Eur J Forest Res (2014) 133:969–982
DOI 10.1007/s10342-014-0815-5
ORIGINAL PAPER
Nutrient and heavy metals in decaying harvest residue needles
on drained blanket peat forests
Zaki-ul-Zaman Asam • Mika Nieminen • Annu Kaila • Raija Laiho •
Sakari Sarkkola • Mark O’Connor • Connie O’Driscoll • Afshan Sana
Michael Rodgers • Xinmin Zhan • Liwen Xiao
Received: 2 August 2013 / Revised: 7 April 2014 / Accepted: 25 April 2014 / Published online: 10 May 2014
 Springer-Verlag Berlin Heidelberg 2014
Abstract Harvest residue decomposition can signifi-
cantly contribute to nutrient and heavy metal exports to
receiving water courses. This study monitors the nutrient
and heavy metal dynamics in decaying Sitka spruce and
lodgepole pine harvest residue needles on Atlantic blanket
peat forests in the west of Ireland. Using the litterbag
method, harvest residue was placed both within and
between furrows in two uncut forest and two clear-cut sites.
On the clear-cut sites, the litterbags were positioned outside
the harvest residue piles (i.e. brash windrows). Over the
2-year monitoring period, the needles decomposed slower
at the clear-cut sites than the uncut forest sites, with mass
losses of 46–55 and 58–77 %, respectively. Approximately
20 % less phosphorous (P) was released from the decaying
needles at the clear-cut sites, while nitrogen (N) was
released only at the uncut sites. Tree species was a sig-
nificant factor contributing to nutrient and heavy metal
Communicated by A. Merino.
Z. Asam  M. O’Connor  A. Sana  M. Rodgers  X. Zhan
Civil Engineering, National University of Ireland, Galway,
Ireland
Z. Asam  M. O’Connor  C. O’Driscoll  L. Xiao (&)
Department of Civil, Structural and Environmental Engineering,
Trinity College Dublin, Dublin, Ireland
e-mail: Liwen.xiao@tcd.ie
M. Nieminen  A. Kaila  S. Sarkkola
Southern Finland Regional Unit, Finnish Forest Research
Institute, Vantaa, Finland
R. Laiho
Western Finland Regional Unit, Finnish Forest Research
Institute, Parkano, Finland
•
release and accumulation patterns, with higher concentra-
tions of aluminium (Al), nickel (Ni), cadmium (Cd) and
zinc (Zn) in the decaying spruce needles than in pine.
Conversely, the spruce needles showed accelerated deple-
tion of calcium (Ca) and magnesium (Mg) relative to the
pine. The harvest residue needle positioning (inside furrow/
between furrows) and the site soil characteristics contrib-
uted significantly to Al transformations in spruce needles
and iron (Fe) in both spruce and pine needles, with more
accumulation occurring inside the furrows where Al and Fe
contents of the peat were high. Manganese (Mn) was
released from the needles in three of the four sites with a
total release of over 90 % within 2 years. In the remaining
site, where the Mn content of the peat was high, an accu-
mulation of Mn in the needles was observed. The decom-
position of needles on blanket peat catchments may be a
significant source of P to receiving water courses, owing to
their fast release of P, but not a likely source for N export.
Keywords Blanket peatlands  Harvest residue 
Litterbag experiment  Needle decomposition  Nutrient
release  Heavy metals
Introduction
Large areas of blanket peat catchments were afforested
between 1950s and 1990s in Ireland and the UK, with the
bulk of plantations either at or approaching commercially
harvestable age. An increase in nutrient leaching from
harvested peatland forests has been reported by several
studies (Rosén and Lundmark-Thelin 1987; Nisbet et al.
1997; Lundin 1998, 1999, 2000; Nieminen 1998, 2003,
2004; Ahtiainen and Huttunen 1999; Cummins and Farrell
2003a, b; Rodgers et al. 2010). Peatland-dominated
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catchments in Ireland contain oligotrophic headwaters,
many of which contain IUCN (International Union for the
Conservation of Nature) Red List species (e.g. salmonids or
freshwater pearl mussels) and are highly sensitive to
increased nutrient input (O’Driscoll et al. 2011, 2012,
2013).
In the conventional stem-only harvesting method, the
harvest residues (needles, twigs and branches) remain on
site post-stem extraction. The decomposition of such har-
vest residues is seen as a potentially high source of nutri-
ents to receiving water bodies from drained peatlands
(Rodgers et al. 2010; Kaila et al. 2012), but may be con-
sidered a particularly important source of nutrients from
blanket peat catchments in Ireland. The stand density in
blanket peat catchments in Ireland is relatively high (about
2,800 trees ha-1 at clear-felling, Rodgers et al. 2010)
compared with those of for example Scandinavia, as no
thinning operations are performed during stand develop-
ment. Stand volume in sites approaching clear-felling
phase is also significantly larger for blanket peat forests
([400 m3 ha-1) than peatland stands in more northern
areas (150–300 m3 ha-1). This means that the amount of
cutting residues in clear-felled blanket peat forests may be
significantly larger than, for example, Scots pine (Pinus
sylvestris)-dominated peatlands in Scandinavia. The above-
ground harvest residues of lodgepole pine in blanket peat
forests in west of Ireland were estimated at over 80,000 kg
(d.w.) ha-1 (Asam 2012; Asam et al. 2014), while the
harvest residues of, for instance, Scots pine in Fenno-
scandia amount to only about 14,000 kg (d.w.) ha-1 at
clear-felling phase (Palviainen and Finér 2012).
Similarly, fresh above-ground harvest residues of lodge-
pole pine in the west of Ireland were estimated to contain
25–33 kg ha-1 of phosphorus (P) and 330–430 kg ha-1 of
nitrogen (N), with the needles accounting for nearly 50 % of
the total stores of N and P in the above-ground harvest res-
idues (Asam 2012; Asam et al. 2014). These figures for N and
P in harvest residues are significantly higher than those found
for Scots pine-dominated sites in Scandinavia, which were
reported to be between 8 and15 kg ha-1 for P, and
77–135 kg ha-1 for N (Hyvönen et al. 2000; Palviainen and
Finér 2012).
During forestry operations on blanket peat catchments,
harvest residues are utilised as brash mats to improve the
poor soil-carrying capacity against heavy machinery that
transports the harvested stems to the roadside (Rodgers
et al. 2010). As the remaining harvest residues make the
replanting of trees difficult, they are collected together to
form windrows, between which the trees are planted.
Reported measurements for these windrows find they have
a width of approximately 4 m, run along the main slope in
parallel rows at intervals of approximately 12 m and cover
about 25 % of the clear-cut area (Rodgers et al. 2010;
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Eur J Forest Res (2014) 133:969–982
Asam et al. 2012). Windrowing of harvest residue (brash)
generally occurs shortly before the replanting of the site,
and therefore, a significant portion of the needles can fall
from the branches and do not end up in the windrows. To
fully understand the processes controlling nutrient and
heavy metal release from harvest residues in blanket peat
catchments, information is required on needles decompo-
sition from both the areas within windrows and areas
outside windrows, where needles fell down before
windrowing.
Forested blanket peats differ from other forestry-drained
peatlands in that they were drained for forestry by closely
spaced drainage furrows (about 1 m wide and at 3–5 m
intervals) which accounts for a considerable percentage of
a site area (Ledger and Harper 1987; Anderson et al. 2000;
Rodgers et al. 2010). In standing forests, litter is deposited
and accumulates in these furrows where the majority of the
water flows before leaving the area. After harvesting, a
significant fraction of harvest residues accumulates in the
furrows. Differences in redox and temperature conditions
within drainage furrows and the areas between furrows
may significantly affect the release and accumulation pat-
terns of nutrients and heavy metals.
Several studies have investigated the decomposition of
conifer needle litter and harvest residues on mineral soils
(Nilsson 1972; Berg and Staaf 1980; Rustad and Cronan
1988; Laskowski and Berg 1993; Rustad 1994; Laskowski
et al. 1995; Lundmark-Thelin and Johansson 1997; Vest-
erdal 1999; Palviainen et al. 2004a, b; Lehto et al. 2010)
and peatlands in Scandinavia and North America (Moore
et al. 2005; Kaila et al. 2012), whereas there is no data
available on harvest residue decomposition and nutrient
dynamics from drained blanket peat forests in Ireland.
Notwithstanding the differences, in both the quantities of
harvest residues and operational forestry practices, the
decomposition of harvest residues in blanket peat catch-
ments in Ireland may differ from that of drained peatlands
in more northern areas due to climatic conditions. Irish
blanket peats exist in a relatively mild climate with no
distinct snow or frozen soil periods. Precipitation
([2,000 mm year-1) in blanket peat areas may be two- to
threefold higher when compared to peatland-dominated
areas in Scandinavia.
Decomposition of conifer needles is strongly related to
tree species (Kaila et al. 2012) and blanket peats with Sitka
spruce (Picea sitchensis) and lodgepole pine (Pinus con-
torta) as the dominant tree species may have different
nutrient release patterns than that reported earlier for other
types of peat soils with different tree species. A general
trend in harvest residue needle decomposition is that ele-
ments such as P and particularly potassium (K) and mag-
nesium (Mg), which are mostly present in the needle cell
solution, are released during the early phases of
Eur J Forest Res (2014) 133:969–982
decomposition, while N is mostly accumulated and im-
mobilised (Kaila et al. 2012). This immobilisation is
interpreted to be because heterotrophic decomposers, par-
ticularly fungal hyphae, transport N from soil to harvest
residues, where decomposition is nitrogen-limited. Simi-
larly, the different heavy metals are also accumulated,
especially in areas with high atmospheric heavy metal
deposition (Kaila et al. 2012). After losing their protective
epiderm and cuticle, needle litter behaves like mosses and
lichens by easily absorbing heavy metals from air and soil,
but the mechanisms behind heavy metal absorption and
accumulation are largely unknown.
In this study, the decomposition of lodgepole pine and
Sitka spruce harvest residue needles was investigated on two
uncut forest sites and two clear-cut sites on Atlantic blanket
peats in west of Ireland. It was expected that the nutrient [N,
P, K, calcium (Ca), Mg, boron (B)] and heavy metal [zinc
(Zn), manganese (Mn), iron (Fe), aluminium (Al), lead (Pb),
copper (Cu), nickel (Ni), cadmium (Cd)] release and accu-
mulation patterns in the clear-cut sites would differ from the
uncut forest sites and that the decomposition patterns were
expected to vary between the needles in the drainage fur-
rows and those between the furrows. We also expected that
under similar conditions, we observe different decomposi-
tion behaviour between the needles of Sitka spruce and
lodgepole pine (Kaila et al. 2012).
Materials and methods
Site description
This experiment was conducted on four sites in two
Atlantic blanket peat-dominated catchments (two sites in
each catchment) in the west of Ireland, the Glennamong
(53580 2.9900 N, 9360 43.5100 W, 69 m a.s.l.) and Skerdagh
(53560 4.0100 N, 9300 36.1700 W, 125 m a.s.l.) located
approximately 5 km apart. Both catchments were drained
with furrows and afforested in 1970s. The dominant tree
species at the Glennamong sites was lodgepole pine (Pinus
contorta) ([95 % of tree volume), and the secondary
species was Sitka spruce (Picea sitchensis). At the Sker-
dagh sites, the dominant tree species was Sitka spruce
([90 %) and secondary species was lodgepole pine. The
assigned names for the four sites used in this study are as
follows: Glennamong clear-cut, Glennamong uncut, Sker-
dagh clear-cut and Skerdagh uncut. The Glennamong clear-
cut site was harvested in August 2009 and the Skerdagh
site in January 2009, using conventional stem-only har-
vesting where the harvest residue was left on site. The
clear-cut sites and uncut sites were about 300 and 150 m
apart in the Glennamong and Skerdagh catchments,
respectively.
971
The two catchments have similar hydrological condi-
tions and receive an average precipitation of approximately
2,000 mm year-1 with average rain days ([ 0.2 mm pre-
cipitation) in excess of 225 per year. From recordings at the
weather station operated by the Marine Institute, about
5 km from the study sites, the 2009–2011 mean annual
temperature was 9.8 C, with means of 4.4 C in January
and 14.4 C in July. The average slope at the Glennamong
sites was 5–10 and 2–5 at the Skerdagh sites.
The peat soil in both the Glennamong sites is highly fibric
with a depth of[1 m and overlies mainly quartzite and schist
bedrock. At Skerdagh, layers of mineral soil have been
deposited within the[1 m peat layer, and there were more of
those layers in the peat of the Skerdagh uncut site compared
to the clear-cut site. The higher mineral content in the
Skerdagh sites is reflected as lower organic matter and car-
bon (C) contents, as well as higher Al and Fe contents in its
peat when compared to the Glennamong sites (Table 1).
However, the most significant difference between the four
sites was that the Skerdagh uncut site had significantly higher
contents of Ca, Fe, Al and Mn than the other three sites.
Decomposition experiment and analysis
The needles for both species were collected from the
Glennamong clear-cut area within a few hours of harvest-
ing. Branches with living green needles, representing the
top 10 m of the tree crown, were collected. The needles
were stripped from the branches by hands, air-dried in the
laboratory and hand-mixed thoroughly. Subsamples of said
needles were drawn for the determination of the initial
nutrient and heavy metal concentrations.
Litterbags of size 10 9 10 cm were made from nylon
sheets with a mesh size of 1 9 1 mm. 1.5–2.0 g (d.w.) of
needles were enclosed in each litterbag and then equipped
with plastic tags labelling the initial needle material
weight. The litterbags were placed systematically in five
sampling positions (at approximate 10 m spacing) between
the furrows (BF) and five inside the furrows (IF) at each of
the four sites, with three pine needle and three spruce
needle bags at each position. The bags were placed directly
on top of the litter layer and anchored using plastic pins. In
the clear-cut sites, the bags were installed in the harvest-
residue-free areas. The incubation study lasted 2 years
(from November 2009 to November 2011), and the litter-
bags were sampled at 0.5, 1.0 and 2.0 years from com-
mencing the experiment. On each sampling occasion, one
bag for pine and one for spruce were systematically col-
lected from each of the 5 sampling positions in BF and 5
sampling positions in IF from each of the four sites. Thus,
the total number of bags was 240 (4 sites, 2 tree species, 2
sampling positions (between furrow/inside furrow), 3
sampling occasions and 5 replicates).
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972 Eur J Forest Res (2014) 133:969–982

Table 1 Peat properties of the Sites Glennamong clear-cut Glennamong uncut Skerdagh clear-cut Skerdagh uncut
study sites at 0–20 cm peat
layer Organic Matter (%) 95.5 95.7 67.4 64.7
C (%) 53.9 53.1 38.5 37.6
N (%) 2.4 2.3 1.7 2.2
P (mg kg-1) 364 348 398 619
K (mg kg-1) 274 228 597 366
-1
Ca (mg kg ) 791 896 600 4,850
Peat samples were collected
systematically from a grid of 27 Mg (mg kg-1) 1,000 930 869 498
sampling positions within each Fe (mg kg-1) 1,080 1,310 3,110 72,100
site, using 5.5-cm-diameter Al (mg kg-1) 1,010 1,340 1,720 3,550
gouge auger, and were pooled to
give one composite sample for B (mg kg-1) 3.52 3.49 1.73 0.026
-1
analysis. Prior to analysis, the Zn (mg kg ) 36.0 4.33 4.65 16.3
pooled samples were dried at Mn (mg kg-1) 10.9 4.59 10.3 2,350
?60 C, milled to pass a 2-mm
Pb (mg kg-1) 16.6 15.8 12.2 11
sieve and analysed for C and N
using LECO CHN-1000 Ni (mg kg-1) 1.89 3.01 6.38 1.94
analyser and the other elements Cd (mg kg-1) 0.14 0.13 0.08 0.026
using ICP/AES after dry ashing Cu (mg kg-1) 1.66 1.95 0.859 0.150
and digestion in HCl

Table 2 Initial element concentrations and ratios of C/N, C/P and

dry mass. For chemical analyses, the contents of five rep-
N/P in harvest residue needles licate bags were combined, leaving 48 samples that were
ground using MF 10 IKA-analytical mill to pass a 2-mm
Element Pine needles Spruce needles
sieve. Subsamples of needles drawn for the initial con-
C (%) 51.8 52.1 centrations of elements were dried and ground the same
N (%) 1.42 1.08 way. The ground samples were analysed for N and C
P (mg kg-1) 1,170 1,100 concentrations using LECO CHN 1000 analyser, and for P,
K (mg kg-1) 5,890 5,900 B, K, Ca, Mg, Fe, Al, Mn, Zn, Pb, Ni, Cu and Cd con-
Ca (mg kg-1) 1,220 3,950 centrations using ICP/AES following dry ashing and
Mg (mg kg-1) 1,010 1,270 digestion in HCl. The analysis results were above the
Fe (mg kg-1) 27 21.1 detection limits of analytical devices, except for 10 sam-
Al (mg kg-1) 255 30.2 ples of 50 tested for Pb, which had concentrations below
B (mg kg-1) 15.2 19.5 the detection limit. In the calculations, those values for Pb
Zn (mg kg-1) 22.2 6.45 were substituted with a value equal to half the detection
Mn (mg kg-1) 177 511 limit.
Pb (mg kg-1) 0.172 0.147 Initial concentrations of nutrients and heavy metals in
Ni (mg kg-1) 0.351 0.144 harvest residue pine and spruce needles are shown in
Cd (mg kg-1) 0.091 0.021 Table 2. We investigated the changes in mass (g g-1) and
-1
Cu (mg kg ) 0.705 1.48 element contents (the content: concentration, mg g-1 9
C/N 36.5 48.2
remaining mass, g) in the decaying needles over 2 years.
C/P 442.7 473.6
Changes in the mass and nutrient contents over the study
N/P 12.1 9.8
period were expressed as percentage values from the initial
content.
The effect of tree species (lodgepole pine/Sitka spruce),
litterbag placement (inside furrows/between furrows) and
After collection and pre-drying at laboratory tempera- stand treatment (uncut/clear-cut) on mass loss and nutrient
ture, the contents of each litterbag were removed and and heavy metal dynamics in decaying needles was tested
placed on a clean tray in order to remove roots, soil, insects using linear mixed model analysis which enables the con-
and any other alien material that had entered the litterbags. sideration of potential autocorrelation between successive
Each needle was inspected separately, and if there was any measurements (see also Kaila et al. 2012). We identified
alien material on the surface of the needle, it was gently two hierarchical levels in our data set that were used as
brushed off. The cleaned needles were then dried at random variables in the models: 1) variation between the
?60 C to constant mass and weighed for the remaining sites (Skerdagh clear-cut, Skerdagh uncut, Glennamong

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Eur J Forest Res (2014) 133:969–982
clear-cut, Glennamong uncut) and 2) variation between the
four measurement time points (November 2009, May 2010,
November 2010 and November 2011). The explanatory
(fixed) variables were time (years), site dummy (Skerdagh,
Glennamong), tree species dummy (lodgepole pine, Sitka
spruce), harvest dummy (clear-cut, uncut) and drainage
dummy (inside furrows, between furrows).
It has been shown that litter may act as a temporary pool
for metals from the soil around and below litter (Scheid
et al. 2009). The Fe and Mn contents of peat in the Sker-
dagh uncut site were significantly higher than in any of the
other three sites, and we expected this to be reflected in the
sites litter Fe and Mn dynamics during decomposition. In
testing the effects of explanatory variables on the contents
of Fe and Mn in needles, we therefore used respective soil
Fe and Mn contents as additional fixed explanatory vari-
ables in the mixed models.
The mixed model had the following form:
yij ¼ aij þ b1 x1ij þ b2 x2ij þ    þ bn xnij þ uj þ eij ð1Þ
where yij is the mass loss or the element (see Table 2)
content (%) in the harvest residue needle sample occasion
i on the site j, a is the intercept, b1 … bn are the model
parameters i.e., the regression coefficients, x1ij … xnij, are
the explanatory variables, u is the random effect of the site
j and eij is the random error that accounts for the variation
among the sample occasion i within the sites. The random
effects were assumed to be independent and to follow
normal distribution with zero mean and constant variances
and covariances at each level. If the relationship between
the explanatory variable and the dependent variable was
nonlinear, the explanatory variable was linearised with
exponent or natural logarithm.
The models were estimated with the restricted iterative
generalised least-square (RIGLS) method using MLwiN
2.0 software (Rasbash et al. 2001). A parameter was
determined to be significant if its absolute value was more
than twice the size of the standard error. The value of
-2(log-likelihood) was used to compare the overall
goodness of fit of the models of increasing number of
explanatory variables. The explanatory variables were
added to the model one after another premising that the
added variable must maximise the decrease of the log-
likelihood value (e.g. Sarkkola et al. 2009; Kaila et al.
2012). The order of the added variables did not affect the
model predictions. The model reliability and accuracy were
evaluated by calculating the systematic error (absolute
bias, Ba), relative systematic error (Br) and the proportion
of variance explained (EV%) as follows:
" n #
1X m
1X j
Ba ¼ ðyij  y^ij Þ ð2Þ
m j¼1 nj i¼1
973
" n  #
1X m
1X j
yij  y^ij
Br ¼ ð3Þ
m j¼1 nj i¼1 yij
 
u^j þ e^ij
EV% ¼ 100 ð4Þ
uj þ eij
where m is the number of sites, nj is the number of obser-
vations of mass loss or the element content in the needles in
site j, and ŷij is the predicted value of mass loss or element
content, uj and eij are the variances of the random effects
before the parameter estimation of the explanatory variables
in the model and ûj and êij are those variances after parameter
estimation of the explanatory variables.
Results
During the 2-year litterbag incubation, only the contents of
Al, Ni, Cd and Cu did not show significant correlation with
time (Table 3). On average, Fe (Fig. 6) and Pb (Fig. 7)
accumulated significantly, while C (Fig. 1; Table 3), N, P
(Fig. 2; Table 3), B, K, Ca, Mg (Fig. 4; Table 3), Mn and
Zn (Fig. 5; Table 3) were released, although there was high
variation between different sites, tree species and sampling
positions, with both release and accumulation for N, P, Ca,
Mn and Zn. Boron and K were released at the very early
phases of incubation, during the first 6 months, with
80–90 % of K and 50–75 % of B being released from the
needles (Fig. 4). The C/N ratio of the needles decreased
significantly during incubation, and the N/P and C/P ratios
increased (Fig. 3). The average mass loss over both needle
types and all incubation positions was 58 %.
The differences in element accumulation or release
behaviour between the two catchments (Skerdagh/Glenn-
among) were mostly masked behind the large within-
catchment variation, and there were significant differences
only for the C/P and N/P ratios, which were higher for the
needles of the Skerdagh catchment than Glennamong
(Table 3). However, the substantially higher Fe and Mn
contents in peat at Skerdagh (Table 1) were reflected in
that the Fe and Mn dynamics in decaying needles corre-
lated significantly with their contents in peat. Thus, Fe and
Mn were either released less or accumulated more in the
needles of Skerdagh than Glennamong. There was also a
significant time–catchment interaction for Cu (Fig. 7); it
accumulated in the needles of the Skerdagh catchment
during the incubation period, but was mostly released from
the needles of the Glennamong catchment (Table 3).
Overall, it was found that lodgepole pine needles
decomposed quicker than those of Sitka spruce (Fig. 1;
Table 3). Specific element observations revealed that Ni,
Zn, Cd and Al were mostly released from pine needles, but
accumulated in spruce needles (Figs. 6, 7; Table 3) and
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Table 3 Model parameters and performance values of the mixed models constructed for the remaining mass (RM) (%) and the element content remaining (%) during 2 years since installation
of litterbags containing fresh needles of Lodgepole pine and Sitka spruce on two drained peatland sites in Ireland
Variable a Site Harvest Furrow Tree sp. Time ln(Time) (Time)3 (Time)-3 Fe peat Mn peat u e Bias Biasr EV%
Site mg kg-1 mg kg-1

RM 92.0 8.5 3.6 -52.7 0.00 27.9 0.001 -0.010 82.2

C 93.0 8.7 3.5 -51.5 0.00 25.9 0.690 0.003 94.5
N 104.2 19.2 -8.5 -0.9 10.4 213.4 -0.001 -0.022 46.0
Al 84.4 54.8 50.1 9,993.5 0.0004 -0.178 5.9
B 8.0 3.3 90.9 2.7 11.7 0.0002 -0.045 98.8
Ca 138.9 -2.4 -24.8 273.6 528.2 0.004 -0.193 38.1
Cd 87.9 79.2 0.00 3,134.0 0.003 -0.153 14.2
Cu 97.2 62.6 1,952.4 35,083.7 0.0003 -0.349 8.4
Fe -2,104.1 -2,186.6 2,045.5 0.03 0.00 11,616,540.0 2.923 -1.000 24.3
K -3.6 99.8 0.00 92.3 -0.000 -0.102 94.0
Mg 94.8 9.8 -9.2 -58.2 0.00 141.8 -0.0001 -0.040 80.7
Mn 79.5 -5.5 -54.9 0.02 0.00 1,105.6 -0.005 -0.975 46.0
Ni 72.8 38.0 0.00 970.9 -114.3 -1.310 27.2
P 91.3 14.8 -50.2 66.15 149.4 0.0003 -0.038 68.0
Pb -142.7 -330.9 339.5 28,639.4 177,362.0 -0.0003 -0.364 29.1
Zn 103.9 30.2 -21.7 0.00 251.2 -0.0006 -0.074 66.1
C/N 35.8 7.7 -17.8 0.00 14.0 -0.0004 -0.012 83.1
C/P 451.1 -66.6 91.8 18.3 1,387.9 3,642.6 -0.0002 -0.028 38.3
N/P 12.6 -1.7 9.2 0.30 3.3 -0.0000 -0.015 79.0

Eur J Forest Res (2014) 133:969–982

Only the statistically significant (p \ 0.05) parameters are presented. a = intercept, Site = site dummy (Skerdagh = 0, Glennamong = 1), Harvest = clear-cut dummy (clear-cut = 1,
uncut = 0); Furrow = non-furrow dummy (between furrow = 1, inside furrow = 0); Tree sp. = tree species dummy (Lodgepole pine = 0, Sitka spruce = 1); Time = years since the
beginning of the incubation with different transformations (starting year is number 1); u and e are the random effects of the sites and the variation among the measurement time points.
Bias = absolute bias; Biasr = the relative bias; EV% = explained share of the total variance. For random part, the parameters of u were non-significant
Eur J Forest Res (2014) 133:969–982 975

Fig. 1 Changes in mass and Glennamong Skerdagh

carbon content in harvest BF-Spruce-CC
120 IF-Spruce-CC 120
residue needles during 2 years BF-Pine-CC
of incubation as % of initial. BF 100 IF-Pine-CC 100
Between drainage furrows, IF BF-Spruce-UC

Mass remaining (%)

IF-Spruce-UC
inside drainage furrows, CC 80
BF-Pine-UC
80
clear-cut, UC uncut. To help the IF-Pine-UC
interpretation of figure; solid 60 60
line = UC, dotted line = CC,
circle = BF, triangle = IF, 40 40

closed circle/triangle = spruce,

20 20
open circle/triangle = pine
0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0

120 120

100 100
C remaining (%)

80 80

60 60

40 40

20 20

0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0
Time (Year) Time (Year)

Fig. 2 Changes in N and P Glennamong Skerdagh

contents in harvest residue BF-Spruce-CC
needles during 2 years of IF-Spruce-CC
BF-Pine-CC
incubation as % of initial. For 180 IF-Pine-CC 180
the explanation of symbols, see 160 BF-Spruce-UC 160
Fig. 1 IF-Spruce-UC
140
N remaining (%)

BF-Pine-UC 140
120 IF-Pine-UC
120
100 100
80 80
60 60
40 40
20 20
0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0

140 140
120 120
P remaining (%)

100 100
80 80
60 60
40 40
20 20
0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0
Time (Year) Time (Year)

that the C/N and C/P ratios remained higher during
decomposition of Sitka spruce than the lodgepole pine
needles (Fig. 3). In contrast, however, Ca and Mg release
was faster from Sitka spruce than lodgepole pine needles
(Fig. 4; Table 3).
Harvesting resulted in slower decomposition of needles
and also slower release of N and P as compared to the
uncut forest sites (Fig. 1; Table 3). On average, there was
no net change in the N content in the clear-cut sites, while
the needles in the uncut sites lost 30–40 % of their initial N

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976 Eur J Forest Res (2014) 133:969–982

Fig. 3 Changes in C/N, C/P Glennamong Skerdagh

and N/P ratios in harvest residue 60 BF-Spruce-CC 60
needles during 2 years of IF-Spruce-CC
BF-Pine-CC
incubation. For the explanation 50 IF-Pine-CC 50
of symbols, see Fig. 1 BF-Spruce-UC
40 IF-Spruce-UC 40
BF-Pine-UC

C/N
IF-Pine-UC
30 30

20 20

10 10

0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0
1000 1000

800 800

600 600
C/P

400 400

200 200

0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0

30 30

25 25

20 20
N/P

15 15

10 10

5 5

0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0
Time (Year) Time (Year)

content. Conversely, Mn was released significantly faster
from the needles incubated in the clear-cut sites (Fig. 5).
Significant changes were additionally observed for the Pb
content of the needles in the clear-cut site, whereby results
show up to 5–30-fold higher content at the end of the study
than observed initially (Fig. 7).
The litterbag placement (inside furrows/between fur-
rows) had a significant effect on the contents of N, B, Fe,
Mg and Pb (Table 3). During decomposition, the N, Mg
and B contents remained at a significantly higher level in
the needles placed between the furrows, whereas the Fe and
Pb contents were significantly higher inside the furrows.
Discussion
We studied the decomposition of harvest residue needles
and subsequent nutrient and heavy metal transformations in
needles placed in drainage furrows and between furrows in
two clear-cut and two uncut forest sites on Atlantic blanket
peats in west of Ireland. The bags in the clear-cut sites were
positioned in windrow-free areas. Clear-cutting resulted in
slower decomposition of needles: over the 2-year study
period, the needles decomposed significantly slower at the
clear-cut sites (46–55 % mass loss) when compared to the
uncut sites (58–77 % mass loss). During the 2 years, N was
not clearly released and even slightly accumulated in the
needles at clear-cut sites, which is consistent with the
results of earlier studies (Hyvönen et al. 2000; Palviainen
et al. 2004b; Kaila et al. 2012). However, considerable N
release took place from the uncut sites (32–37 % of initial
N content), which has not been reported previously on
peatlands. This high net release of N from the needles at
the uncut sites could be linked to their higher mass loss
when compared to the needles from the clear-cut sites, as
the N concentrations in needles during decomposition (data

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Eur J Forest Res (2014) 133:969–982 977

Fig. 4 Changes in B, K, Ca and Glennamong Skerdagh

Mg contents in harvest residue 120 BF-Spruce-CC 120
needles during 2 years of IF-Spruce-CC
incubation as % of initial. For 100 BF-Pine-CC 100
the explanation of symbols, see IF-Pine-CC

B remaining (%)
80 BF-Spruce-UC 80
Fig. 1
IF-Spruce-UC
60 BF-Pine-UC 60
IF-Pine-UC
40 40

20 20

0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0

120 120

100 100
K remaining (%)

80 80

60 60

40 40

20 20

0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0

200 200
Ca remaining (%)

150 150

100 100

50 50

0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0

120 120

100 100
Mg remaining (%)

80 80

60 60

40 40

20 20

0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0
Time (Year) Time (Year)

not shown) did not indicate differences between uncut and
clear-cut sites. Kaila et al. (2012) presented contrasting
results from drained and fertile Norway spruce-dominated
peatlands in Finland, where clear-cutting did not signifi-
cantly affect the decomposition of spruce and pine needles.
Palviainen et al. (2004b) in their study on a mineral soil
forest reported that clear-cutting decreased the decompo-
sition of spruce needles but not of pine needles.
There could be a number of reasons for the lower mass
loss of needles in the clear-cut sites compared to the uncut
sites. Decomposition of litter and harvest residues is a
complex phenomenon influenced by the activity and
nutrient demand of heterotrophic decomposers. The activ-
ity is regulated by environmental conditions such as soil
temperature, nutrient availability and moisture conditions
(Gosz et al. 1973; Prescott 2005; Laiho 2006). A rise in the
water table level after harvesting may lead to water satu-
ration in parts of the clear-cut area, especially on blanket
peat soils and regions associated with high precipitation.
Water-saturated conditions could decelerate the aerobic

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978 Eur J Forest Res (2014) 133:969–982

Fig. 5 Changes in Zn and Mn Glennamong Skerdagh

contents in harvest residue
BF-Spruce-CC
needles during 2 years of IF-Spruce-CC
incubation as % of initial. For 200 200
BF-Pine-CC
the explanation of symbols, see IF-Pine-CC
Fig. 1 160 160

Mn remaining (%)
BF-Spruce-UC
IF-Spruce-UC
120 BF-Pine-UC 120
IF-Pine-UC

80 80

40 40

0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0

150 150

120 120
Zn remaining (%)

90 90

60 60

30 30

0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0
Time (Year) Time (Year)

Fig. 6 Changes in Al and Fe Glennamong Skerdagh

contents in harvest residue BF-Spruce-CC
needles during 2 years of IF-Spruce-CC
300 1000
BF-Pine-CC
incubation as % of initial. Note IF-Pine-CC
that logarithmic scale is used for 250 BF-Spruce-UC
Al remaining (%)

Skerdagh. For the explanation IF-Spruce-UC

200 BF-Pine-UC
of symbols, see Fig. 1 IF-Pine-UC
150 100

100

50

0 10
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0

1000 100000
Fe remaining (%)

800

10000
600

400
1000

200

0 100
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0
Time (Year) Time (Year)

decomposition of harvest residues. However, Laiho et al. forest sites in Finland. So despite the rise in water level
(2004) and Domisch et al. (2000) observed no clear dif- after harvesting, higher air (Kubin and Kemppainen 1991)
ferences in pine litter decomposition in drained (with low and soil (Huttunen et al. 2003) temperatures and wind
water table) and undrained (with high water table) peatland velocity in the clear-cut sites can lead to drying out of the

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Eur J Forest Res (2014) 133:969–982 979

Fig. 7 Changes in Pb, Cu, Ni Glennamong Skerdagh

and Cd contents in harvest 1200 BF-Spruce-CC 10000
residue needles during 2 years IF-Spruce-CC
of incubation as % of initial. 1000 BF-Pine-CC

Pb remaining (%)
IF-Pine-CC
Note that for Pb and Cu, 800 BF-Spruce-UC
logarithmic scale is used for IF-Spruce-UC
Skerdagh. For the explanation 600 BF-Pine-UC 1000
of symbols, see Fig. 1 IF-Pine-UC
400

200

0 100
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0
400 10000
Cu remaining (%)

300
1000

200

100
100

0 10
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0

300 300

250 250
Ni remaining (%)

200 200

150 150

100 100

50 50

0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0

350 350
Cd remaining (%)

300 300
250 250
200 200
150 150
100 100
50 50
0 0
0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0
Time (Year) Time (Year)

topmost peat layers, especially during dry summer periods,
thus inducing moisture stress among microbial communi-
ties. Peat soils generally have a capillary fringe to move
water to the surface in dry periods. However, if the topmost
humus or peat eventually dries out, the harvest residues can
lose their capillary connection to the subsurface water
(Laurén and Heiskanen 1997; Laurén and Mannerkoski
2001). In contrast, the variation in temperature and mois-
ture in the surface soil under forest canopy is smaller than
treeless peatland areas (Straková et al. 2012), thus main-
taining more optimal moisture conditions for heterotrophic
decomposers.
Apart from the initial accumulation in the needles inside
the furrows of the Skerdagh clear-cut site, P was clearly
released (38–53 % of initial P content) from all sites and
sampling positions. This study therefore supports the
hypothesis presented in previous decomposition studies
(Palviainen et al. 2004b; Kaila et al. 2012) that P release
from decaying harvest residues can be a significant source
of P to receiving water courses, especially in the case of
blanket peats and other peat soils with low P adsorption
capacity (Tamm et al. 1974; Nieminen and Jarva 1996;
Asam et al. 2012). The initial accumulation of P inside the
furrows of the Skerdagh clear-cut site could partly be

123
980
explained by the fact that the needles inside the furrows
were also accumulating Fe and Al, which can act as
sorption sinks for P in furrow waters.
In contrast, negligible release of N from needles on the
harvested sites indicates that N release from harvest resi-
dues may not be an explanation for the enhanced N export
that has been reported to occur from peat soils soon after
harvesting (Cummins and Farrell 2003b; Nieminen 2004).
Thus, increased mineralisation of N from peat could be a
more probable cause for the reported high N exports. It
should be noted, however, that even if harvest residues did
not release N during the early stages of decomposition, the
harvest residue piles can increase N export by enhancing N
mineralisation and release from underlying soil (Rosén and
Lundmark-Thelin 1987).
High release of easily soluble nutrients, for instance B,
K and Mg, in the early phases of decomposition is in
accordance with earlier studies (Palviainen et al. 2004a;
Kaila et al. 2012). However, the release in the presented
study was observed to be faster, possibly reflecting the
effects of climatic variables on nutrient leaching from
harvest residues. Calcium began to be released after about
half a year. Its release was slower, compared to K and Mg,
as it is a structural component of needle litter, and hence,
its release depends on microbial activity rather than direct
leaching from needle cell solution (Blair 1988). Other
studies on mineral soil forests (Palviainen et al. 2004a) and
forestry-drained peatlands (Kaila et al. 2012) have reported
the Ca release to begin after a lag period of approximately
1 year.
Among the different heavy metals, Mn was clearly
released from needles from three of the four sites, with a
net release of [90 % of the initial Mn content over the
2 years. In contrast, it was accumulated at the Skerdagh
uncut site, especially in the pine needles. Other studies on
needle litters have shown mostly release of Mn (Laskowski
and Berg 1993; Kaila et al. 2012). The accumulation of Mn
in the needles of the uncut Skerdagh site is most likely
related to the high Mn contents in peat. Manganese content
in peat was over 200 times higher at the Skerdagh uncut
site compared with the other three study sites (Table 1),
and Mn absorption from soil could explain the high Mn
accumulation in needles. Similarly, higher accumulation of
Al in spruce needles at the Skerdagh uncut site compared
with the other three sites, and significantly higher accu-
mulation of Fe in the furrows of both Skerdagh sites
compared with the Glennamong sites, could be due to high
contents of these elements in the soils of respective sites
(Table 1). However, the mechanisms behind Fe, Al and Mn
transport from soil into needles require further studies. The
contact of metal-rich soil with decomposing needles is one
possible explanation, and microbial translocation of metals
by fungi from underlying soil is another plausible factor
123
Eur J Forest Res (2014) 133:969–982
(Van Nevel et al. 2014 and the references therein). Fungi
may accumulate significant amounts of metals, and as fungi
may constitute a significant pool of decomposing organic
material, it is likely that fungi could influence metal
translocations from soil into litter. In addition, polyvalent
metal ions form stable complexes with humic substances,
and as the amount of humic substances in organic material
increases in the course of decomposition, more metals may
be bound in decomposed needle litters than freshly fallen
needles (Van Nevel et al. 2014).
Copper also clearly accumulated in the needles inside
the furrows of the uncut Skerdagh site, even though Cu
contents in the soil were very low (Table 1). However,
particularly inside the furrows, the accumulation of ele-
ments may be related more to the soil characteristics
upstream of the study site than the in situ soil character-
istics. The litters decaying inside the furrows are in close
contact with the water discharging from the upslope blan-
ket peat area, and after losing their protective epiderm and
cuticle, the litters may easily absorb metals and other ele-
ments from this water (Tyler 1972). Other than the dif-
ferences in soil characteristics, the different accumulation
and release behaviour of elements between Glennamong
and Skerdagh could be due to topographical factors. Gen-
tler-sloping terrain in Skerdagh can increase water resi-
dence time and hence the contact time between needle litter
and elements in through-flowing water.
Zinc, Ni and Cd were mostly accumulated in the spruce
needles, but released from the pine needles, which is in
accordance with the results of Kaila et al. (2012) from
nutrient-rich Norway spruce-dominated peatlands in Fin-
land. Among the different heavy metals, Pb was the only
one showing any difference between the uncut and clear-
cut sites (Fig. 7). Higher accumulation of Pb in the clear-
cut sites is difficult to explain. There were no differences in
peat Pb contents between the study sites, thus different
accumulation from the soil is not a likely explanation. The
present-day atmospheric Pb deposition in Ireland may not
be significantly more than 10–20 g ha-1 year-1 (Schell
et al. 1997), and more atmospheric Pb deposition reaching
the soil surface at clear-cut sites over uncut sites also may
not be a likely mechanism behind different Pb accumula-
tion behaviour. Thus, further research is needed to assess if
the differences in Pb accumulation between the clear-cut
and uncut blanket peat sites, as observed in the present
study, are a general phenomenon, and if so, what could be
the driving mechanism behind higher accumulation in
clear-cut sites.
In conclusion, the study showed that clear-cutting
resulted in the slower decomposition of needles than in
uncut forest sites on Atlantic blanket peats. The tree spe-
cies was a significant factor contributing to nutrient and
heavy metal release and accumulation patterns from
Eur J Forest Res (2014) 133:969–982
needles, as was their position (inside furrow/between fur-
row). The site soil characteristics contributed significantly
to Al, Fe and Mn transformations during decomposition.
The decomposition of needles on blanket peat catchments
may be a significant source of P to receiving water courses,
owing to their fast release of P. However, needles are not a
likely source for N export, or most heavy metals contained
within, following harvest operations. This was the first time
the decomposition of harvest residues was studied on
Atlantic blanket peat catchments, and further studies are
still needed as to the effects of decomposition-contributing
factors not studied here, such as harvest residue piles (i.e.
windrows) or topographic variation, i.e., drier high-gradi-
ent upslope sections or wetter gentler-sloping areas close to
water pathways.
Acknowledgments The authors gratefully acknowledge the funding
(SANIFAC/ForSite) from the Department of Agriculture, Fisheries
and Food in Ireland. We thank the help and support from the Envi-
ronmental Protection Agency (EPA) of Ireland, Coillte Teoranta for
permission to use the sites and the Marine Institute for the use of their
facilities over the duration of the study. We also wish to thank our
colleagues at Finnish Forest Research Institute (FFRI) and NUI
Galway for all their assistance, and finally, the useful comments and
suggestions of three anonymous reviewers.
References
Ahtiainen M, Huttunen P (1999) Long-term effects of forestry
managements on water quality and loading in brooks. Boreal
Environ Res 4:101–114
Anderson AR, Ray D, Pyatt DG (2000) Physical and hydrological
impacts of blanket peat afforestation at Bad a Cheo, Caithness:
the first 5 years. Forestry 73:467–478
Asam Z (2012) Cycling and transport of phosphorus and nitrogen
from harvested peatland forests and possible mitigation methods.
National University of Ireland, Galway. Academic dissertation
Asam Z, Kaila A, Nieminen M, Sarkkola S, O’Driscoll C, O’Connor
M, Sana A, Rodgers M, Xiao L (2012) Assessment of
phosphorus retention efficiency of blanket peat buffer areas
using a laboratory flume approach. Ecol Eng 49:160–169
Asam Z, Nieminen M, O’Driscoll C, O’Connor M, Sarkkola S, Kaila
A, Sana A, Rodgers M, Zhan X, Xiao L (2014) Export of
phosphorus and nitrogen from lodgepole pine (Pinus contorta)
brash windrows on harvested blanket peat forests. Ecol Eng
64:161–170
Berg B, Staaf H (1980) Decomposition rate and chemical changes of
scots pine needle litter. II. Influence of chemical composition.
Ecol Bull 32:373–390
Blair J (1988) Nutrient release from decomposing foliar litter of three
tree species with special reference to calcium, magnesium and
potassium dynamics. Plant Soil 110:49–55
Cummins T, Farrell EP (2003a) Biogeochemical impacts of clearf-
elling and reforestation on blanket peatland streams I. Phospho-
rus. For Ecol Manag 180:545–555
Cummins T, Farrell EP (2003b) Biogeochemical impacts of clearf-
elling and reforestation on blanket-peatland streams II. Major
ions and dissolved organic carbon. For Ecol Manag 180:557–570
Domisch T, Finer L, Laiho R, Karsisto M, Laine J (2000)
Decomposition of Scots pine litter and the fate of released
981
carbon in pristine and drained pine mires. Soil Bio Biochem
32:1571–1580
Gosz JR, Likens GE, Bormann FH (1973) Nutrient release from
decomposing leaf and branch litter in the Hubbard Brook Forest,
New Hampshire. Ecol Monogr 43:173–191
Huttunen JT, Nykänen H, Martikainen PJ, Nieminen M (2003) Fluxes
of nitrous oxide and methane from drained peatlands following
forest clear-felling in southern Finland. Plant Soil 255:457–462
Hyvönen R, Olsson BA, Lundkvist H, Staaf H (2000) Decomposition
and nutrient release from Picea abies (L.) Karst. and Pinus
sylvestris L. logging residues. For Ecol Manag 126:97–112
Kaila A, Asam Z, Sarkkola S, Xiao L, Laurén A, Vasander H,
Nieminen M (2012) Decomposition of harvest residue needles
on peatlands drained for forestry: implications for nutrient and
heavy metal dynamics. For Ecol Manag 277:141–149
Kubin E, Kemppainen L (1991) Effect of clear cutting of boreal
spruce forest on air and soil temperature conditions. Acta For
Fennica 225:1–42
Laiho R (2006) Decomposition in peatlands: reconciling seemingly
contrasting results on the impacts of lowered water levels. Soil
Biol Biochem 38:2011–2024
Laiho R, Laine J, Trettin CC, Finér L (2004) Scots pine litter
decomposition along drainage succession and soil nutrient
gradients in peatland forests, and the effects of inter-annual
weather variation. Soil Bio Biochem 36:1095–1109
Laskowski R, Berg B (1993) Dynamics of some mineral nutrients and
heavy metals in decomposing forest litter. Scand J For Res
8:446–456
Laskowski R, Niklinska M, Maryanski M (1995) The dynamics of
chemical elements in forest litter. Ecology 76:1393–1406
Laurén A, Mannerkoski H (2001) Hydraulic properties of mor layers
in Finland. Scand J For Res 16:249–441
Laurén A, Heiskanen J (1997) Physical properties of the mor layer in
a Scots pine stand I. Hydraulic conductivity. Can J Soil Sci
77:627–634
Ledger DC, Harper SE (1987) The hydrology of a drained, afforested
peat bog in southern Scotland, 1977–1986. Trans R Soc Edinb
Earth Sci 78:297–303
Lehto T, Aphalo P, Saranpää P, Laakso T, Smolander A (2010)
Decomposition and element concentrations of Norway spruce
needle litter with differing B, N, or P status. Plant Soil
330:225–238
Lundin L (1998) Alternative peatland forestry: impacts on hydrology
and surface water chemistry. In: Proceedings of the international
peat symposium, the spirit of peatlands, 7–9 September 1998,
Jyväskylä, Finland, pp 76–78
Lundin L (1999) Effects on hydrology and surface water chemistry of
regeneration cuttings in peatland forests. Int Peat J 9:118–126
Lundin L (2000) Water environment care at peatland forestry
practices. In: Rochefort L, Daigle J-Y (eds) Sustaining our
peatlands. Proceedings of the 11th international peat congress,
Quebec City, Canada August 6–12, 2000, Vol II, pp 952–961
Lundmark-Thelin A, Johansson M (1997) Influence of mechanical
site preparation on decomposition and nutrient dynamics of
Norway spruce (Picea abies (L.) Karst.) needle litter and slash
needles. For Ecol Manag 96:101–110
Moore TR, Trofymow JA, Siltanen M, Prescott C, CIDET Working
Group (2005) Patterns of decomposition and carbon, nitrogen,
and phosphorus dynamics of litter in upland forest and peatland
sites in central Canada. Can J For Res 35:133–142
Nieminen M (1998) Changes in nitrogen cycling following the
clearcutting of drained peatland forests in southern Finland.
Boreal Environ Res 3:9–21
Nieminen M (2003) Effects of clear-cutting and site preparation on
water quality from a drained Scots pine mire in southern Finland.
Boreal Environ Res 8:53–59
123
982
Nieminen M (2004) Export of dissolved organic carbon, nitrogen and
phosphorus following clear-cutting of three Norway spruce
forests growing on drained peatlands in southern Finland. Silva
Fennica 38(2):123–132
Nieminen M, Jarva M (1996) Phosphorus adsorption by peat from
drained mires in southern Finland. Scand J For Res
11(4):321–326
Nilsson I (1972) Accumulation of metals in spruce needles and needle
litter. Oikos 23:132–136
Nisbet T, Dutch J, Moffat A (1997) Whole-tree harvesting: a guide to
good practice. Forestry Commission Practice Guide, Forestry
Commission, Edinburgh
O’Driscoll C, De Eyto E, O’Connor M, Asam Z, Rodgers M, Xiao L
(2013) Biotic response to forest harvesting in acidic blanket peat
fed streams: a case study from Ireland. For Ecol Manag
310:729–739
O’Driscoll C, De Eyto E, Rodgers M, O’Connor M, Asam Z, Xiao L
(2012) Diatom assemblages and their associated environmental
factors in upland peatforest rivers. Ecol Indic 18:443–451
O’Driscoll C, Rodgers M, O’Connor M, Asam Z, De Eyto E, Poole R,
Xiao L (2011) A potential solution to mitigate phosphorus
release following clearfelling in peatland forest catchments.
Water Air Soil Pollut 221:1–11
Palviainen M, Finér L (2012) Estimation of nutrient removals in stem-
only and whole-tree harvesting of Scots pine, Norway spruce,
and birch stands with generalized nutrient equations. Eur J For
Res 131:945–964
Palviainen M, Finér L, Kurka A-M, Mannerkoski H, Piirainen S, Starr
M (2004a) Release of potassium, calcium, iron and aluminium
from Norway spruce, Scots pine and silver birch logging
residues. Plant Soil 259:123–136
Palviainen M, Finér L, Kurka A-M, Mannerkoski H, Piirainen S, Starr
M (2004b) Decomposition and nutrient release from logging
residues after clear-cutting of mixed boreal forest. Plant Soil
263:53–67
Prescott CE (2005) Decomposition and mineralization of nutrients
from litter and humus. In: BassiriRad H (ed) Nutrient acquisition
by plants. Springer, Berlin, Heidelberg, pp 15–41
Rasbash J, Browne W, Goldstein H, Yang M, Plewis I, Healy M,
Woodhouse G, Draper D, Langford I, Lewis T (2001) A user’s
guide to MlwiN. Centre for Multilevel Modeling, Institute of
Education, University of London, London
123
Eur J Forest Res (2014) 133:969–982
Rodgers M, O’Connor M, Healy MG, O’Driscoll C, Asam Z,
Nieminen M, Poole R, Muller M, Xiao L (2010) Phosphorus
release from forest harvesting on an upland blanket peat
catchment. For Ecol Manag 260(12):2241–2248
Rosén K, Lundmark-Thelin A (1987) Increased nitrogen leaching
under piles of slash: a consequence of modern forest harvesting
techniques. Scand J For Res 2:21–29
Rustad L, Cronan C (1988) Element loss and retention during litter
decay in a red spruce stand in Maine. Can J For Res 18:947–953
Rustad LE (1994) Element dynamics along a decay continuum in a
red spruce ecosystem in Maine, USA. Ecology 75:867–879
Sarkkola S, Koivusalo H, Laurén A, Kortelainen P, Mattsson T,
Palviainen M, Piirainen S, Starr M, Finér L (2009) Trends in
hydrometeorological conditions and stream water organic carbon
in boreal forested catchments. Sci Total Environ 408:92–101
Scheid S, Gunthard-Goerd MS, Schulin R, Nowack B (2009)
Accumulation and solubility of metals during leaf litter decom-
position in non-polluted and polluted soil. Eur J Soil Sci
60:613–621
Schell WR, Tobin MJ, Novak MJV, Wieder RK, Mitchell PI (1997)
Deposition history of trace metals and fallout radionuclides in
wetland ecosystems using 210Pb chronology. Water Air Soil
Pollut 100:233–239
Straková P, Penttilä T, Laina J, Laiho R (2012) Disentangling direct
and indirect effects of water table drawdown on above- and
belowground plant litter decomposition: consequences for accu-
mulation of organic matter in boreal peatlands. Glob Change
Biol 18:322–335
Tamm CO, Holman H, Popovic B, Wiklander G (1974) Leaching of
plant nutrients from soils as a consequence of forest operations.
Ambio 3:221–222
Tyler G (1972) Heavy metals pollute nature, may reduce productivity.
Ambio 1:52–59
Van Nevel L, Mertens J, Demey A, De Schrijver A, De Neve S, Tack
FMG, Verheyen K (2014) Metal and nutrient dynamics in
decomposing tree litter on a metal contaminated site. Environ
Pollut 189:54–62
Vesterdal LL (1999) Influence of soil type on mass loss and nutrient
release from decomposing foliage litter of beech and Norway
spruce. Can J For Res 29:95–105