Ecography 36: 800808, 2013

doi: 10.1111/j.1600-0587.2012.07922.x
2013 The Authors. Ecography 2013 Nordic Society Oikos
Subject Editor: Catherine Graham. Accepted 27 November 2012

Geomorphological disturbance is necessary for predicting ne-scale

species distributions
Peter C. le Roux, Risto Virtanen and Miska Luoto
P. C. le Roux (peter.leroux@helsinki.) and M. Luoto, Dept of Geosciences and Geography, Univ. of Helsinki, FI-00014 Helsinki, Finland.
R. Virtanen, Dept of Biology, Univ. of Oulu, FI-90014 Oulu, Finland.

Disturbances related to geomorphological processes are frequent, widespread and often intense at high latitudes and altitudes, aecting the ne-scale distribution of many plant species. While the inclusion of physical disturbances into models
of species geographic ranges is widely recommended, no studies have yet tested the utility of eld-quantied geomorphological disturbances for terrestrial species distribution modelling. Here we apply generalized additive models and boosted
regression trees to examine if the explicit inclusion of terrestrial and uvial geomorphological variables alters species distribution models for 154 vascular plant, bryophyte and lichen species in north European mountain tundra. The inclusion of these disturbances signicantly improved both the explanatory and predictive power of distribution models, with
consistent results for all three species groups. Spatial distribution predictions changed considerably for some species after
the inclusion of disturbance variables, with uvial disturbances generating strongly linear features for species inuenced by
erosion or sediment deposition. As a consequence, models incorporating geomorphological variables produced markedly
more rened distribution maps than simpler models. Predictions of species distributions will thus benet strongly from
the inclusion of ne-scale geomorphological variables, particularly in areas of active earth surface processes, enabling more
accurate forecasting of future species ranges under changing conditions.

A wide variety of community properties are inuenced

by disturbance events, including species richness (Connell
1978, Huston 1994), abundance (Sousa 1984) and ecosystem productivity and functioning (Cardinale and Palmer
2002, Kreyling et al. 2008). Disturbances can strongly
alter the spatial and temporal heterogeneity of conditions
within landscapes, favouring some species and excluding
others (Huston 1994). As a result, explicitly including
variables that reect disturbance processes in species distribution models should improve model performance (Guisan
et al. 1998, Dirnbck and Dullinger 2004, Guisan and
Thuiller 2005), and thus make a step towards the modelling
of realistic species distributions. However, despite the clear
link between disturbance regimes and spatial variation in
species abundance and occurrence, few studies incorporate
ne-scale disturbances of any kind when modelling the
distribution of individual species (Austin and Van Niel 2011).
Disturbance regimes at high latitudes and altitudes may
be dominated by earth surface (i.e. geomorphological)
processes, which are known to strongly aect the ecological
conditions in these environments (Sigafoos 1952, Krner
2003, Nagy and Grabherr 2009) and have been hypothesized
to be important determinants of species distributions
(Randin et al. 2009). Water- (uvial erosion and sedimentation), frost- (cryoturbation and soliuction), snow(nivation) and wind-related (deation) processes can reduce
800

the biomass of individual plants and the size of populations

and alter the competitive environment (Sigafoos 1952,
Grime 1977, Nagy and Grabherr 2009). These geomorphological processes also aect a diversity of environmental
characteristics, including soil stability, chemistry and nutrient availability (Jonasson and Skld 1983, Jonasson 1986,
Kreyling et al. 2008), and temperature and moisture regimes
(Krner 2003, Walker et al. 2004, Malanson et al. 2012).
As a result, these processes may exert strong control over
plant performance, as well as the nature of interactions
between species (Grime 1979). For example, uvial processes, including erosion, sedimentation and ooding,
can cause partial or total destruction of vegetation (Hupp
1988, Wohl 2000). Indeed, at high latitudes and altitudes
the landforms associated with these geomorphological
processes essentially characterise the micro- and mesoscale
environmental heterogeneity of landscapes (Dahl 1956,
Hupp and Osterkamp 1996), and therefore, can be expected
to be drivers of ne-scale species distributions.
Several studies have examined the inuence of biotic
disturbances (e.g. herbivory, digging, trampling) and infrequent, large-scale disturbances (e.g. res, storms, ooding)
on species geographic distributions (e.g. Moretti et al.
2006), but the inuence of geomorphological processes
on species distribution patterns has received much less
attention. The study by Randin et al. (2009) was the rst to

attempt to explicitly include geomorphological processes

in species distribution modelling. However, that study did
not show improvement in models predictive power, probably, as highlighted by the authors, due the semi-quantitative
disturbance variable being insuciently precise and too
coarse-scaled to be truly ecologically meaningful. Similarly,
Mellert et al. (2011) developed distribution models incorporating estimates of uvial activity based on soil maps,
but found this variable to be largely redundant for model
performance. Therefore, the potential for ne-scale eldquantied geomorphological variables to contribute to species distribution models remains poorly understood.
There are broad generalities in species responses to disturbances (and environmental factors in general; Grime 1979)
but only limited consistency in their specic reactions to
changes in environmental conditions (Chapin and Shaver
1985, Dormann and Woodin 2002). Geomorphological
factors are unlikely to be an exception to this trend
and, therefore, markedly idiosyncratic responses may be
expected. It is possible, for example, that frost-heave may
preclude some species from habitats (Prez 1987), but
simultaneously be essential for the occurrence of others
(Nylhn and Totland 1999). These potentially important
individualistic relationships between species distributions
and geomorphological processes remain essentially uninvestigated. Thus, it is necessary to examine the inuence of
geomorphological variables on a large and representative
set of species if community-level responses are to be estimated. Furthermore, modelling the distribution of multiple
species oers the possibility of identifying general patterns
that can be applied to taxonomical, biogeographic or functional categories (Guisan and Zimmermann 2000).
Here we examine if the incorporation of eld-measured
geomorphological variables can signicantly improve the
accuracy and predictive power of species distribution models
for 154 bryophytes, lichens and vascular plant species in
north-western Finland and Norway. This is done by applying
generalized additive models and boosted regression trees to
a unique dataset comprising ne-scale vegetation surveys of
vascular plants, bryophyte and lichen species and eldbased quantication of active geomorphological processes,
recorded in 1080 survey plots spanning a wide range of
environmental conditions characteristic of mountain systems in high northern latitudes. Additionally, the impact
of geomorphological disturbance on species richness patterns is examined by summing individual species distribution predictions from models with and without the
disturbance variables included.

Methods
Data collection
Data were collected from 270 sites in north-western
Finland and Norway, along 18 altitudinal transects in the
sub-continental northern Scandes (ca 69N, 2050E).
Four 1 m2 plots, located 5 m from the centre of each site,
were surveyed at each site, resulting in a total of 1080
plots for analysis. The cover and identity of all vascular
plants, bryophytes and lichens in each plot were recorded

(see Virtanen et al. 2010, le Roux et al. in press for more

details). Species were classied into two broad categories
by biogeographic distribution: arctic-alpine (main distribution in cold arctic-alpine areas) or boreal (having a broader,
often Holarctic, distribution).
Environmental variables for modeling species distributions were selected from three dierent groups of commonly
used variables (macro-climate, soil resources and solar
radiation), after a priori exclusion of strongly collinear variables. Three climate variables were calculated for each plot:
mean temperature of the hottest month (July), mean
temperature of the coldest month (January), and total
annual precipitation. Climatic data were obtained from the
Finnish Meteorological Inst. data set (Venlinen and
Heikinheimo 2002), with average values from 1971 to
2000 downscaled from the original 10-km grids following
the methodology of Vajda and Venlinen (2003; linear
regression using latitude, longitude and altitude terms).
This approach accounts for adiabatic lapse and has
performed well in this region (Sormunen et al. 2011). As
all three downscaled climate variables were strongly correlated (|r|  0.75, p  0.001), only the mean temperature
of the hottest month (hereafter TempJuly) was used in subsequent analyses.
Soil quality was estimated from the proportion of
calcareous (nutrient-rich) and silicaceous (nutrient-poor)
bedrock at plots. Bedrock data was extracted from Korsman
et al. (1997) at a resolution of 100 m and interpolated
using the ArcGIS interpolation tool (ESRI 1991). Analyses
across a subset of the plots show that soil quality is signicantly correlated with soil nutrient content (Vuollet and
Luoto unpubl.). Relative soil moisture at each plot was
estimated by calculating the topographic wetness index
(TWI; Beven and Kirkby 1979) from a 10 m resolution
digital elevation model (DEM; National Land Survey of
Finland) using the ArcView Spatial Analyst extension (ESRI
1991). TWI was calculated as:
TWI  ln (upslope catchment area per unit width
orthogonal to the ow direction  tan[slope angle])
with slope angle measured in radians. Potential solar radiation (Mj/cm2/a; Radiation) for each plot was calculated
from the DEM using Solar Analyst extension in ArcView
(ESRI 1991, McCune and Keon 2002). This value reects
the maximum potential radiation, based on slope aspect
and angle, assuming clear skies. Plot slope and soil moisture
balance (i.e. the dierence between precipitation and
potential evapotranspiration; calculated following Maggini
et al. 2006) were both considered as potential explanatory
variables, but were not included in analyses due to being
strong correlated with TWI (r  0.89, p  0.001) and
climate variables (|r|  0.81, p  0.001) respectively.
Geomorphological activity was estimated for each site
by recording the cover of two sets of variables at each site
(10  10 m). The cover of active terrestrial (Disturbterr:
including frost-related cryoturbation and soliuction, and
wind-related nivation and deation) and uvial (Disturbuv:
uvial erosion and deposition) geomorphological processes
was estimated in situ during the summers of 20092011 as
percentage cover following the methodology of Hjort and
801

Luoto (2009, 2011) and Virtanen et al. (2010). All surveys

were performed by the same geomorphologist to exclude
variation resulting from observer dierences, with the
investigator focusing only on geomorphological variables
to ensure independence of the botanical and geomorphological data. The activity of features was dened based
on observations of topsoil material (e.g. frost heaving and
cracking, mass wasting, soil displacement, as well as uvial
erosion and sedimentation). This geomorphological disturbance measure thus combines many physical disturbance
factors potentially causing biomass loss, excluding the
biotic disturbances caused by grazing animals (Virtanen
et al. 2010). Disturbance values were log-transformed
prior to analysis to reect the non-linear relationship
between the cover and the intensity of geomorphological
activity (i.e. a doubling of the cover of a geomorphological
disturbance was assumed to result in more than a two-fold
increase in its eect on the vegetation). In this system
we consider geomorphological processes to be chiey disturbances (i.e. processes removing biomass; sensu Grime 1979),
with any additional eects on resource availability assumed
to have only a secondary impact relevant to a subset of
species that tolerate disturbance. We did not investigate res
as an additional physical disturbance in this study as wild
res are very infrequent in the northern European mountain
tundra.
Statistical analysis
Since dierent species distribution models may potentially
produce dierent predictions (Franklin 2009), two alternative state-of-the-art methods were used: generalized additive models (GAMs) and boosted regression trees (BRTs).
Both techniques are data-driven, capable of modeling
non-linear functions, and widely used to model species distributions (Elith et al. 2006, Franklin 2009). The original
dataset was randomly split into a calibration (i.e. training;
75% of records) and a validation (i.e. testing; remaining
25% of records) dataset. Only species with at least 15
records in the calibration dataset and 7 records in the validation dataset were included in analyses (limiting analyses to
only more abundant species did not give dierent results).
GAMs are a non-parametric extension of generalized
linear models that use smoothers to estimate the relationship between response and predictor variables (Yee and
Mitchell 1991). GAMs were tted in R statistical software
(R Development Core Team), using the mgcv package
(Wood 2011) which calculates the extent to which the
degrees of smoothness for each smoother can be reduced
during model tting without signicantly lowering model
t. These models were tted assuming a quasibinomial
distribution of errors (applying a logit link function), setting
the initial degrees of smoothness for each univariate term
at four.
BRT is a machine learning method that estimates the
relationship between a response variable and its predictors
without a priori specication of a data model (Death
2007, Elith et al. 2008). This technique combines large
numbers of simple tree-based models to form a nal model
optimized for prediction, using cross-validation for model
802

building. BRTs can model complex functions and automatically incorporate interactions between predictors. As
GAMs and BRTs gave similar results, only those from the
GAM analyses are presented here (detailed BRT methodology and results are presented in Supplementary material
Appendix 1, Table A1 and Fig. A1, A2).
To test if the inclusion of the two disturbance variables
signicantly improved the t of species distribution models,
we followed the methodology of Zimmermann et al.
(2009) and le Roux et al. (in press), rst modeling each species occurrence as a function of simple topographysoil
climate variables:
Presence/Absence  Soil moisture  Radiation
 Soil quality  TempJuly
(simple model)
Next, we incorporated terrestrial and uvial geomorphological disturbances into the model
Presence/Absence  Soil moisture  Radiation
 Soil quality  TempJuly  Disturbterr
 Disturbuv
(full model)
Simple and full models for each species were tted to the
calibration dataset and an ANOVA (implementing a Chi2
test) was used to test if the full model provided a signicantly
better t to the calibration data than the simple model.
The explanatory power of both models was assessed by
comparing the adjusted R2 for all relationships (i.e. the
proportion of variance explained, after accounting for the
dierent numbers of explanatory variables in simple and
full models). The relative importance of predictor variables
was assessed from calibration models from each variables
drop contribution (i.e. change in deviance associated
with exclusion of a given variable from a model containing
all the other variables). Variables contributions were
scaled so as to sum to 100, with higher numbers indicating
stronger inuence on the response.
The predictive power of the models was determined
by testing the accuracy of predictions made for the validation dataset (i.e. data that was not used in model development), calculating the area under the curve of a receiver
operating characteristic plot (AUC; Fielding and Bell
1997) and the true skill statistic (TSS; Allouche et al.
2006). A non-parametric Wilcoxons matched pairs test
was used to compare if the explanatory power and predictive accuracy of the simple and full models diered
signicantly. Both explanatory accuracy and predictive
ability of models are considered, so as to evaluate models
ability to explain local phenomena and to predict to independent data.
Predicting species ne-scale distribution
To visualize the distribution patterns predicted by each
species best-t distribution models (and the resulting patterns of species richness), an area of 3.1  5.2 km in the
centre of our study region (covering the Saana and, partially,
Jehkats massifs) was divided in 25  25 m cells. A 1 m2

plot in the centre of each cell was used for all calculations
and predictions. Soil moisture and solar radiation for each
plot was calculated from the 10 m resolution DEM, and soil
quality and mean temperature of the hottest month were
downscaled from the available data (following the methods
described above). The intensity of geomorphological disturbances were mapped across the prediction area using
the DEM, false-colour aerial photographs (0.5 m spatial
resolution), and extensive eld surveys. From the cover
of these active terrestrial and uvial geomorphological processes the intensity of geomorphological disturbances was
estimated for each plot in the prediction area (following
Hjort and Luoto 2009, 2011, Virtanen et al. 2010;
Supplementary material Appendix 2, Fig. A3). Best-t
simple and full models were then used to predict the occurrence of each species across this landscape, with species
richness for each cell calculated as the sum of predicted
species occurrences (repeated for simple and full models).
Predicted probabilities of occurrence were converted to
presence/absence predictions using the threshold value maximizing sensitivity  specicity (Liu et al. 2005).

Results
Models including the two geomorphological disturbance
variables provided a better t to calibration data than simple
topographysoilclimate models, with 81% of full models
(124 of 154 species with adequate records in the calibration

Table 1. Results from Wilcoxons matched pairs tests, testing if

the explanatory accuracy and predictive power of full models
(including geomorphological variables) differs from that of simple
models (containing topography, soil and climate variables). Explanatory accuracy is measured as adjusted R2, and predictive power
as the area under the curve of a receiver operating characteristic plot
(AUC) and the true skill statistic (TSS). n  154.
Metric
2

Adjusted R
AUC
TSS

Simple model
(mean SE)

Full model
(mean SE)

Wilcoxon
V

19.5 1.0
82.5 0.7
51.0 1.3

25.7 1.1
84.5 0.6
54.1 1.3

27
1888
3461

 0.001
 0.001
 0.001

and validation data sets) having signicantly lower deviance

than equivalent simple models (ANOVA test on GAM models; Table 1). Full models also had better explanatory power,
with signicantly higher adjusted R2 (mean increase  6%,
max. increase  33%; Table 1, Supplementary material
Appendix 2, Table A2). Temperature of the warmest month
was on average the most important predictor of species
occurrence in both simple and full models (Fig. 1a). However,
when including terrestrial and uvial disturbance in models
the importance of TempJuly declined strongly (e.g. average
importance dropping from 58 to 32%), with Disturbterr
becoming the second most important variable. The importance of the two geomorphological disturbance variables
diered strongly between species, although their combined
importance ranged between 10 and 40% for most species
(Fig. 1b). Thus the contribution of the disturbance variables

Figure 1. The relative importance of explanatory variables as calculated from generalized additive models: (a) variable importance for the
simple (D: topographysoilclimate variables) and full (D: topographysoilclimatedisturbance) models, and (b) the distribution
of the combined importance of terrestrial and uvial disturbances for all species. In (a), empty circles indicate the maximum importance
for each variable across all 154 species modeled and standard error bars are omitted as they are too small to display clearly (all SE  2).
Rad.  solar radiation, Moist.  soil moisture, Qual.  soil quality, Temp.  mean July temperature, Fluv.  uvial disturbance, Terr. 
terrestrial disturbance.

803

varied considerably between species, but was strong enough

on average to signicantly improve model explanatory
power.
Model predictive performance improved with the
inclusion of the two disturbance variables, with small but
signicant improvement in AUC values for full models
(mean increase  0.02, max. increase  0.23; Table 1).
Mean TSS values were also signicantly higher for
models incorporating the two disturbance variables (mean
increase  0.03, max. increase  0.47; Table 1). Overall,
most GAM models had higher AUC (77% of models) and
TSS values (66%) for full models than simple topography
soilclimate models. Analysing species subsets based on
taxon (bryophyte, lichen or vascular species) or biogeographic
distribution (arctic-alpine or boreal species) gave similar
results to the initial analysis of all species (Supplementary
material Appendix 2, Table A3).
Response curves for the disturbance variables varied
greatly between species (i.e. from strongly positive to strongly
negative impacts on species probabilities of occurrence; see
e.g. Saussurea alpina in Fig. 2; all data in Supplementary
material Appendix 2, Table A2). Vascular plant species
response curves for uvial disturbance were most commonly
positive, with lichens predominately responding negatively
to increasing uvial disturbance, and bryophytes exhibiting an intermediate response (Supplementary material
Appendix 2, Table A4). Vascular plants and bryophytes
generally responded positively to increasing terrestrial

Figure 2. Response curves for Saussurea alpina as estimated by

generalized additive models. Solid lines indicate the response
curves for the simple model, with dashed lines indicating the
response curves from the full models. Temp. July  mean July temperature, Dist. uv.  uvial disturbance, Dist. terr.  terrestrial
disturbance.

804

disturbance, with more mixed responses from lichen species.

Comparing the response curves of arctic-alpine and boreal
species showed the former group to be positively aected
by terrestrial disturbance, while the latter species were
clearly negatively impacted by higher cover of terrestrial
geomorphological disturbances (full results and additional
analysis in Supplementary material Appendix 2, Table A4).
Across our interpolation extent the clearest dierence to
the predicted distributions between simple and full models
is the addition of linear elements into species occurrence
patterns, reecting some species strong (positive or negative)
responses to uvial disturbance (see for example Saussurea
alpina in Fig. 3). Changes in species occurrence patterns due
to terrestrial disturbances were more evenly distributed
throughout the landscape, and thus not as noticeable.
Predictions of species richness (derived by summing predictions for individual species) were similar between simple
and full models, although the eects of uvial disturbance
were clearly visible, having a positive eect on the richness of
vascular plants, but a negative eect on the number of
species of bryophytes and lichens (Supplementary material
Appendix 2, Fig. A4).

Discussion
The results from this study clearly support previous
hypotheses (Dirnbck and Dullinger 2004, Randin et al.
2009) that the inclusion of geomorphological processes
improves predictions of species distributions. This is in
agreement with the perceived role of geomorphology in
inuencing plant ecology in high latitude and altitude
ecosystems (Krner 2003, Nagy and Grabherr 2009,
Malanson et al. 2012). Our results show that incorporating
measurements of terrestrial and uvial geomorphological
processes strengthens species distribution models explanatory accuracy (i.e. ability to model the observed data)
and predictive power (i.e. ability to extrapolate results to
independent datasets). These ndings were consistent
for vascular plants, bryophytes and mosses, despite fundamental dierences between these groups of species (Molau
and Alatalo 1998), highlighting that the inclusion of
geomorphological variables benets species distributions
models across multiple taxa. Thus, geomorphological variables have non-trivial eects for the predictions of the current species distributions, and these should be carefully
considered in the context of constructing climate change
impact models.
Geomorphological variables exhibited considerable
importance for the majority of species, and very high
importance for a subset of the species (consistent with
Randin et al. 2009). Therefore, in agreement with other
studies examining the inuences of disturbances (Tucker
et al. 2012), species showed a wide range of responses to the
same geomorphological disturbances. Consistent improvements in model performance across all species grouping
(vascular plantsbryophyteslichens, and borealarcticalpine) suggests that inclusion of geomorphological variables can improve species distribution models for dierent
taxa. However, species response curves reveal that these
groups dier in their dominant responses to uvial and

Figure 3. Modeled distribution of Saussurea alpina in a 3.1  5.2 km area in the centre of our study region, based on the predictions of
(a) simple and (b) full generalized additive models. Grey indicates predicted areas of occurrence, and black indicates lakes and ponds.
Latitude and longitude are indicated in the Finnish coordinate system with contours representing altitude (m a.s.l.).

terrestrial geomorphological disturbances. For example,

arctic-alpine species showed more strongly positive responses to high levels of geomorphological disturbance than
species with boreal distributions. This agrees with previous
ndings that geomorphological disturbances are necessary
for the occurrence of some species outside their typical
ranges (Jonasson and Skld 1983) and suggests that earth
surface processes may be a prerequisite for many of the small
arctic-alpine species (e.g. Saxifraga foliolosa, Ranunculus
nivalis and Cardamine bellidifolia; Hmet-Ahti et al.
1998). By contrast, boreal species which are known to be
disturbance-intolerant and restricted to relatively undisturbed sites, including shrubs and dwarf shrubs (e.g. Betula
nana; Jonasson 1986, Oksanen and Virtanen 1995), generally exhibited a negative response to geomorphological
disturbance. As our study site encompasses the ecotone
between the arctic-alpine and boreal biomes it is interesting
how these two co-occurring species groups dier strongly
in response to geomorphological disturbances, suggesting
important dierences in their relative sensitivity to these
processes. Thus, while some species are clearly more strongly
aected by geomorphological disturbances than others, we
are presently unable to a priori predict species sensitivity to
these processes, and therefore these geomorphological variables should be incorporated whenever possible in ne-scale
species distribution models.
The geographic distribution projections including
geomorphological variables produced much more detailed

distribution maps than traditional models with only

topographysoilclimate variables. For instance, the inclusion of these disturbance processes markedly rened the
areas predicted to be suitable for many species. If not
included, model predictions grossly exaggerated or underestimated some species potential ranges (e.g. Fig. 3), emphasizing the need for the inclusion of ecologically meaningful
and suciently ne-scale variables for realistic species distribution models (Sormunen et al. 2011). However, we believe
it is still necessary to test the robustness of these results
with data from other environments to fully understand the
generality of these ndings. In agreement with Randin
et al. (2009), we also observed changes in the connectivity
of suitable habitat when disturbance variables were
included, chiey due to greater ne-scale patchiness. Due
to the strong contagion of cells containing uvial disturbance, it appears that habitats with uvial inuence
have clear potential to act as migration corridors for those
species tolerant of disturbance and responsive to warming
temperatures.
Geomorphological processes frequently generate low
intensity small-scale disturbances. Such geomorphological
disturbances maintain similar conditions from year to year,
with the spatial positions of these processes being relatively
stable over decades (French 2007). The scale at which these
disturbances operate remains unknown but it is likely that
their incorporation at a resolution of 1 m is adequate for
many purposes. Even though species distributions within
805

disturbed systems are essentially unpredictable at very ne

scales (Levin 1992), the incorporation of geomorphological
disturbances at this scale was useful. Indeed, the scale
sensitivity of disturbance for species distributions remains
to be studied further. Furthermore, in systems characterized
by infrequent and large-scale disturbances (e.g. avalanches
or landslides) more complex modelling approaches may
be necessary to capture the successional dynamics initiated
by disturbance events (Franklin 2010). Species distribution
models would, for example, at least require the inclusion
of time since the disturbance as a covariate (Moretti et al.
2006). Recent studies have suggested that biotic disturbances should also be incorporated into species distribution
models (Munier et al. 2010), although this too may require
further methodological developments. Indeed, the frequent low intensity impacts of burrowing, browsing
and grazing animals or the infrequent (but often devastating) outbreaks of insects and lemmings can strongly inuence high altitude and latitude vegetation (Fox 1985,
Ravolainen et al. 2011). However, due to uctuations of
herbivore populations and the patchy occurrence of their
eects, documenting the spatio-temporal variability of
biotic disturbances has been dicult (but see Virtanen et al.
2002) and their inclusion in species distribution modelling
remains a challenge.
Due to on-going climate change, physical disturbance
regimes are predicted to change rapidly in the future as
many disturbance processes have a signicant climate
forcing (IPCC 2007, Hjort and Luoto 2009). Although this
consequence of global climate change is recognized, there
is an urgent need for more comprehensive evaluation of
scenarios of future disturbance regimes because they may
strongly mediate the ecological consequences of climate
change (Walther 2004, Cannone et al. 2007). For example,
the response of the forb Euphrasia frigida to warming
will strongly depend on how changing climatic conditions
aect the distribution of cryoturbation (Nylhn and
Totland 1999). With declining frost-related disturbances
forecast for northern Fennoscandia (Fronzek et al. 2010),
it is likely that strongly competitive species (including
shrubs) may increase in abundance under the milder disturbance regime, with negative consequences for less competitive species (Pajunen et al. 2011). Thus community
composition may be simultaneously impacted by warming
temperatures and declining frost-disturbance frequency.
Moreover, from a conservation point of view, areas with
intense earth surface processes may be more important
than previously understood since traditional species distribution models have not explicitly accounted for geomorphological disturbances. It is, therefore, imperative that
spatial and temporal variation in disturbance processes is
incorporated into studies of global change. While frost- and
snow-related disturbances are particularly important at
high latitudes and altitudes, in temperate systems a greater
focus could be placed on disturbances driven by wind,
owing water and slope processes. For example, in coastal
and riverine systems species distributions may be especially
sensitive to wind and uvial disturbances which can strongly
alter habitat conditions and generate pronounced heterogeneity in environmental conditions over short distances
(Osterkamp and Hupp 2010).
806

In conclusion, this study demonstrates that the explicit

incorporation of geomorphological processes improves
species distribution models explanatory and predictive
power in a high latitude environment. Based on the relative
importance of geomorphological variables, these processes
are on average as important (if not more important) than
topography, climate and soil variables, highlighting the
signicance of disturbance measures in species distribution
models. Our results could thus be a step towards forecasting
more accurate and realistic species distributions and range
shifts.
Acknowledgements We acknowledge funding from the Academy
of Finland (project no. 1140873).

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