Professional Documents
Culture Documents
Forest restoration with Betula ssp. and Populus ssp. nurse crops increases
productivity and soil fertility
Hendrik Stark a,, Arne Nothdurft b, Joachim Block c, Jrgen Bauhus a
a
Chair of Silviculture, Faculty of Environment and Natural Resources, University of Freiburg, Tennenbacher Str. 4, 79085 Freiburg, Germany
Institute of Forest Growth, Department of Forest and Soil Sciences, University of Natural Resources and Life Sciences (BOKU), Peter-Jordan-Strae 82, 1190 Vienna, Austria
c
Department of Forest Monitoring and Environmental Care, Research Institute for Forest Ecology and Forestry Rhineland-Palatinate (FAWF), Hauptstrae 16, 67705
Trippstadt, Germany
b
a r t i c l e
i n f o
Article history:
Received 14 May 2014
Received in revised form 30 November 2014
Accepted 3 December 2014
Keywords:
Nurse crops
Populus ssp.
Betula ssp.
Biomass production
Nutrient cycling
Forest restoration
a b s t r a c t
The rapid re-establishment of forests following large disturbances is being seen as one option to increase
the contribution of forests to climate change mitigation. The temporary inclusion of pioneer trees as
nurse crops on disturbed sites can facilitate the establishment of target tree species and may additionally
benet productivity and soil fertility. In this study we compared productivity and nutrient cycling
between stands of oak target species (Quercus robur and Quercus petraea) that were established with
and without widely spaced Betula ssp. or Populus ssp. nurse crops. Simulation results for a full rotation
of oaks (180 years) indicated that both types of forests, with and without nurse crops, have a comparable
total productivity. However, stands with nurse crops supplied 5996 Mg ha1 harvestable biomass after
20 years, whereas the rst harvest of biomass from stands without nurse crops would occur at least
30 years later. Nutrient element costs associated with the removal of Betula ssp. wood were low compared to Populus ssp. Also, nurse crop stands had up to 2.5 times larger pools of exchangeable base cations
in top mineral soils (030 cm) compared to mono-specic oak stands. The high soil cation pools may have
resulted from reduced leaching under nurse crops or the increased recycling of cations, also from deeper
soil depth, via litter fall and ne-root turnover. Our results show that forest reestablishment with pioneer
tree species may be a suitable tool for the rapid recovery of forest productivity and mitigation potential
following disturbances while simultaneously helping to maintain or increase soil fertility.
2014 Elsevier B.V. All rights reserved.
1. Introduction
Climate change mitigation strategies in Europe aim to off-set
fossil fuels and replace energy-intensive materials with wood.
Therefore, the supply of biomass from forests is considered a crucial component within the set of renewable energy options
(Wiesenthal et al., 2006; Stupak et al., 2007).
However, during the last few decades the intensity and frequency of disturbances caused by storms, droughts, res, pest
insect outbreaks or wet snow events in central European forests
has risen measurably (Schelhaas et al., 2003; Nilsson et al.,
2004). Thus, approximately 16,000 ha or 0.15% of the total forest
area in Germany are annually disturbed considering the time
between 1950 and 2012 (EFI, 2013). At the European level such
Corresponding author. Tel.: +49 761 203 3678.
E-mail addresses: hendrik.stark@waldbau.uni-freiburg.de (H. Stark), arne.nothdurft@boku.ac.at (A. Nothdurft), joachim.block@wald-rlp.de (J. Block), juergen.
bauhus@waldbau.uni-freiburg.de (J. Bauhus).
http://dx.doi.org/10.1016/j.foreco.2014.12.003
0378-1127/ 2014 Elsevier B.V. All rights reserved.
58
ssp., Quercus ssp. or Picea ssp. Often, forest restoration with these
species is realized by planting mono-specic stands, which can suffer amongst others from late frost events in large forest openings
(Lundmark and Hllgren, 1987; rlander, 1993; Groot and
Carlson, 1996; rlander and Karlsson, 2000; Agestam et al.,
2003). Therefore, the capacity of forests to provide biomass may
be substantially reduced, if, following disturbances, regeneration
is delayed and thus productivity reduced (Osterburg et al., 2013).
On the other hand, large open areas in forests offer opportunities for new, alternative and more resistant silvicultural systems.
Hence, pioneer trees can be used as nurse crops, which are temporary mixtures of fast growing and light-demanding tree species in
the overstorey and shade-tolerant target tree species in the understorey (Pommerening and Murphy, 2004). During the initial years
of stand development, the pioneer trees facilitate the establishment of other tree species beneath their canopy. One possible
nurse effect is the rapid recovery of forest micro-climate and thus
the amelioration of environmental extremes, such as frost (Carlson
and Groot, 1997; Schmidt-Schtz and Huss, 1998). Outside of central Europe, nurse crops are therefore commonly used for large or
small scale reforestations (Pommerening and Murphy, 2004) following clearfelling (Keenan et al., 1995; Lieffers et al., 1996) or natural disturbances (Perala and Alm, 1990; Drouineau et al., 2000;
Vallauri, 2005; Nelson et al., 2012) and for the afforestation of
abandoned agricultural lands (Mander and Jogman, 2000;
Gardiner et al., 2004; Uri et al., 2007).
Once trees of the target species have been successfully established and start to suffer from inter-specic competition from
nurse trees, the latter are typically removed (Cotta, 1828;
Shepperd and Jones, 1985). Often, however, the nurse trees are just
killed and not commercially harvested, because it is feared that
their felling and extraction might damage the established regeneration of target species.
To ensure that this biomass of nurse trees is not wasted but
used, a new approach to establish widely-spaced nurse crops for
the purpose of woody biomass production has been developed
(Unseld et al., 2012). This approach is based on the assumption that
the temporary inclusion of fast growing pioneer tree species may
facilitate an early harvest of biomass and thus an earlier return
of investment of stand establishment costs to the land owner.
Additionally, the overall stand productivity may increase through
competitive reduction between tree species (Vandermeer, 1989;
Man and Lieffers, 1997) and thus a complementarity in resource
use, which typically occurs when mixing light demanding overstorey with shade-tolerant understorey trees (Man and Lieffers, 1999;
Richards et al., 2010; Forrester, 2014). To achieve this, nurse crops
are managed on rather short rotations and a wide tree spacing is
employed that facilitates harvesting of nurse trees with minimal
damage to the regeneration (Unseld et al., 2012).
Nurse crops may also improve nutrient cycling following forest
disturbances, when the situation is often characterized by high
mineralization rates of organic matter and reduced uptake of water
and nutrients. This can lead to high rates of leaching of nutrient
elements such as calcium (Ca), potassium (K), magnesium (Mg)
and nitrogen (N) and consequently a reduction in soil fertility
(Hornbeck et al., 1986; Hendrickson et al., 1989; Bauhus and
Bartsch, 1995; Yanai, 1998). These nutrient element losses may
be aggravated, if the sites are subsequently used for intensive production and harvesting of biomass in nurse crops and nutrients
contained therein (Sverdrup et al., 2006; Worrell and Hampson,
1997).
Owing to the rapid growth and canopy closure of pioneer tree
species, they have a higher nutrient and water uptake, thus reducing leaching, soil temperatures and mineralization rates, when
compared to slower growing tree species (Prescott, 2002). Interactions between nurse and target tree species may further accelerate
59
Fig. 1. Chart of the study sites Kirchberg and Sobernheim as originally established
in 1991 (modied from Schmidt-Schtz and Huss, 1998). The set of plots with
mono-specic oak (oak-mono), aspen-oak and birch-oak stands assessed in the
present study were marked with bold lines and color codes.
To determine relevant base cation pools in forest oor and mineral soils, soil cores were collected from 12 randomly selected sampling positions per plot. We used larger cylinders for the forest
oor ( 170 mm) and smaller ones for the mineral soil (
50 mm). Mineral soil cores were divided into 010 and 1030 cm
depth layers. To obtain information from a greater soil depth, additional samples were collected from 3070 cm ( 24 mm) at four
out of 12 sampling positions. Samples were oven dried at 40 C
and their individual dry weight was recorded.
Subsequently, to reduce laboratory costs, pairs of samples
were bulked to yield six samples (two samples for the layer
3070 cm) per layer and plot. These bulked samples were sieved
(2 mm), ground and 25 ml of 1 M NH4Cl were used to extract
exchangeable base cations from 2.5 g soil sub-samples. We used
the mass of ne soil per unit of sample volume multiplied by
the respective nutrient element concentration to estimate nutrient element contents per sample volume and subsequently per
area. Likewise, forest oor samples were mixed, sub-samples
were ground to powder and subsequently digested in 65%
HNO3 at 190 C (pressure vessels, Loftelds Analytical Solutions,
Neu-Eichberg, Germany). The diluted solutions from extractions
and digestions were analyzed for element concentrations using
ICP-OES (Spectro, Kleve, Germany) (Knig, 2005). Organic carbon
(C) and nitrogen (N) concentrations were measured using dry
combustion (Truspec, Leco, St. Joseph, USA). Soil pH was measured in CaCl2.
To determine base cation pools within the woody biomass, at
each plot and for each tree species ve trees were destructively
sampled across the entire range of all tree diameters at breast
height, respectively. Diameter at breast height (cm) and tree height
(m) were recorded immediately after felling, and stem discs were
extracted at 2 m intervals along aspen and birch stems. The length
60
of oak stems with a diameter larger than 4 cm was divided into six
equally long sections and stem discs were extracted at these relative intervals. From each sample tree with a diameter at breast
height thicker than 4 cm three branches were sampled from the
lower, middle and upper crown section, respectively. Subsequently, the fresh weight of all remaining branches and stem sections was recorded. Stem disc fresh weight was recorded
separately for wood and bark. Oak trees with a diameter at breast
height thinner than 4 cm were assigned to the branch compartment (Stark et al., 2013).
For element analysis, wood shavings of stem discs were collected from two perpendicularly cross-sectional transects while
stem bark fragments were chipped. Both types of sub-samples
were oven dried at 40 C until constant weight, mixed at the tree
level, and ground. Likewise, sample branches were chipped, mixed
at the tree level and ground. Finally, compartment-specic element
analysis was undertaken as described above for forest oor
samples.
The respective compartment-specic aboveground woody biomass of all trees was estimated using allometric biomass equations
(Stark et al., 2013). The species-specic biomass of roots between 2
and 5 mm was measured using the soil cores (0 and 30 cm depth)
collected for this study, while the biomass of roots larger 5 mm and
stumps was estimated according to equations provided in Wang
(2006). The biomass of birch regeneration with diameters at breast
height ranging between approximately 1 and 5 cm was estimated
using equations from Uri et al. (2007). The nutrient element pools
contained within sample tree stem wood, stem bark and branches
were nally calculated using the measured compartment-specic
biomass and element concentrations.
We did not measure nutrient element concentrations in understorey birch regeneration. In order to estimate the nutrient element pools within the aboveground woody biomass of birch
regeneration we used the concentrations measured in sample tree
branches, assuming that the stem-wood to stem-bark ratio and
thus the nutrient element concentrations both in sample branches
of nurse crop trees and birch regeneration were comparable. Also,
we did not measure nutrient element concentrations in roots.
Therefore, we used information provided in the literature to infer
the average ratio between nutrient element concentrations in tree
branches and roots. Based on such comparisons, we assumed tree
roots with diameters between 2 and 5 mm to have comparable
nutrient element concentrations as measured in our sample tree
branches (Jacobsen et al., 2003), and we assumed stumps and roots
>5 mm to have 50% of these branch concentration (Rademacher,
2005; Block et al., 2007).
For planted aspen and birch in aspen-oak and birch-oak plots
Pn
we calculated the biomass and nutrient
element pools per ha on
t
j1 j
each sample plot separately as nab
10; 000, with t j being the
single tree biomass or nutrient element pool of each planted aspen
and birch tree on a specic plot, and a and b being the width of the
regular plant spacing in x and y direction. The biomass and nutrient element pools per ha of mono-specic oak, natural regeneration and understory oaks were estimated by means of 2 m wide,
diagonal transect samples.
b3 Ifsite Sobernheimg
1
b6 Iftreatg Ifdepthg
1
61
biomass
Ca
100
50
50
300
300
150
150
K
Mg
200
200
100
100
60
60
30
30
300
300
150
150
40
40
20
20
birch
underst. oak
aspen
underst. oak
oakmono
birch
underst. oak
aspen
underst. oak
100
oakmono
Sobernheim
3. Results
Kirchberg
62
63
Sobernheim
Kirchberg
biomass [Mg/ha]
Table 1
Total biomass and nutrient element removals from mono-specic oak stands and
aspen-oak and birch-oak stands in Kirchberg and Sobernheim. In all cases the biomass
produced by a beech understorey is included, too.
Site
Treatment
Biomass
Ca
Mg
Oak-mono
Aspen-oak
Birch-oak
Mg/ha
898
928
911
kg/ha
1733
1896
1746
kg/ha
1291
1362
1286
kg/ha
255
271
257
kg/ha
1748
1871
1841
kg/ha
127
152
144
Oak-mono
Aspen-oak
Birch-oak
823
828
820
1643
1728
1651
1259
1313
1234
203
216
202
1773
1855
1814
122
137
131
350
300
250
200
150
100
50
350
300
250
200
150
100
50
100
80
60
40
20
0
100
80
60
40
20
0
ci 0.0250.975
aspenoak mean
birchoak mean
oakmono mean
ci 0.0250.975
oaknursecrop mean
oakmono mean
10
15
20
50
100
150
Fig. 3. Left panels: Reconstructed aboveground woody biomass development of aspen, birch and mono-specic oak (oak-mono) stands between the ages 0 and 20 years at
both study sites, respectively. Mind that the biomass of understory oak could not be included here. They contributed about 3.2 Mg biomass per ha at the age of 20 years. Right
panels: Biomass development of oak stands established with (black) and without (red) a nurse crop. These pools include a beech understorey, which is commonly installed to
suppress the development of epicormic branches in oak stems. Mind that the age of oaks established without a nurse crop corresponds to the stand age given at the x-axis,
while oaks established with a nurse crop were ve years younger. For details please refer to the Supplementary Tables S9S12. Presented are means and 95% condence
intervals of 199 model iterations. (For interpretation of the references to color in this gure legend, the reader is referred to the web version of this article.)
Kirchberg
Sobernheim
1000
1000
aspenoak
800
birchoak
800
600
oakmono
600
400
400
200
200
0
0
50
100
200 0
50
100
200
age [years]
Fig. 4. Cumulative biomass removals derived from three exemplied model runs for aspen-oak, birch-oak and mono-specic oak stands across the complete rotation period.
Included is a beech understorey, which would be introduced at the oak height of 12 m to prevent epicormic branches in oak stems. For details please refer to the
Supplementary Tables S9S12.
64
Ca [mg/g]
12
10
8
ab
a
6
4
2
birchoak
aspenoak
oakmono
4.0
pH CaCl2
3.8
3.4
3.2
birchoak
aspenoak
oakmono
50
40
20
birchoak
aspenoak
10
oakmono
Fig. 6. Soil acidity (pH) measured in mineral soils in 010 cm depth under aspenoak, birch-oak and mono-specic oak (oak-mono) plots. Different letters indicate
signicant differences between treatments and were produced from multiple pairwise comparisons of means.
30
140
130
120
110
100
90
oakmono
aspenoak
birchoak
80
70
3.6
14
Fig. 7. Base cation saturation of the exchange complex as measured in the mineral
soil in 010 cm depth aspen-oak, birch-oak and mono-specic oak (oak-mono)
plots. Different letters indicate signicant differences between treatments and were
produced from multiple pair-wise comparisons of means.
40
60
80
100
120
C [mg/g soil]
Fig. 8. Regression curves for cation exchange capacity (CEC) dependent on the
organic carbon concentration in the mineral soil in 010 cm depth (Eq. (3)). The pH
(CaCl2) is xed at its overall mean of 3.4.
C*1000
Kirchberg
Sobernheim
measured predicted
[kg/ha]
[%]
measured predicted
[kg/ha]
[%]
100
400
100
400
50
200
50
200
Ca
1200
1200
400
600
400
600
200
0
1000
200
0
1000
400
500
400
500
200
Mg
0
700
400
350
200
0
700
400
350
200
0
oakmono
aspenoak
birchoak
oakmono
aspenoak
birchoak
400
8000
400
200
4000
200
oakmono
aspenoak
birchoak
4000
200
oakmono
aspenoak
birchoak
0
8000
65
forest floor
Fig. 9. Left bar triplet of each panel: Measured nutrient element pools per soil
depth layer with standard deviations (error bars) given for the overall element pool
of all depth layers, respectively. Right bar triplet of each panel: To increase the
visibility of treatment effects, the relative difference between the treatment aspenoak and the mono-specic oak reference stand as well as the relative difference
between the treatment birch-oak and the mono-specic oak reference stand was
calculated from layer-specic nutrient element pool predictions using signicantly
parametrized models (1). Thus, for each depth layer (for each shade of gray) the
element pool under mono-specic oak plots (oak-mono) was set to 100% and the
element pools under aspen-oak or birch-oak systems could be smaller or larger
than that, respectively. Test results of pair-wise comparisons of means of empirical
data can be found in Tables S3S6.
oak, while in Sobernheim only those in aspen-oak were signicantly larger than in mono-specic oak stands.
In Kirchberg, the exchangeable pools of Mg were 40 (25), 25
(25) and 14 (6) kg ha1 in aspen-oak, birch-oak and mono-specic
oak plots, respectively; the corresponding pools in Sobernheim
were 26 (6), 20 (5) and 17 (8) kg ha1 in mono-specic oak,
aspen-oak and birch-oak, respectively. Also the Mg pools in
Kirchberg under aspen-oak and birch-oak stands were signicantly
larger than in mono-specic oak, while in Sobernheim only in
aspen-oak stands signicantly larger pools than in mono-specic
oak stands were observed.
The pools of N were 1680 (417), 1613 (266) and 1595
(353) kg ha1 in aspen-oak, birch-oak and mono-specic oak plots
in Kirchberg and 1122 (187), 1171 (158) and 1293 (165) kg ha1 in
aspen-oak, birch-oak and mono-specic oak in Kirchberg. No signicant difference were observed in Kirchberg, but in Sobernheim
the N pools in 010 cm mineral soil under mono-specic oak plots
were signicantly larger than under aspen-oak plots. P pools in
mineral soils were not analyzed.
Fig. 10. Comparison of the amount of biomass (bulk wood including bark with
diameter > 7 cm) that can be harvested per mass unit nutrient element from several
tree species of central Europe. The larger the biomass per unit nutrient the lower
the nutrient cost of biomass harvest. Error bars indicate the standard deviation
among sample trees. Different letters indicate signicant differences between tree
species and were produced from multiple pair-wise comparisons of means. Data
were compiled from this study, Jacobsen et al. (2003) and Pretzsch et al. (2013)
(sample sizes: Betula pendula x Betula pubescens 12, Fagus sylvatica 195, Picea abies
123, Pinus sylvestris 94, Populus tremula x Populus tremuloides 14, Pseudotsuga
menziesii 72, Quercus ssp. 101).
In model (1) the xed study site effect generally explained a signicant share of variation (Table S16) for all elements except Ca, and
soil depth and often the interaction of soil depth treatment and
soil depth study site were signicant xed effects, too. According to this model the Ca pools were up to 140%, K up to 56% and
Mg pools up to 165% higher in mineral soils between 0 and 10 cm
depth under aspen-oak or birch-oak plots compared to soils under
mono-specic oak plots (Tables S3 and S4). A similar pattern was
observed in the mineral soil between 10 and 30 cm depth.
3.3. Nutrient element costs of wood
According to model (2) the xed effects tree species and
biomass compartment both signicantly affected the nutrient element costs of the harvestable woody biomass (Table S17). On average 25%, 41%, 39% and 2% less total aboveground woody biomass
66
per unit Ca, K, Mg and P could be harvested from aspen when compared to birch; 9% more aboveground woody biomass could be
harvested per unit N from aspen than from birch. Alternatively,
69%, 47%, 54% and 13% less stem wood and stem bark per unit
Ca, K, Mg and N might be removed from aspen than from birch
and 5% more stem wood and stem bark could be harvested per unit
N from aspen compared to birch. An additional analysis of variance
using our empirical data supported these model predictions, indicating signicant differences in K and Mg costs between aspen
and birch (Fig. 10, Tables S18 and S19).
Considering the harvest of woody biomass compartments with
diameters larger than 7 cm of major tree species in central Europe,
woody biomass of Pseudotsuga menziesii had the lowest Ca costs,
whilst aspen and birch took a moderate rank among most other
species; Ca costs of aspen and birch stems were signicantly lower
than those of Quercus ssp. K costs, on the other hand, were moderately high in birch and Picea abies. In contrast, K costs of aspen and
oak stems were comparatively high and Mg costs were lowest for
stems of Pseudotsuga menziesii. In general, N and P costs of aspen
and birch woody biomass were comparatively high.
Finally, harvesting without branches increased the amount of
woody biomass per unit Ca, K, Mg, N and P in aspen by 12%, 11%,
17%, 30% and 44% and in birch by 23%, 39%, 32%, 35% and 63% irrespective of the study site (Eq. (2)).
4. Discussion
4.1. Effects of nurse crops on productivity
Many trials have studied the effect of tree species mixtures on
stand productivity compared to pure plantations (Man and
Lieffers, 1999; Binkley, 2003; Forrester, 2014). Only few studies
have focused on tree species mixtures with aspen or birch (Man
and Lieffers, 1999; Bergqvist, 1999; Johansson, 2003; Comeau
et al., 2009). We are not aware of studies that have considered
aspen and birch as nurse crops with rotations of only two decades.
In this study, aspen and birch at 4 4 m plant spacing in aspenoak and birch-oak plots produced between 59 and 96 Mg of aboveground woody biomass per ha. This biomass was harvestable after
20 years (or earlier) and corresponded to a mean annual increment
between 3 to 5 Mg ha1 year1 (Fig. 2, Tables S1 and S2). Despite
the water logged soil conditions and the cool mountainous climate,
the productivity of the studied nurse crops were comparable to
that of other widely spaced nurse crops in Germany (Nelson
et al., 2012; Unseld et al., 2012). They were also comparable to productivities reported in the literature for some aspen or birch shelter woods in Estonia or Sweden (Maard, 1996; Bergqvist, 1999;
Johansson, 2003; Karacic et al., 2003), but signicantly lower,
approximately only 50%, than the productivity reported for plantations in Denmark and Germany (Nielsen et al., 2014; Liesebach
et al., 1999). However, the difference may be explained by the
much lower stocking of the studied nurse crops and also the lower
fertility of forest soils when compared to former agricultural soils
that are typically used for woody biomass plantations.
Despite of their comparatively slow growth, mono-specic oak
stands had produced as much biomass at the age of 20 years as
aspen or birch nurse crops. This may be mostly attributable to their
much higher stand densities of approximately 5000 N/ha. Additionally, it may also be related to the contrasting, species-specic
growth dynamics of aspen, birch and oak. During the rst decade
post establishment, high biomass growth in aspen and birch stands
would have exceeded those in mono-specic oak stands, but also
peaked early to subsequently decline below the level of monospecic oak stands leading to comparable biomass pools at the
age of 20 years (Gutsell and Johnson, 2002).
67
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