLIMTOPO vs. ALL p = 2.4-5 (ALL) p = 0.285 (ALL)
GD vs. ALL p = 0.007 (GD) p = 0.004 (GD)
GD explanatory variables from the geodiversity group
CLIMTOPO explanatory variables from the climate and topography groups
ALL explanatory variables from all the three groups, geodiversity, climate and topography
a
Model calibration and evaluation in the same study area, e.g. Kevo
b
Model calibration and evaluation in different study areas, e.g. Kevo models evaluated in Oulanka
c
model with higher predictive ability is indicated in brackets

with a high probability of harbouring high biodiversity, even if the species composition of
these areas would change over time (e.g. Hunter et al. 1988). Consequently, geodiversity
conservation has the potential to provide a useful approach for long-term preservation of
biodiversity that may be applied across a range of spatial scales (Anderson and Ferree
2010; Beier and Brost 2010; Parks and Mulligan 2010).
The measures of geodiversity used here, i.e. diversity of geology, hydrology and geo-
morphology, were rather clearly linked to plant species richness, probably because of their
contribution to one or more important resource factors (cf. Vanreusel et al. 2007). Geology
(e.g. rock type) and geological processes (e.g. weathering) contribute to the availability of
important resources for plants by providing them with a substrate and nutrients (e.g. Birks
1996; Heikkinen et al. 1998; Moser et al. 2005). Moreover, different rock types and

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Fig. 5 Scatter plots between predicted and observed vascular plant species richness in the intra-area
evaluation data. The predictions were derived using generalized additive modelling (GAM) (see the text for
further details). The example plots were selected to describe good (a–c, Oulanka) and moderate (d–f, Kevo)
predictive success. Plots (a) and (d) were derived using GAMs based on geodiversity variables (i.e.
measures of geodiversity); plots (b) and (e) were derived using GAMs based on the climate and topography
variables; and plots (c) and (f) were derived using GAMs based on the geodiversity, climate and topography
variables

surficial deposits increase the environmental heterogeneity and number of different habi-
tats in an area. The positive relationship between plant species richness and hydrological
diversity supports earlier findings that moisture availability is one of the critical factors for
plant species distribution and richness (Gosselink and Turner 1978). However, the linkage
between species richness and hydrological diversity goes beyond this general maxim and
supports the observations of Nichols et al. (1998) that the within-area variation of water
availability also significantly contributes to the number of species that a given landscape
can support (see also Lavers and Field 2006). For example, streams and rivers were
considered as different features in hydrological diversity because their characteristics can
differ significantly: drastic floods in river environments and ephemerality of small streams
give rise to different growing conditions for plants.
In the context of space and spatial heterogeneity, geomorphological diversity is linked
to plant species richness because it partly controls the variability of habitats. Moreover,
geomorphological diversity contains features of various sizes, from small-sized fluvial
features to extensive fracture valleys, which provide habitats with different sizes and
ecological characteristics. A more diverse landscape consisting of various different abiotic
habitats and structural organisms provides a setting for a wider number of niches available
for species to occupy (Menge and Sutherland 1976; Tilman 1994; Moser et al. 2005).
Physical disturbance is regarded as an important driver of biodiversity (e.g. Tilman
1982; Huston 1994) which may have a particularly notable significance in high-latitude
landscapes (Birks 1996; Virtanen et al. 2010). Physical disturbance associated with soil

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instability and other geomorphological processes causes a decrease in biomass (Grime

2001), and changes in resource availability (Tilman 1982) and species interactions
(Crawley 1986). Disturbances often counteract competitive exclusion and enhance the
spatial heterogeneity of vegetation, and both of these factors may have a positive influence
on species richness (Huston 1994). In our study, geomorphological diversity included
several process types that actively modify the landscape under current conditions (Hjort
and Luoto 2009, 2010). Physical processes, especially those related to running water and
slopes, generate fine-scale dynamical disturbances and patches free of competition in the
vegetation cover. These processes often results in overall enhanced biodiversity on a given
site. In an extreme case, geomorphological processes may create novel ecospaces that can
be colonized by species new to the area (e.g. Benton 2009).
The measures of geodiversity used in this study are simple and easy and rapid to
employ, but yet rather explicit. Moreover, they seem to convey information that is not
covered by the commonly utilized abiotic variables (Whittaker et al. 2001; Willis and
Whittaker 2002; Field et al. 2009). However, our measures of geodiversity did not include
direct estimates of climate or other energy-related components (e.g. Serrano and Ruiz-
Flaño 2007; Benito-Calvo et al. 2009). Earlier studies have shown that the energy-related
factors (e.g. evapotranspiration, temperature and productivity) may explain up to 90 % of
the variation in biodiversity patterns (Mittelbach et al. 2001), and the importance of
energy-related factors is especially pronounced in broad-scale studies (Currie and Paquin
1987; Wright et al. 1993; Kerr and Packer 1997; Parks and Mulligan 2010). However, our
analyses were performed at the landscape scale at which the abiotic heterogeneity (e.g.
geomorphology and hydrology), in addition to properties of vegetation, controls micro-
climatological variability (Geiger 1965; Hayden 1998). Thus, the energy component was
indirectly included in our measures of geodiversity. In addition, it should be noted that
other factors often begin to show relatively more importance for species richness than
energy-related factors when switching the resolution from broad biogeographical scales to
a landscape or local scale (Wright et al. 1993; Whittaker et al. 2001; Willis and Whittaker
2002; Luoto et al. 2007). Nevertheless, we acknowledge that when the exploration of
process-based links between geodiversity and biodiversity (i.e. mechanistic analysis) or
prediction of biodiversity at national or continental scales is the focus of the study, the
incorporation of energy-related variables in geodiversity will no doubt have a higher
relevance (Parks and Mulligan 2010).
Our analyses indicate that measures of geodiversity were the best single variable group
for explaining plant species richness at the landscape scale. However, a well-known
limitation is that species richness is a measure that conveys no information on the actual
species composition of a community or on the habitat composition of a landscape (Luoto
et al. 2004). Correspondingly, geodiversity is a landscape summary measure that does not
take into account the uniqueness or potential ecological importance of different habitats.
This may limit its usefulness under certain circumstances. Further, there are situations in
which increasing geodiversity may contradict management objectives for threatened spe-
cies that require large and homogeneous habitat patches of a specific habitat type, since
higher habitat diversity may actually be a reflection of undesirable fragmentation of critical
habitats (McGarigal and Marks 1994). These challenging features of the geodiversity
measures may give rise to biased predictions in the models from the conservation and
management point of view, especially for species that require large areas of suitable habitat
(see Luoto et al. 2004; Hanski 2005).
There are also some other potential data- and method-related limitations to be
acknowledged. First, our study included plant species data from only two study areas with

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particular environmental conditions. Thus, without testing, our findings cannot be applied
to other types of environments, landscapes and biomes. For example, vegetation in more
topographically uniform boreal landscapes can be rather homogenous when compared with
mountainous regions. Second, the utilized air temperature and precipitation variables were
downscaled from a coarser resolution, which probably underestimated the importance of
the climate variables and may have influenced the results. On the other hand, it should be
noted that the solar radiation and topographical variables employed in our study very likely
cover part of the microclimatological variability at the modelling resolution (Geiger 1965;
Hayden 1998).
A general assumption in biogeographical species richness modelling studies has been
that climate plays a less significant role than other abiotic conditions at the landscape scale
(Pearson and Dawson 2003). Thus, the concept used in this study should be developed
further and tested on a broader sampling scale, particularly on a wider sampling extent.
Also the approach to measure geodiversity could be developed further. For example, more
ecologically relevant measures of geodiversity might be outlined by considering the habitat
requirements of the plants in the target area. This target could be approached by weighting
different features of geodiversity according to their importance for plants (e.g. higher
weights for limestone and gully than granite and erratic block, respectively) (cf. Ruban
2010). Overall, the development of ecologically relevant measures of geodiversity can be
arguably included among the key tasks in geodiversity research in future (Gordon et al.
2012).
Methodologically, the relatively small data sets employed here hampered the fully
independent evaluation of the models. Given this, our statistical test results may be over-
optimistic. Moreover, the presence of SAC in model residuals may increase type I error
rates (falsely rejecting the null hypothesis of no effect). However, SAC in the residuals of
GLMs and GAMs developed in this study were rather low (Dormann et al. 2007; Bini et al.
2009). Thus, we consider that the potential influence of residual SAC for the interpretation
of our results would be low. Another point which should be acknowledged is that the
relationships between species richness and measures of geodiversity were in all cases
positive, but they were not always linear throughout (cf. Nichols et al. 1998). If the
threshold is not known, the asymptotic levelling off effect may complicate the accurate
prediction of high species diversity sites.
Despite these limitations, our results provide a clear signal that the inclusion of explicit
and directly measured geodiversity variables have the potential to improve species richness
models and provide novel insights into biodiversity surveys (Swanson et al. 1988; Nichols
et al. 1998; Lobo et al. 2001; Pausas et al. 2003; Illán et al. 2010). When the interpretation
of the variability of geological, geomorphological and hydrological features is done sys-
tematically and with caution, they may provide a more reliable basis for transferring
biodiversity models to new areas than topographical or climatic variables especially, at
scales, where the effect of climate is expected to be rather weak.

Conclusions

A long-term challenge in biodiversity research has been the development of robust

methods for cost-effective targeting of conservation actions. Here we compared the
explanatory and predictive performance of directly measured variables of geodiversity and
commonly used abiotic surrogates of biodiversity in modelling and have shown that the
inclusion of explicit measures of geodiversity can improve the explanatory power,

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predictive ability and robustness of the mesoscale plant species richness models in the
boreal zone. Thus, the measures of geodiversity employed, namely geological, geomor-
phological and hydrological diversity, appear to be promising surrogates of biodiversity
which both directly and indirectly reflect important abiotic resource factors. In areas with
insufficient climate and topography data, simple measures of geodiversity may provide the
best surrogates for biodiversity patterns.

Acknowledgments We would like to express our gratitude to Paul Beier and two anonymous reviewers
for their helpful and constructive comments. Moreover, we would like to acknowledge Lauren Martin for
her help in checking the English of the revised manuscript.

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