Earth-Science Reviews 126 (2013) 250265

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Earth-Science Reviews
journal homepage: www.elsevier.com/locate/earscirev

The role of trees in the geomorphic system of forested hillslopes

A review
ukasz Pawlik
Department of Geomorphology, Institute of Geography and Regional Development, University of Wrocaw, Poland

a r t i c l e

i n f o

Article history:
Received 10 January 2013
Accepted 12 August 2013
Available online 27 August 2013
Keywords:
Tree
Hillslope
Biomechanical weathering
Biochemical weathering
Tree uprooting
Biotransport

a b s t r a c t
Forested hillslopes form a special geoecosystem, an environment of geomorphic processes that depend strongly
on forest ecology, including the growth and decay of trees, changes in structure, disturbances and other uctuations. Hence, the following various functions of trees are reviewed here: their role in both biomechanical and biochemical weathering, as well as their importance for the hillslope geomorphic subsystem and for transport of soil
material via tree uprooting and root growth. Special attention is paid to tree uprooting, a process considered the
most efcient and most frequent biogeomorphological indicator of bio-physical activity within forest in complex
terrain. Trees have varied implications for soil formation in different environments (boreal to tropical forests)
and altitudes. In this paper an attempt has been made to emphasize how trees not only modulate geomorphic
processes, but also how they act as a direct or indirect agent of microrelief formation, the most striking example
of which being widespread and long-lasting pit-and-mound microtopography. Based on the analyzed literature it
seems that some problems attributed to forest ecology can have a fundamental effect on forested hillslope dynamics, a relationship which points to the need for its integration and interpretation within the eld of geomorphology. The biology of individual trees has a key inuence on the development of e.g. rock faces, weathering
front migration and changes in the soil biomantle within upper and lower forest belts. Additionally, forms and
sediments depend largely on the horizontal and vertical extent, volume and structure of root systems, as well
as on active processes taking place in the root zone and rhizosphere. Furthermore, although trees to a large extent
stabilize slope surfaces, their presence can also have a dual effect on slope stability due to tree uprooting, a process which in some circumstances can trigger mass movements (e.g. debris avalanches). So far, several attempts
at quantifying the inuence of trees on slopes have been made via the use of mathematical equations, enabling
researchers to calculate: 1) the root plate volume of uprooted trees, 2) the amount of soil displacement due to
tree root growth, and 3) rates of erosion, sedimentation and soil creep. In light of the reviewed literature, the
most urgent issue appears to be the need for a thorough study of the interactions and feedbacks occurring between trees and geomorphic systems (e.g. soil mixing and biotransport by trees) in different climate zones, altitudes and time frames, especially in terms of the development of forest ecosystems during the Holocene.
2013 Elsevier B.V. All rights reserved.

Contents
1.
2.

3.
4.

5.
6.

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . .
Trees and rock weathering . . . . . . . . . . . . . . . . . . .
2.1.
Biomechanical weathering . . . . . . . . . . . . . . . .
2.2.
(Bio)chemical weathering . . . . . . . . . . . . . . . .
Trees, soil mass displacement and soil processes . . . . . . . . .
3.1.
Tree uprooting . . . . . . . . . . . . . . . . . . . . .
Trees and supercial processes on hillslopes . . . . . . . . . . .
4.1.
Trees and accumulation processes
. . . . . . . . . . . .
4.2.
Trees and slope stability . . . . . . . . . . . . . . . . .
Trees and the development of landforms and surface rock structures
Geomorphic processes and trees attempts at quantication . . .
6.1.
Calculation of the root plate volume of uprooted trees . . .
6.2.
Calculation of soil displacement due to tree root growth
. .

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0012-8252/$ see front matter 2013 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.earscirev.2013.08.007

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 Pawlik / Earth-Science Reviews 126 (2013) 250265

6.3.
The damming effect of trees, erosion and exposed roots . . .
6.4.
Trees and palaeogeographical reconstructions . . . . . . . .
7.
Potential directions for future research conclusions and nal remarks
Acknowledgements
. . . . . . . . . . . . . . . . . . . . . . . . .
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1. Introduction
Since their rst expansion, trees have remodelled landscapes and
have been an important element of the Earth's natural history from at
least the Devonian (Meyer-Berthaud et al., 1999). Were early terrestrial
plants including trees able to create new niches and habitats? And, in
turn, did they help in the evolution of new plant and animal species?
This question remains to be answered satisfactorily (Kelly et al., 1998;
Gabet and Mudd, 2010). Others, while considering the evolution of
trees, seed plants and the colonization of land by forests during the
Devonian, have hypothesized that the development of terrestrial
vegetation induced a brief late Devonian glaciation by causing a drawdown in atmospheric pCO2 (the partial pressure of CO2) (Algeo and
Scheckler, 1998; Berner, 1998, as cited by Goudie and Viles, 2012). Forests currently cover at least 30% of the global land surface (FAO, 2005).
As a component of vegetation cover, they form a biological membrane
(also known as the Earth's biospheral envelope) that absorbs solar energy (a concept rst proposed by Vernadsky, 1926, 1944; see also reviews
by Ghilarov, 1995 and Lapo, 2001) and which also acts as a large reservoir of rainwater (Osuch, 1998; Phillips, 2009). Through their growth
and decay, trees have a critical and continuous effect on the land
surface of the Earth, and, as components of forest ecosystems, their
structural development and exchange of species and stands must
also be stressed. These natural phenomena occasionally happen in
a catastrophic manner, when slow changes are intensied and quickened (e.g. via windstorms).
Since at least the end of the 19th century, trees have been considered agents of soil disturbance and mixing (Shaler, 1891; Hack and
Goodlett, 1960; see reviews by Johnson, 1993; Wilkinson et al., 2009).
However, for a long time the forest environments of the upper and
lower montane belts were assumed to be static, or at least with infrequent changes (Jahn, 1989; but see for example Dietrich and Dunne,
1978), and were thus neglected in geomorphic studies (e.g. Klimek
and Latocha, 2007). Nevertheless, some exceptions exist. For instance,
the above-mentioned remark does not apply to humid tropical denudation systems where the study of the geomorphology of the humid
tropics cannot be divorced from a consideration of the vegetation
(Douglas, 1969, p. 13; also Thomas, 1994). Similarly, interactions between vegetation and hillslope geomorphology have recently been
emphasized as crucial to our understanding of linkages between
different ecoregions (Marston, 2010). In retrospect, biological inuences on geomorphological processes have gained much more attention (see, for example, Table 1 in Phillips, 2009). At present, the need
for such an interdisciplinary approach to geomorphology is undeniable,
with this approach's character and methods formalized in the subject of
biogeomorphology (Viles (Ed.), 1988a, 1988b, 1990; Corenblit et al.,
2008; Corenblit and Steiger, 2009). This sub-discipline aims at full integration of the biotic and abiotic aspects of geomorphic systems at all
levels of complexity. Forest geomorphology denes the functions of forests and of individual trees, and has already been appreciated as an important discipline within the frame of geomorphology, dealing with
many problems typical of mountainous areas (protective forests: FAO,
2005; Sakals et al., 2006). The most important issue currently facing
forest geomorphology appears to be the integration of the efforts of different disciplines towards producing an explanation of forested hillslope

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261
262
262
262
263

dynamics. This theory will be based on tendencies in forest ecosystem

changes and will thus involve integrating knowledge of forest ecology
(White, 1979). Similar efforts attempt to bridge the gap between pedology and forest ecology (amonil et al., 2010a), as well as between climate change and sediment transport (Constantine et al., 2012).
The main aim of this review is to explore, based on existing literature, the role of trees within the forested hillslope domains of mountain
geoecosystems, with the most important question arising being: How
do trees contribute to the activity and modication of geomorphic processes, and thus also to landscape evolution at different spatial and temporal scales?

2. Trees and rock weathering

Vegetation contributes to weathering via various mechanisms
over a full range of scales, from the microscale (e.g. the interaction
of ne roots with minerals) to the macroscale, where the physical
fragmentation of large rocks may take place in the root zone. Trees
contribute to weathering processes in many ways and their action
can be considered biotic weathering which, as proposed by Selby
(1993), is a combination of chemical and physical weathering effects
(Fig. 2). Biological weathering is, as proposed by Yatsu (1988), the

Fig. 1. Forest geomorphology as a subdiscipline within biogeomorphology. Only natural

processes are included, although Rosenfeld (2004) also considered the effects of forest
management activities within the frame of forest geomorphology. See text for explanation
(gure not yet published).

252

Pawlik / Earth-Science Reviews 126 (2013) 250265

Fig. 2. Traditional division of weathering types, distinguishing biological weathering and

its effects (original).

process brought about by living organisms and their decomposition

products (p. 287). Similarly, bioweathering covers the range of biological contributions to processes of rock and mineral weathering (Naylor
et al., 2002). Undoubtedly, although every plant inuences soil properties, as stated by Miles (1986), trees tend to have greater effects than
other plant life forms because of their size and longevity (p. 55). This
last statement refers to the hidden part of treestheir roots.
2.1. Biomechanical weathering
Trees bring about physical, chemical, biological and morphological
changes to the soils in which they grow (Retzer, 1963). A single tree is
a factor of biomechanical weathering inuencing soil and regolith properties (Phillips and Marion, 2006; Phillips et al., 2008a, 2008b). There
are several ways in which trees and their roots act as agents of biomechanical weathering, with the most important including (Lutz and
Griswold, 1939; Retzer, 1963; Schaetzl et al., 1989a; Gabet et al., 2003;
Phillips et al., 2005; Phillips and Marion, 2006; Gabet and Mudd, 2010):
1. tree uprooting;
2. physical anchoring of soil on slopes by tree roots (in Johnson, 1993,
considered a biomechanical process);
3. physical displacement of soil by root and trunk growth;
4. formation and inlling of stump rot depressions and burned-out
stump holes;
5. decay and inlling of former root channels;
6. penetration of bedrock by tree roots (Fig. 3);
7. swaying of trees during high winds which promotes root movement
and agitation of soil (causing changes in soil bulk density, porosity
and permeability).
Direct bedrock disruption is caused by the mining of mineral fragments from the root zone via tree uprooting (Small et al., 1990; Phillips
and Marion, 2006; Phillips et al., 2008b). The ability of trees to perform
such an activity depends on many factors, such as: 1) bedrock features,
2) characteristics of tree root system (e.g. rooting depth relative to regolith thickness), 3) edaphic conditions and 4) tree species. In many instances, regolith can be locally deepened through tree root growth
along bedding planes and penetration into rock ssures; these processes
quicken biomechanical weathering and facilitate both moisture migration and water availability. Consequently, chemical weathering can
penetrate deeper into solid rock, thus contributing to migration of the
weathering front. These tendencies draw attention to bedrock type, especially when the initial stage of slope cover development is considered
(Phillips et al., 2008a). Tree roots attack and penetrate the joints or
ssures of fractured sandstone, deeply weathered coarse-grained granite

and marble in different ways. Indeed, several studies have underlined

the fact that root system architecture, while genetically controlled, is
strongly inuenced by the characteristics of the substrate (Stone and
Kalisz, 1991; Matthes-Sears and Larson, 1995). As well as bedrock features, deep rooting can also be controlled by the presence of a fragipan,
excessive stoniness or a clay-rich B horizon (Schaetzl et al., 1989a).
Other important biomechanical effects of trees include: (1) physical
displacement of soil by root and trunk growth, (2) inlling of stump rot
depressions (Fig. 4) and (3) tree uprooting. The above mentioned effects have a considerable inuence on forest soils (Retzer, 1963;
Phillips and Marion, 2006) and can cause problems in terms of the
latter's classication (Retzer, 1963). Although all of these processes
are ubiquitous in forested ecosystems, only tree uprooting has received
much attention (amonil et al., 2010a). The uprooting of trees may contribute to continual mixing of the biomantle and has been recognized as
a key factor in soil evolution and forest dynamics (Lutz, 1940; Lyford
and MacLean, 1966; Faliski and Faliska, 1986; Scatena and Lugo,
1995; Phillips and Marion, 2006; amonil et al., 2009; Lenart et al.,
2010; amonil et al., 2010b). The physical displacement of soil by
expanding tree roots and trunks, although less spectacular, can bring
about considerable changes in soil mantle porosity and bulk density
which, for instance, during heavy downpours may affect slope stability.
However, in most cases roots reinforce hillslopes through root cohesion
and their anchoring into bedrock (e.g. Roering et al., 2003).
Tree roots can exert an axial and radial pressure as high as 1.45 and
0.91 MPa, respectively (Bennie, 1991, as cited by Gabet et al., 2003 and
Gabet and Mudd, 2010), potentially enough to push up a column of soil
approximately 100 m thick (Gabet et al., 2003). Birot (1966), as cited by
Pitty (1971), provided an example of rock fragment displacement and
suggested that a living root 10 cm in width and 1 m in length can
move a block weighing 40 tons. Such theoretical assumptions, when
added to information regarding potential rooting depth and root zone
extent, present a highly impressive view of root power, especially
with respect to the wedging and fracturing of rocks. In his excellent
book, Yatsu (1988) recalled a classic and, in his view, still unresolved,
problem associated with the wedging effect of growing plant roots
(p. 365). He stated, after analyzing the radial pressure (instead of the
axial pressure, which is frequently wrongly included in theoretical considerations) of the plant root and the tensile strength of a rock block,
that such an ability cannot be conrmed and that new data are urgently
required in order to solve the problem. This is probably still true in the
case of the fracturing of fresh (unweathered) rock by roots, with the
process continuing to suffer from a lack of attention and thorough explanation 25 years after Yatsu's statement. However, it is interesting
to note that in the case of rock wedging, such an ability had already
been ascribed to tree roots in early books on geomorphology. Indeed,
Baulig stated as early as 1940 that trees are able to crumble rocks and
comminute coarse particles (Baulig, 1958). Scheidegger (1970) later argued that plants penetrate rock ssures and cause their widening
(p. 122). However, none of the above-cited authors discussed the
issue thoroughly or referred to a dedicated article containing empirical
data. It might be possible that the only source of explanation comes
from Jackson and Sheldon (1949) and Matthes-Sears and Larson
(1995). Investigating a limestone cliff-like valley side in Derbyshire,
England, the rst authors concluded that woody plants caused recession
of the cliff face by detaching rocks from the edge of the cliff. However,
the authors doubted this was solely a consequence of ssures widening
as a result of root thickening and suggested that the action of drainage
water was also involved. Although very important, the results of this
study were based only on eld observations, not measurements of
root pressure or long-term monitoring. The second authors found that
the roots of an eastern white cedar (T. occidentalis) growing on a limestone cliff penetrated almost exclusively those rock ssures in the softer,
more weathered rock layers. Both articles effectively just continue the
discussion about rock wedging by tree roots and thus do not allow for
a denite rejection of Yatsu's statement. In this context, an interesting

 Pawlik / Earth-Science Reviews 126 (2013) 250265

piece of evidence was collected in underground archaeological monuments in Rome, Italy, by Caneva et al. (2009), who documented several
examples of damage caused by growing roots. Gabet et al. (2003) illustrated the mechanical disruption of bedrock (Fig. 6, p. 261), with Gabet
and Mudd (2010) later supporting their own biogeomorphic model of
soil production through the statement that a tree's roots can penetrate
bedrock and split it apart. It seems that such processes are indeed active
but only in conjunction with: 1) rock wedging (facilitated by chemical
weathering, as well as nutrient and water availability along ssures)
and 2) tree throw during which a part of fractured bedrock is extracted,
evidence of which can be easily observed in the root plates of fallen
trees. Direct mechanical fracturing of rock faces by tree roots is rather
unrealistic; root growth is such a slow process that it can be overlapped
by other physical effects such as frost wedging.
The above-described issue may have a greater meaning when the
spatial development and extent of tree roots are taken into account.
Although the reported average maximum depth of rooting is 3 m for
temperate deciduous forest and 4 m for temperate coniferous forest
(Gregory, 2006), it is estimated that merely 1 to 10% of a tree's total
root length occurs in soil below 1 m depth (Crow, 2005; see also
Schenk and Jackson, 2002). As a result, root wedging is frequently absent
and physical and chemical activity of roots is limited to the topsoil and
subsoil. Finally, whereas tree roots are able to penetrate rock ssures as
small as 100 m (Zwieniecki and Newton, 1995), individual trees may
create different patterns in soils at horizontal scales of 515 m (Binkley
and Giardina, 1998).
In conclusion, the pedological effects of trees are related to at least
three types of mechanisms of change: 1) biomechanical, 2) hydrological
and 3) mechanical (Phillips and Marion, 2005). According to various
authors, the pedological inuence of trees represents an important
control on local soil variability, especially in terms of changes due to
tree throw. A similar conclusion was drawn by amonil et al. (2011)
after studying soils within pit-and-mound microtopography in the
Novohradsk Mts., Czech Republic. Hydrological mechanisms of change
are beyond the scope of this review, but a synthesis can be found for instance in Ohte and Tokuchi (2011). The rst order mechanism in this
case is rainwater transmission through concentrated stem ow and inltration along main living roots (Selby, 1993), with tree root channels
(macropores) also facilitating water inltration and percolation (Gabet
et al., 2003).
2.2. (Bio)chemical weathering
Chemical weathering caused by biota is potentially much more effective than weathering under abiotic conditions (e.g. Berner, 1998).
As recently reviewed by Lucas (2001), plants directly control water dynamics, weathering and the chemistry of weathering solutions (Kelly
et al., 1998). Biota, from microbes through to vascular plants, also enhance chemical weathering through: 1) stabilizing soil, 2) producing
humic and other organic and inorganic acids and chelating agents

253

(Yatsu, 1988), 3) acting as a sink for the nutrients freed by weathering

and 4) contributing to physical weathering through the microfracture
of mineral grains (Schwartzman and Volk, 1989). Factor 1 points to
the ability of soil to act as a high surface area reservoir in which water
is stored and used as a medium for acid attack (Schwartzman and
Volk, 1989). Other factors which control rates of biotic weathering include: 1) biomass, 2) surface area of contact and 3) the capacity of organisms such as plants (roots) and mycorrhizal fungi to interact
physically and chemically with minerals (Taylor et al., 2009). The role
of vegetation has been considered, for instance, in silicate weathering
phenomena (Goudie and Viles, 2012), while lichens have gained special
attention in studies examining biological (biochemical) weathering
(Viles, 1995; Lee and Parsons, 1999; Chen et al., 2000; Lisci et al.,
2003; Goudie and Viles, 2012). Higher plants (vascular plants) may
have a more signicant impact on soil (see Cochran and Berner, 1996
and Berner, 1998) because they develop high potential root systems
that control soil properties (e.g. soil pH; Gruba, 2009). Berner (1998) argued that during the Devonian, higher plants were able to affect the
chemical weathering of silicate rocks because of deep rooting and
good drainage. This effect can be also attributed to trees, which have
been proven to enhance chemical weathering thanks to: 1) the extent
of their root systems, 2) biochemical processes in the rhizosphere in
which alteration of minerals takes place (Yatsu, 1988) and 3) the transmittance of water and moisture along root channels. The rhizosphere
has been dened as the volume of soil surrounding the roots which
is affected by it (Gregory, 2006; Calvaruso et al., 2009). Studies investigating Hawaiian basalts (USA) have shown that chemical alteration is much more advanced under higher-plant communities than
at unvegetated or lichen-encrusted sites (Cochran and Berner, 1996).
When compared to abiotic conditions, root activity is also likely to increase weathering rates of basaltic rocks by up to 5 times, as was proved
experimentally by Hinsinger et al. (2001). Other experimental data indicate that under articially planted red pines (Pinus resinosa Aiton),
weathering rates of primary minerals are more than 10 times higher
than those in a non-vegetated environment (Bormann et al., 1998).
However, it should be remembered that weathering rates determined
in the laboratory are generally orders of magnitude greater than eld
predictions (Kelly et al., 1998, p. 33). Nevertheless, tree roots play a
very important role because they contribute to both physical and biochemical weathering. The latter contributes to a greater extent within
the upper soil horizons and rhizosphere for two reasons. First, the
rapid turnover of ne roots affects all the soil material via the action
of rhizospheric processes. Second, variation in biological activity is
greater in the upper soil horizons (Lucas, 2001). For instance, Yatsu
(1988) recalled the results of experiments conducted by Spyridakis
et al. (1967) which showed that the rhizospheric activity of coniferous
and deciduous tree seedlings led to the transformation of biotite to
kaolinite (most effectively by the white cedar Thuja occidentalis) and
biotite to vermiculite (by the Monterey pine Pinus radiata). Moreover,
it is interesting to note that in the rhizosphere, invading plant roots

Fig. 3. Examples of the biomechanical inuence of trees on bedrock, rock faces and individual blocks (original).

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Pawlik / Earth-Science Reviews 126 (2013) 250265

Fig. 4. A conceptual model of the displacement of rock fragments and soil mass by root system growth. After complete decomposition of a stump, some rock fragments fall into the stump
hole and root channels are inlled. Figure inspiration drawn from Phillips and Marion (2006).

affect mineral grains both mechanically and chemically, with some part
of this work attributed to root-associated microorganisms such as fungi
and bacteria (Bonneville et al., 2009; Calvaruso et al., 2009 and references cited therein). Fungi and bacteria produce organic acids which
promote chemical weathering (Cochran and Berner, 1996); recent
data have revealed that the most important for biotic weathering is
the combined activity of roots and mycorrhizal fungi driven by carbon
uptake from photosynthetic organisms (Taylor et al., 2009). This set of
complex interactions can lead for instance to the formation of root
groves, which are common in karst areas (Wall and Wilford, 1966;
also root karst, see Viles, 1988a, 1988b) and are also known in sandstone regions (Dorn et al., 2013). Root groves form mainly as a result
of biochemical inuences, although biophysical processes have also
been mentioned (e.g. enlargement of microfractures; Viles (Ed.),
1988a, 1988b; Dorn et al., 2013).
Gabet et al. (2003) argued that, through breaking bedrock, tree roots
can cause a six-fold increase in the surface area available for chemical
weathering (by creating spaces for water and different weathering factors; Gabet and Mudd, 2010). Although this view (see Fig. 6, p. 261,
Gabet et al., 2003) is really only a simplication made for modelling purposes, to support it the authors also referred to the work of Lutz (1960)
who noted many instances of fresh rock torn out of bedrock (p. 261).
For comparison, Schaetzl and Anderson (2005) state that as rocks are fractured due to physical weathering factors other than tree roots, their
surface area increases geometrically. Tree root exploration is also considered a primary mechanism of the downward extension of the ecosystem
boundary (Bormann et al., 1998). Important data at the watershed scale

were published by Moulton et al. (2000), who calculated a 4 times greater

weathering release of Mg2+ from forested areas and proved pyroxene
weathering to be 9 to 10 times higher there than in barren but geologically similar areas. Additionally, when considering individual forest communities, different tree species have varying effects on both weathering and
soil formation, e.g. podsolization. For instance, a spruce ecosystem was
found to cause a threefold increase in calcium denudation from monzonite granite in comparison to a beech coppice on the same parent material
(Bormann et al., 1998). Conifers, especially spruces: 1) accelerate podsolization, 2) cause surface acidication and 3) cause higher rates of organic
matter accumulation. In contrast, broadleaved species are associated
with: 1) different soil fauna which may decrease podsolization via physical intermixing of the A and B horizons, 2) a reduction in soil acidity and
3) mull-humus formation (Miles, 1986). The rst of these processes can
refer to the soil homogenization sequence proposed by Hole (1961). In
their analysis of chemical weathering and denudation in a forest ecosystem based on a 20-year sandbox experimental study, Balogh-Brunstad
et al. (2008) concluded that disturbance can be an important factor controlling chemical denudation rates (in this case a rapid increase in denudation rates was observed after tree harvesting).
Another important issue is linked to palaeogeographical reconstructions and chemical weathering. Chemical weathering is thought to have
been a dominant process under forest cover in the Holocene (Starkel,
1977; Bieroski et al., 1992), in contrast to the dynamic mechanical
denudation which took place during cold phases in the Pleistocene
(e.g. Migo, 2006). The postglacial succession of forest led to the stabilization of slope mantles which then became a potential source of

 Pawlik / Earth-Science Reviews 126 (2013) 250265

255

Fig. 5. Sequence of pit and mound pair development due to tree uprooting caused by strong winds. Note: years taken from amonil et al. (2009). Author's original work.

weathering material. Slope mantle volume in the Sudety Mts., SW

Poland, has been calculated to be at least 1,000 000 m3 km-2 (Klimek
and Latocha, 2007).
3. Trees, soil mass displacement and soil processes
3.1. Tree uprooting
Although the signicance of the tree uprooting process has been
underlined in several research articles (Phillips et al., 2008b; amonil

Fig. 6. Boulder trapped in the root system of a fallen tree in an advanced stage of decomposition. An example found in the lower belt forest of the Karkonosze Mts., Sudety,
SW Poland.

et al., 2010a), review publications (Schaetzl et al., 1989a, 1989b, 1990;

Viles, 1990; Johnson, 1993; Gabet et al., 2003; Marston, 2010; amonil
et al., 2010b) and scientic books (Huggett, 1995; Schaetzl and
Anderson, 2005), an appreciation of its importance is not equally shared
among the different Earth Science disciplines. Nevertheless, the process
has been thoroughly studied from the point of view of forest ecology
(Putz, 1983; Jonsson and Dynesius, 1993; Everham and Brokaw, 1996;
Ulanova, 2000; amonil et al., 2009; ebkov et al., 2012), forestry
(Steven, 1953; Vicena et al., 1979; Brzdil, 1998; Brzdil et al., 2004)
and soil science (in terms of soil production: Gabet and Mudd, 2010;
amonil et al., 2010a, 2010b; soil chemistry: Liechty et al., 1997;
amonil et al., 2008a, 2008b; and soil disturbance and pit-and-mound
microrelief: Lutz and Griswold, 1939; Lutz, 1940; Denny and Goodlett,
1956; Lyford and MacLean, 1966; Schaetzl, 1986; Lenart et al., 2010;
amonil et al., 2010a). For the most recent and comprehensive review
considering tree uprooting vs. soil interactions, see amonil et al.
(2010a). Although a number of papers focus on the geomorphic consequences of the tree uprooting process (Lutz, 1960; Kotarba, 1970;
Schaetzl and Follmer, 1990; Norman et al., 1995; Embleton-Hamann,
2004; Osterkamp et al., 2006; Phillips et al., 2008b; Gallaway et al.,
2009), many forest environments have not yet been fully studied and
thus further work is required for a more complete evaluation of forested
hillslope dynamics.
Tree uprooting (also tree throw, tree tip, tree saltation; Faliski and
Faliska, 1986; similar in meaning is root throw, but see the explanation
in Osterkamp et al., 2006, p. 7) (Fig. 5) is not only one of the most important active biogeomorphological processes taking place on forested
hillslopes, it is also considered an important link between atmosphere,
biosphere and pedosphere (Small et al., 1990), undoubtedly contributing, at least to some extent, to the Critical Zone paradigm (Anderson
et al., 2008; Lin, 2010). Its efciency in breaking ground (Wilkinson
et al., 2009) has been widely evaluated and exists under several

 Pawlik / Earth-Science Reviews 126 (2013) 250265

256
Table 1
Reported mean root plate volumes.
Mean root plate
volume (in m3)

Place

Tree species or forest type

Altitude
(m a.s.l.)

1.83.6

Tatra Mts., Poland

lower and upper belt spruce forest

10001500 Kotarba, 1970

1.22.8

New York, USA

different species in second-growth

forests

Beatty and Stone, 1986

4.0

Washington, USA

western hemlock (Tsuga heterophylla)

and Pacic silver r (Abies amabilis)

701020

Reid, 1981

2.0

Arkansas, Ouachita
Mts., USA
Tatra Mts., Slovakia

75530

Phillips et al., 2008a, 2008b

9231284

Dbrowska, 2009

mixed shortleaf pine (Pinus achinata)

and hardwood forest
1.9
mixed spruce-larch forest
(Lariceto Piccetum)
0.10.7
British Columbia, Canada
lodgepole pine (Pinus contorta Loudon
var. latifolia Engelm.), Engelmann spruce
(Picea engelmannii Parry ex. Engelm.)
1.7
Tatra Mts., Slovakia
mixed spruce-larch forest (Lariceto Piccetum)
0.3
Puerto Rico
dry, moist, wet and rain forest with
broadleaf, needleleaf and palm trees
4.0
Sudety Mts., Poland
lower belt spruce forest
4.0 (beech)5.6 (spruce) Outer Western Carpathians, natural r-beech forest
Czech Republic

Reference

13303086 Gallaway et al., 2009

8001300
01000

Rojan, 2010
Lenart et al., 2010

700900
600800

Pawlik, 2013
amonil P. (unpublished
data)

Notes
According to
Norman et al. (1995) calculated
volumes are twice too large
Root plate volumes were
calculated as cylinders
(V = r2h)
Calculation made with a model
of cylinder with a segment
missing

Survey was carried out in

post-re conditions

For the calculation a formula

for half an ellipsoid
were used as proposed by
Norman et al. (1995)

Mean values obtained from different plots.

terms, e.g. biomechanical weathering, bioturbation, pedoturbation,

oralturbation or oralpedoturbation (Pawluk and Dudas, 1982;
Johnson, 1993; Gabet et al., 2003; Phillips and Marion, 2005;
Schaetzl and Anderson, 2005), as well as biogenic transport
(Swanson et al., 1982a; Burns and Tonkin, 1987; Schaetzl et al., 1990),
which has been dened as the movement of debris as a result of biological activity (Swanson et al., 1982a). Tree uprooting is also known as biological saltation (Birot, 1966) and bioporting transport by biota (see
Phillips and Marion, 2005) and can be divided into two categories:
1) catastrophic local transport (caused by e.g. tree throw, animal
burrowing) and 2) slow transport (e.g. root growth, wind stress on
trees) (Swanson et al., 1982a). The average volume of soil mass
transported in root plate form is between 0.1 and 5.6 m3 (Table 1).
Tree uprooting is a major disturbance factor in most natural forests
(Schaetzl et al., 1989a; Huggett, 1995; Everham and Brokaw, 1996;
Brzdil et al., 2004) and is considered to be an important variable in so
called disturbance ecology and gap-phase dynamics (White, 1979;
Pickett and White (Eds.), 1985; Ulanova, 2000; Linke et al., 2007).
Simultaneously, it is also an agent altering hillslope stability and affecting other geomorphological processes, changing both their course and
intensity (see reviews by Schaetzl et al., 1990, p. 285, and Viles, 1990,
p.12). For instance, after severe windthrow in the Swiss Alps caused
by extra-tropical cyclone Vivian in 1990, countless erosion scars were
formed (Gerber et al., 2002). Additionally, due to heavy rainfall in subsequent years, the combination of construction works carried out after
the storm and the lack of vegetation led to further scar development.
The authors concluded that the areas affected by windthrow were highly susceptible to erosion and shallow landslides, a reciprocal effect also
considered in previous work examining hillslope stability and evolution
(e.g. Schaetzl et al., 1990; Selby, 1993; Johnson and Wilcock, 2002).
The uprooting of trees contributes to the downslope movement of
soil (seasonal biogenic creep; Kirkby, 2004; Wilkinson et al., 2009)
and rock mantle, and locally may initiate gully erosion (Lutz, 1960;
Wilkinson et al., 2009). Published results, which range from 0.02 to
1.3 t ha1 year1, demonstrate that tree uprooting should be seen as
an important factor inuencing sediment transport in a watershed
(Schaetzl et al., 1990). As shown by Swanson et al. (1982a), sediment
transfer driven by tree uprooting can be close in magnitude to soil
creep rates (see also Clment, 1993). Interactions between the tree

uprooting process and soil creep have been studied in the Stoowe
range of the Sudety Mts., SW Poland (Pawlik et al., 2013). Probably
less frequent but more important is the transport of large boulders. Published data indicate that a wind-uprooted tree may move rocks with a
volume of as much as 1.4 m3 and a weight of up to 4 tons (Lutz,
1960). Additional results show that uprooting may contribute to sediment transfer via several other mechanisms (Schaetzl et al., 1990):
1. Tree-throw pits absorb water which leads to a reduction in soil shear
strength and, as a consequence, can result in debris avalanches or debris ows;
2. During tree fall a sort of vibration is induced which can cause soil materials to reach their liquid limit and fail, triggering mass movement.
Tree uprooting is a very common forest process (e.g. Stephens, 1956)
that signicantly inuences the structure and layering of slope covers
(e.g. Phillips and Lorz, 2008), and especially applies to mountain areas
for the following reasons. Firstly, wind, which is the primary factor of
tree damage (Everham and Brokaw, 1996; Brzdil, 1998; Peterson,
2007; Phillips et al., 2008b), speeds up with altitude (see review by
Mitchell, 2012), while another important factor of tree damage is the
occurrence of ice storms (see, for instance, Bragg et al., 2003). The proportional area covered by pit-mounds after a large catastrophic blowdown may be up to 11% (Peterson et al., 1990), suggesting that a
substantial part of a hillslope may be affected by the tree uprooting process. Secondly, due to the high slope gradients, up to 50% of soil material
from the exposed root plate falls outside the pit (Burns and Tonkin,
1987; Gallaway et al., 2009), partly because trees tend to fall downhill
on steeper slopes (Burns and Tonkin, 1987; Norman et al., 1995); this
phenomenon has been assessed at 90% probability for a 45 slope
(Gabet and Mudd, 2010). Thirdly, soil attached to the root system is subject to further deterioration due to subsequent supercial processes
such as rain splash, wash or mass wasting (Small et al., 1990; Pawlik,
2013; Pawlik et al., 2013), thus prolonging its downslope transfer. The
material removed from root balls covers an area lying directly below
and affects, for instance, undergrowth vegetation. However, it has to
be kept in mind that rock fragments of different sizes can be trapped
at some height above the ground within the root system, and remain
there for many years after the toppling of the tree (Fig. 6).

 Pawlik / Earth-Science Reviews 126 (2013) 250265

Table 2
Tree uprooting and its inuence on soils, pedogenesis and transport of sediments (based
on Phillips and Marion, 2005; Phillips et al., 2008b).
Effects
Direct

Indirect

Soil mixing
Soil prole inversion
Local redistribution of soil
substance
Formation of pit-and-mound
microtopography (windthrow
morphology)

Action of erosion and mass wasting processes

on exposed root plate. Microscale variations
within windthrow morphology in terms of:
- Weathering intensity,
- Moisture ux and water availability,
- Dynamics of organic matter
(preferable litter accumulation in pits),
- Chemistry,
- Microclimate.
Intensied organic activity in pits
(e.g. by earthworms)

An important parameter which allows for better evaluation of tree

uprooting frequency is the ratio of uprootings to broken trees, both as
a consequence of long-term variation and a single strong wind event
such as an extra-tropical cyclone. Yamamoto (2000) discovered
uprooting to be the least recorded cause of tree mortality in Japan, a
nding which is consistent with studies from Europe, including the following: 1) In the old-growth r-beech forests of the Czech Republic, the
ratio between bole breakages and uprootings was 23:1 (amonil et al.,
2013), 2) in the beech-r old-growth forests of Bosnia and Herzegovina,
gures of 14% uprooted trees, 60% snapped stems and 26% standing
dead trees were recorded (Bottero et al., 2011). Both given examples
come from mountain forests and document a phenomenon associated
with gap phase dynamics. The proportion of uprootings is distinctly different after catastrophic bora and foehn wind events. In the High Tatra
Mts. of Slovakia after a bora wind event in November 2004, a ratio of
41% windsnaps to 59% uprootings was recorded (Rojan, 2010), while
similar gures were observed on the northern side of the Tatra Mts.
after a strong foehn event in 1966 (48% windsnaps and 52% uprootings)
(Bzowski and Dziewolski, 1973). These latter results likely reect the
presence of tree species such as the generally shallow-rooted Norway
spruce which dominate in the Tatra Mts. massif. Peterson (2007)
found a good relationship between vulnerability to uprooting and tree
species/features (mainly tree diameter, wood properties) in several
natural forests in North America. Generally, coniferous trees are
more susceptible to windthrow, with tree resistance to uprooting
negatively correlated with their diameter at breast height. For data
regarding other inuencing factors such as topography or forest stand
structure, see also reviews by Everham and Brokaw (1996) and
Mitchell (2012).
After tree uprooting, soil horizons are extensively disturbed by: (1)
the overturning of the tree and soil mass upheaval above the ground
in root plate form, (2) the mixing or partial inversion of horizons during
mound formation (Schaetzl, 1986) and (3) the subsequent erosion of
the tree-throw mound and concentration of rock fragments within the
topsoil and/or the surface (e.g. so called gravel armours; Small et al.,
1990; Phillips et al., 2005). Rock fragment veneers derived from coarse
rock fragments falling forwards from the root plate beyond the pit
area are of similar origin (Osterkamp et al., 2006). An equally important
modication made to soil mantles caused by tree throw is the uneven
redistribution of soil thickness across a hillslope (Gabet and Mudd,
2010) in the form of pit-and-mound microtopography, which can persist for more than 2000 years (Schaetzl and Follmer, 1990), although
more frequently for only 100200 years (amonil et al., 2009;
ebkov et al., 2012) and up to 300 years (Denny and Goodlett, 1956,
see review by amonil et al., 2010a). So far, the highest documented
age based on 14C dating was recorded in Michigan, USA, and exceeds
6000 years (amonil et al., 2013). The reported variation in pit-mound
longevity depends on climatic conditions and site characteristics; on
well-drained sandy Podzols such topography can persist for more than

257

a thousand years (amonil et al., 2010a). All the potential effects of

tree uprooting on soils are summarized in Table 2.
A nal issue to consider is the contribution of tree uprooting and
sediment transport as a self-reinforcing mechanism. Compared to pits
and undisturbed sites, tree-throw mounds are more frequently and easily occupied by the next generation of trees and if uprooted lead to further downslope transport of soil material from the mounds (Lyford and
MacLean, 1966; Kabrick et al., 1997). The same factor, i.e. tree growth on
mounds, causes their conservation (stabilization), whereas the adjacent
pits are lled much more rapidly.
4. Trees and supercial processes on hillslopes
4.1. Trees and accumulation processes
Trees, either alive or dead standing (snags) or lying (logs) can
prevent and delay the downslope movement of mineral and organic
matter via the physical obstruction of particle transport on hillsides
(Fig. 7). In a similar manner they also affect surface runoff (Harmon
et al., 1986; Maser et al., 1988), protect hillslope surfaces against erosion
(Raka and Orulk, 2009) and contribute to soil stabilization by controlling the ow of water, soil and litter across the forest oor
(Stevens, 1997). Such functions, according to biogeomorphic nomenclature, can be considered bioprotective (Naylor, 2005). An important variable accelerating soil material accumulation within a hillslope is the
number of snags and logs per unit area. The amount of logs accumulated
on the forest oor is a function of tree uprooting and tree snapping intensity (Harmon et al., 1986; Maser et al., 1988; Stevens, 1997) modulated in most cases by strong wind events (Everham and Brokaw,
1996; Schelhaas et al., 2003) and is thus positively correlated with
the amount of trapped sediment, assuming that the tree trunks preferentially fall and/or lie perpendicularly to the slope prole, i.e. along
the contour. However, according to this author's knowledge, the latter
relationship has not yet been veried, and indeed virtually not studied.
Soil particles in transport accumulate on the upslope sides of trees due
to various processes such as sheet wash, soil creep, debris ow and shallow landsliding. Although the amount of sediment can be calculated
(Measeles, 1994), interpretation is probably restricted to the cumulative
effect of all potentially active processes within the forested hillslope.
Coarse woody debris which lies parallel to contour lines plays a similar
function; however, because there is no root anchoring in bedrock and/
or regolith, soil creep cannot be measured in this case. Another important source of weathering material deposited on the upslope side of
trees, snags and logs is the contemporaneous development of frostriven cliffs enhanced by vegetation (tree roots causing the disintegration of frost-riven cliffs) (Raka, 2007). Recently, the geomorphological
role of trees has been analyzed using the concepts of sedimentation
traps (Matyja, 2007) and dam-like effects (Raka, 2007; Raka and
Orulk, 2009); however, detailed description of the damming effect of
trees in fact dates back to 1968 (LaMarche, 1968).
In general and conceptual models, both upper and lower forest belts
have traditionally been considered as accumulation zones where sediments from upper mountain zones (subalpine and alpine) are trapped
and stored. Such a conclusion was reached for the Karkonosze range
of the Sudety Mts., SW Poland, after a study of mechanical denudation
in the region (Bieroski et al., 1992). However, the authors did not consider the biotic components of hillslope environments; for instance, the
tree uprooting process (as a source of mechanical denudation and biogenic transport) was not included in their analysis. In contrast,
Swanson et al. (1982a), after referencing published rates of transport
due to tree throw (between 1.5 and 2.0 mm year1), concluded that
the latter are comparable in magnitude to soil creep measurements on
forested hillslopes and should thus be taken into account in sediment
budgets (p. 8).
Accumulation processes were recognized in the Tatra Mts., Western
Carpathians, where they were described as biological aggradation by

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258

Fig. 7. Examples of the conguration between trees, relief and surcial processes causing transport of rock particles (gure not yet published).

Jahn (1979), who discovered a change in the type of sedimentation in

front of a dwarf pine clump from debris to sand and supposed it to be
caused by the regeneration of plant cover on the upslope side. Another
case study focused on a 2002 debris ow that occurred in the Babia Gra
Massif, Beskidy Mts., S. Poland. This study revealed that trees (standing
or decaying) and the coarse woody debris of dead tree trunks lying on
the hillslope inuenced transport and depositional processes. These
natural elements of the slope surface not only caught part of the debris
ow material, reducing its energy and range (Matyja, 2007), the coarse
woody debris also formed a step-like surface on the slope, increasing
surface roughness and again affecting ow. Assessing the time necessary for complete decomposition of a tree trunk, such an effect could
last for up to 100150 years (Matyja, 2007) and potentially even more
than 400 years (the time of decomposition of the Douglas Fir; Maser
et al., 1988).
The above examples permit the formulation of the general conclusion that trees temporarily modify transport and sediment ux on
hillslopes, simultaneously affecting their long-term transfer between
hillslope and river valley geomorphic systems. Furthermore, trees are
also associated with the direct addition of mass to river channel systems
via coarse woody debris, litter and organic matter of various types (Reid,
1981; Marston, 1982; Swanson et al., 1982b). For instance, coarse
woody debris can control erosion-sedimentation processes and thus
changes in river channel morphology.
4.2. Trees and slope stability
It is widely accepted and proven that vegetation, especially forest, can
stabilize steep slopes (Ziemer, 1981; Marden, 2004; Rickli and Graf,
2009). This issue has been comprehensively reviewed by O'Loughlin
(2005), who pointed to: 1) the modication of soil moisture and porewater pressure by trees, 2) the formation of a protective permeable organic forest oor layer and an increase in soil conductivity, enhancing
soil drainage (e.g. via the rapid transmittance of water through decayed
tree roots) and 3) the mechanical reinforcement of soil by tree roots. The
latter function, which is enabled through root anchoring in bedrock,
crossing zones of weakness in the soil and providing long brous binders
within a weak soil mass (Ziemer, 1981; Roering et al., 2003), is recognized as the most important benecial effect of vegetation on slope stability (Kuriakose and Beek, 2011). Such benecial effects of trees are
mostly recognized in shallow soils (Montgomery et al., 2000; Gabet
and Dunne, 2002).
Rickli and Graf (2009) documented more than 500 shallow landslides (b2 m deep) after heavy rainfall events in Switzerland, concluding that shallow landslides were less frequent in forest than in open
land. However, when forest condition had decreased, for instance due
to a past disturbance event (e.g. windthrow, fungi disease or bark beetle
outbreak), a markedly higher intensity of landslides was observed. Although especially visible on steeper slopes, these landslides were
caused mainly by a reduction in soil cohesion due to loss of root strength

(Selby, 1993; Johnson and Wilcock, 2002; Rickli and Graf, 2009). Clearcutting has a similar effect on root strength and slope stability, after
which landslide frequency also increases (e.g. Montgomery et al., 2000).
For instance, it has been hypothesized that 90% of root reinforcement
is lost within 9 years after logging (Ziemer, 1981) (Fig. 8). Notwithstanding this, in some natural environments (i.e. not anthropogenically
altered) trees can lead to the destabilization of slopes while the latter are
oversaturated with rainwater. Such conditions frequently appear in the
tropics where trees and dense root mats enhance inltration and cause
an increase in pore-water pressures (Thomas, 1994).
Another stabilizing phenomenon is connected with tree uprooting,
which leads to surface or near-surface rock fragment layering. Layers
such as gravel armours or rock fragment veneers limit erosion and thus
stabilize hillslopes (Schaetzl and Follmer, 1990; Osterkamp et al.,
2006). Similarly, coarse woody debris inhibits downslope sediment
transport and erosion below tree trunks. However, some authors have
documented a very interesting dualistic effect of forests, through which
the tree uprooting process also contributes substantially to hillslope sediment ux (e.g. Hughes et al., 2009; Constantine et al., 2012). Another
two-fold effect is known to promote the occurrence of mass wasting
due to tree uprooting in short-term intervals, but in some cases this
can be hindered by the widespread and continual thinning of soils by
downslope transport in the long-term (e.g. Gabet and Mudd, 2010).
One of the rst reports documenting windthrow-induced debris avalanches (caused by a rapid decrease in the positive effect of root
strength) came from Alaska (Swanston, 1967). Such results contradict
the common opinion that forest cover stabilizes hillslopes and inhibits
erosion.
5. Trees and the development of landforms and surface
rock structures
Although Dietrich and Perron (2006) were unable to dene a
topographic signature of vegetation or fauna on the Earth's surface,
Gabet and Mudd (2010) have pointed out that pit-and-mound
microtopography is one of the most spectacular forms resulting from biological activity and is clear evidence of the tree uprooting process. As
the size of individual forms (Table 3) and the density of pit-mound
pairs vary between sites, it is difcult to draw any rm conclusion regarding the general validity of this theory. However, such microrelief
seems to be more pronounced and to last for longer on steeper slopes
(Schaetzl and Follmer, 1990; Norman et al., 1995) and in temperate forests than in tropical forests (Putz, 1983). The density of pit-and-mound
microtopography ranges between 50 (Cremeans and Kalisz, 1988) and
1200 (Lyford and MacLean, 1966) pit-mound pairs per hectare, with
the features potentially persisting for up to 2500 years (Schaetzl and
Follmer, 1990) or even more than 6000 years (amonil et al., 2013).
Pit-and-mound topography has been found on mesoforms of various
morphogenesis, including glacial features such as drumlins (e.g. Kabrick
et al., 1997) and kames (Norman et al., 1995). The uprooting of trees,

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259

Fig. 8. A conceptual model of forested hillslope stability before and after a disturbance event. Source: after Selby (1993), Ziemer (1981) and Sidle (2008), modied.

subsequent deterioration of root plates and levelling of tree-throw

mounds can lead to the formation of structures and forms traditionally
attributed to other geomorphic activities, e.g. frost processes (see examples given e.g. by Embleton-Hamann, 2004). For instance, the erosion
and levelling of mounds have been found to leave rock fragment structures similar to patterned ground (Denny and Goodlett, 1968). Whereas
this is a direct imprint of mound attening by surcial processes, pitand-mound microtopography itself was in the past incorrectly
interpreted as the result of contemporaneous congeliuction
(Dylikowa, 1956) or frost processes of Pleistocene or Holocene age
(e.g. Denny and Goodlett, 1956; Embleton-Hamann, 2004). Structures
similar to patterned ground were documented by Phillips and Marion
(2006), Fig. 8, p. 245) and attributed to tree root expansion, rock fragment displacement and stump hole inlling. Some authors have suggested that whereas the development of gravel armour reects the
complete erosion and elimination of any topographic expression of
the treethrow mound (Small et al., 1990), rock fragment veneers can
develop via root throw on immature soils (Osterkamp et al., 2006). Finally, although tree uprooting in the short-term disrupts layering and
horizonation, on a longer time scale it produces distinct surface or
near-surface layering due to continual mining of rock fragments from
the subsoil (Phillips and Lorz, 2008).

A number of authors have suggested that trees can contribute to the

development of larger landforms. For instance, it has been hypothesized
that due to the biomechanical and biochemical action of their roots,
trees can inuence tor formation in karst regions. Gams (1966) proposed a two-stage formation process for tors in the Dinaric and Alpine
Karst of Slovenia, comprised of 1) deep subsurface weathering followed
by 2) denudation of weathered material and excavation of tors. This hypothesis has probably never been thoroughly studied, nor has that of
weathering pit formation (known as oriangas in Portuguese) under
trees, as proposed by Freise (1938). Moreover, trees, through their
root systems, disintegrate rock fragments and widen ssures in rock
faces and bedrock (see Fig. 3). As Jackson and Sheldon (1949) observed:
There are good reasons for believing that these woody plants, (), play an
important part in detaching rocks from the edge of the cliff, so causing
recession of the cliff face. The authors also pointed out that the action
of the trees was quickened by the properties of the rock cliff in question
(soft limestone with numerous joints and bedding planes). However, as
with tor and weathering pit formation, this process has not been intensively studied (despite the fact that it is frequently observed). This general remark may also be applied to all potentially active biomechanical
processes in mountain environments (see also Yatsu, 1988). In trying to
explain such neglect, one could point to: 1) problems regarding the

Table 3
Reported mean treethrow mound and/or pit volumes.
Mean mound volume (m3)

Mean pit volume (m3)

Place

Tree species or forest type

Altitude (m a.s.l.)

Reference

0.7

Colorado, USA

2900

Osterkamp et al., 2006

0.6
0.2

0.2

Wisconsin, USA
New York, USA

Kabrick et al., 1997

Denny and Goodlett, 1968

2.23.0

1.7
2.9

3 1.3
1.31.9
0.8
1.6
2.2

USA
N Iran
Blue Mountains, Australia
Stoowe Mts., Poland
Outer Western Carpathians,
Czech Republic

Pine (Pinus ponderosa and P. contorta),

Douglas r
(Pseudotsuga menziesii) and aspen
(Populus tremuloides)
Sugar maple-basswood forest
The area was forested in presettlement
times only.
Douglas-r
Temperate forest of Mazandaran Province
Eucalypt
Fertile mountain beech forest
Natural r-beech forest

1001700

600760
600800

Gabet and Mudd, 2010

Kooch et al., 2012
Richards et al., 2011
Pawlik et al., 2013
amonil P., unpublished data

Values calculated using data from the cited reference. For this purpose the equation from Norman et al. (1995) were used.

 Pawlik / Earth-Science Reviews 126 (2013) 250265

260

availability of tools allowing for quantication of process magnitude e.g.

for each type of rock and tree species and 2) measurement difculties.
In terms of the latter, it may be difcult, for instance, to distinguish the effects of tree root action on rock ssures from co-existing frost weathering.
6. Geomorphic processes and trees attempts at quantication
The main aim of many analyses is not only the qualitative description of processes, but also an attempt at their quantitative expression.
As trees are involved in many different hillslope processes, various
ways (both measurement methods and physical models) of quantifying
these processes have been developed. The most important of these
methods include:
1. Calculation of the root plate volume of uprooted trees;
2. Calculation of soil displacement due to tree root growth;
3. The damming effect of trees, erosion and exposure of roots.
6.1. Calculation of the root plate volume of uprooted trees
So far, relatively few researchers have tried to measure and calculate
rates of transport caused by tree uprooting, i.e. the amount of soil material upheaved along with the root system of a fallen tree onto the ground
surface (e.g. Kotarba, 1970). However, such information plays a crucial
role in any attempt at the measurement of watershed sediment balance.
Recently, established methods developed for measuring root plate volume were reviewed by Richards et al. (2011). The most frequent method of calculation was based on the formula for half of an ellipsoid in
which the axes are not equal, which is a close approximation of the
shape of a root plate (Dbrowska, 2009; Rojan, 2010; Pawlik, 2013).
This equation can be written as follows:

so-called partitioning method. Along a baseline, the pit is hypothetically partitioned into rectangular prisms (the dimensions of which
are noted), with the latter's volumes then calculated and summed
(p. 1244). However, this method cannot be applied if: 1) there is
no typical pit but only a shallow uncovered surface, 2) the root
plate is partly or entirely placed within a pit (e.g. complex treefall
with backward displacement; Schaetzl et al., 1989a) and 3) treefall
was incomplete (Beatty and Stone, 1986).
Others have proposed a method of root plate volume estimation
from DBH (diameter at breast height), based on a linear regression
model (Burns and Tonkin, 1987). The authors found statistically significant relationships between DBH, root plate volume and the dimensions
of the resulting pits and mounds.
In a similar manner, as shown in Figs. 8 and 9, the volume of
upheaved soil material can be approximated by measuring pit and
mound dimensions, with the latter's volume then calculated using the
equation for half of an ellipsoid Eq. (2) (Norman et al., 1995; Kabrick
et al., 1997) (Fig. 11) or quarter of an ellipsoid (not shown here; Putz,
1983).
This method can be simplied if we assume that width and height
are equal (Gabet and Mudd, 2010), then:
V

2 2
r d
3

where w is width, h is height and t is depth of the root plate (Fig. 10).
Richards et al. (2011) also proposed a new method with which to
estimate the displaced soil volume by measuring pit volume, using the

where r and d are pit radius and depth, respectively.

Although pits are normally lled to some extent with organic and
mineral matter, and mounds already partly levelled, this method
seems to be much more precise in estimating upheaved soil volume
than does the direct measurement of soil material in root plates, because it does not take into account the volume of root wood (normally
already decomposed). Although this part of a root plate is commonly
not excluded because of difculties associated with its volume assessment, calculation attempts have been made. According to some authors,
the root volume can be derived from tree diameter measured at breast
height (DBH) or stump height (Santantonio et al., 1977; Thies and
Cunningham, 1994), while others have attempted to calculate the volume of tree wood that penetrates into the root plate or continues
through it as a taproot. This latter parameter was determined based
on the cylinder volume equation, i.e. Vw = r2p, where r is the radius
of the tree trunk and p is the penetration depth of the trunk into the
soil-root plate (Richards et al., 2011). The authors also calculated the
volume of both large (N1.5 cm in diameter) and ne roots; together
with Vw these three values were then subtracted from the pit volume
calculated using the partitioning method described above.
The above information is essential in order to use the model of sediment transport via tree uprooting proposed by Gabet et al. (2003),
which was later developed into a bio-geomorphic model of soil production by Gabet and Mudd (2010) and incorporated into a biologically-

Fig. 9. Main dimensions of a root plate used for volume calculation. Author's original work.

Fig. 10. Denition of root plate dimensions proposed by Reid (1981, p.153).


1
4 w h


 d
2
3
2 2

whd
6

where w is width, h is height and d is depth of the root plate (Fig. 9).
Other methods have also been used, such as the equation for the volume of a cylinder (e.g. Beatty and Stone, 1986) or for a cylinder with a
missing segment (Reid, 1981) (Fig. 10).
In this case the equation is written as shown below:
8
0
1
2
39
>
>
=
< w2
 
 q
 2

whC w
1 B
6 w 2
7
wh 
w hw wh2 5
4
cos @1 w A
>
2
2
;
: 2
2

V t>

 Pawlik / Earth-Science Reviews 126 (2013) 250265

261

6.3. The damming effect of trees, erosion and exposed roots

Fig. 11. Main dimensions of a pit and mound essential for their volume calculation as illustrated by Norman et al. (1995), modied.

based forest-gap model (Constantine et al., 2012). Additional relevant

data include event frequency (number of trees toppled per hectare
per year) and slope angle.
6.2. Calculation of soil displacement due to tree root growth

Trees are natural obstacles to washed or gravitationally-transported

rock particles of different fractions. The highest amount of trapped sediment on the upslope sides of coarse woody debris or standing trees is
typically observed in the close vicinity of rock faces on scree slopes, especially when the supply of fresh material is quite intense (Fig. 12).
However, because they are constantly under physical stress, trees
may exhibit lower growth rates or much faster decay rates than under
normal conditions on a forested slope. The amount of material accumulated against standing trees or coarse woody debris can be quantied.
For logs lying perpendicular to the slope prole such an attempt was
made by Raka and Orulk (2009), who employed a method developed
for the quantitative assessment of total accumulation rate (TAR), with
results of 1 to 4 m3 observed. A similar attempt at quantication by
Measeles (1994) used standing trees and stumps. However, in this
case the amount of trapped sediment was interpreted only as an effect
of soil creep, with values of up to 0.12 kg m1 year1 recorded. The
author also provided no detailed information regarding the formula
used.
Tree roots are frequently observed above ground, a phenomenon
often interpreted as an effect of their erosion and simultaneous exposure. Although this assumption allows for assessment of the amount
of eroded material (e.g. Dunne and Dietrich, 1982), certain constraints
should be taken into account (Dunne et al., 1978). A detailed description
of the method was provided by LaMarche (1968), who pointed to the
following limitations: 1) near-surface roots may be uncovered, which
is the result of an increase in diameter over time, 2) the depth of soil removed before the roots are uncovered is not known and 3) asymmetrical root exposure. The last limitation corresponds to a gently sloping
terrace on the uphill side of a tree, as well as a concave hollow on the
downhill side. Such a conguration is especially common on steep
slopes covered by old trees, and causes large discrepancies between
the maximum depths of root exposure and actual slope degradation
(LaMarche, 1968). Some trees are able to develop aboveground roots
(aerial roots) as a response to site conditions and the effect of morphological and structural adaptation e.g. within oodplain forest
(Wittmann and Parolin, 2005). All the above-mentioned problems
must be taken into account and caution must be kept when choosing
an appropriate site and trees for measurement.

Another attempt at quantication of tree growth and mortality is

based on the observation of tree roots that expand horizontally and vertically, causing physical displacement of soil mass. After root decay and
disappearance, the soil material collapses vertically (Gabet et al., 2003,
Fig. 1, p. 253) and thus contributes to net downslope soil ux (qsx)
Eq. (5). The variables included in the calculation are as follows: 1)
rooting depth, 2) root turnover time (growth and dieback of roots), 3)
density of root material and 4) slope angle. A general slope-dependent
equation calculating soil mass displacement via root growth and decay
was proposed by Gabet et al. (2003):

qsx

xr
r

where x (m) is the net horizontal displacement of soil, r (kg m2) is the
root mass per unit area, (year1) is the root turnover rate and
r (kg m3) is the density of root material. Not only do the root turnover rate and the density of root material vary between different ecosystems and tree species, these values also change signicantly throughout
the year, especially for ne roots, the growth and decay times of which
can be very rapid. For instance, in an oakwood ecosystem, ne-root biomass increased from nearly 13 t ha1 in April to 21 in July, then decreased to 10 t ha1 in November (Santantonio et al., 1977).

Fig. 12. Pile of loose material accumulated on the upslope side of a tree. This tree is growing on a scree slope in close vicinity to a rock face built of mudstones in the Stoowe Mts.,
Sudety, SW Poland. The material is derived from weathering processes deteriorating the
rock face.

262

Pawlik / Earth-Science Reviews 126 (2013) 250265

2010), 810 times (1250 years; amonil et al., 2009) or up to as many

as 30 times (300500 years, Denny and Goodlett, 1956) (but see
Norton, 1989, for a different methodological approach). A similar attempt at calculation of turnover time has been made with respect to
the formation and inlling of stump rot depressions (pits) and soil displacement via root growth (Phillips and Marion, 2006). The authors
found that given an average disturbance density of 23 m2 ha1 in the
case of stump rot pits, the time required for the entire surface to be affected was around 43,000 years, compared to around 6000 years for
soil displacement with a disturbance density of 160 m2 ha1 (Phillips
and Marion, 2006). These data suggest that the above processes can
have a cumulative effect on the geomorphology of forested hillslopes
and should not be neglected if we expect our results to be close to
reality.
7. Potential directions for future research conclusions and
nal remarks
Fig. 13. General scheme of feedbacks and interactions between a forest ecosystem and the
geomorphic domain of a forested hillslope (previously unpublished).

Recently, Raka and Orulk (2009) presented the following

formula allowing for the quantication of eroded material: TER =
[(ER + x) rr] (x rr) / 2, where ER = [hd (td tg)] / 2,
x = rr * tg er, TER is total erosion rate, ER is erosion rate, hd is
the difference in height between downslope and upslope ground
level at the tree trunk, td is trunk diameter, is slope inclination
and rr is root radius. Another method of solving the same problem
was described by Grtner (2007), who employed a dendrochronological approach focused on changes in the anatomical structure of
the annual rings of exposed roots. This latter technique seems to be
unburdened by the limitations associated with the traditional approach. With the use of dendrochronological methods, it is also possible to detect episodes of faster soil creep, which is frequently
evident in the shape of bent trees (Braam et al., 1987). Via treecoring, a set of proxy data regarding the moisture conditions of the
hillslope can be obtained, since soil creep is more intense during periods of higher rainfall (e.g. Pawlik et al., 2013).
6.4. Trees and palaeogeographical reconstructions
After collecting basic information regarding the present-day
magnitude and frequency of processes caused or modied by trees,
certain palaeogeographical reconstructions can be carried out. The
palaeogeographical signicance of trees can be considered on two
levels: 1) individual trees and 2) forest ecosystems. It has been suggested, for instance, that forest soils have been extensively mixed
during the Holocene and historic time periods (Phillips et al.,
2005). If so, the following features of montane forests must be
taken into account: 1) the spatial extent of trees and migration of
the timber line and 2) changes in forest type and tree species, with
a potentially different impact on slope cover (e.g. deeply rooting species vs. trees with shallow root systems). Together with the migration of trees onto previously unvegetated areas, a slope may be
affected simultaneously by the new factor of mechanical and chemical weathering directly attributed to the root zone. It seems that
during the Holocene, the most important process has been tree
uprooting leading to the rapid transport of rock mass. Some authors
have pointed to the possibility that uprooting-derived whole slope
surface remodelling may have occurred not only once, but even several times in the last 10,000 years at some sites, up to a depth of at
least 1 m. The rate of such forest turnover (the time required for an
area equivalent to 100% of the ground surface to be uprooted; Phillips
et al., 2008b) for the Holocene could be between 23 times (every
37515000 years; Brewer and Merritt, 1978; 2777 years; Lenart et al.,

Traditionally, trees are viewed as a stabilizing factor for slopeweathering cover against mass movement, with forests considered a
vegetation formation inhibiting surface wash and erosion. However,
given the contribution of individual trees to rock weathering, sediment
transport and accumulation are still rather apparent, even if quantitative methods are limited and eld study poses many difculties. This
thesis is supported by a body of literature that is particularly focused
on biomechanical and biochemical weathering, with the accumulative
and bioprotective functions of trees less well-studied. At present, the
most important issue is the integration of the efforts of different disciplines towards achieving an explanation of forested hillslope dynamics
based on variation in forest ecosystem changes, incorporating forest
ecology, palynology, anthracology, dendroecology etc. and research
methods adequate for each (White, 1979; Pickett and White (Eds.),
1985). The general assumptions of the proposed approach are shown
in Fig. 13.
A similar effort has already been made to bridge the gap between pedology and forest ecology (amonil et al., 2010a), as well as between climate change and sediment transport via tree uprooting (Constantine
et al., 2012). The patterns and interactions shown in Figs. 1 and 13
point to integration on different levels of complexity, with the problems
analyzed in terms of forest ecology having a fundamental effect on forested hillslope dynamics. At the same time the biology of individual
trees has to be considered, as this factor plays a key role in the development of e.g. rock faces, weathering front migration and changes in soil
biomantles within upper and lower forest belts. In this context, forms
and sediments depend in large measure on the extent (horizontal and
vertical), volume and structure of the root system, as well as the processes active in the root zone and rhizosphere. As a general overview,
it is suggested that the end product of root activity, together with the
presence of trees in different states of growth and decay within a forest community, should be viewed as a potential source of matter
supplying the geomorphic system as a whole. Finally, one of the
most urgent objectives is to utilize knowledge (even if still limited)
regarding contemporaneous processes and forms inuenced by
trees for palaeogeographical reconstruction, with special attention
focused on the Holocene.
Acknowledgements
I would like to thank Dr Ian E. Evans (Durham, U.K.), Dr Pavel
amonil (Brno, Czech Republic) and M.S. Krzysztof Urbaniak for their
many invaluable comments and suggestions which considerably improved the quality of this paper. Insightful comments from two anonymous reviewers are gratefully acknowledged. I also thank Dr Pavel
amonil (VUKOZ, Brno, Czech Republic) for giving me permission
to use his database from the Razula National Nature Reserve, Czech

 Pawlik / Earth-Science Reviews 126 (2013) 250265

Republic. Financial support was provided by research project no.

NN306032940.
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