全 文 ：Journal of Forestry Research (2010) 21(2): 119−128
DOI 10.1007/s11676-010-0020-4
Stem flow chemistry of Picea glehnii, Abies sachalinensis and Alnus
japonica and its effect on the peat pore water chemistry in an
ombrogenous mire in Ochiishi, eastern Hokkaido, Japan
Tsutomu Iyobe • Akira Haraguchi
Received: 2009-08-05; Accepted: 2009-12-22
© Northeast Forestry University and Springer-Verlag Berlin Heidelberg 2010
Abstract: We investigated the chemical properties of stemflow of Picea
glehnii, Abies sachalinensis and Alnus japonica as well as peat pore
water chemistry, including the distance and depth profiles of pore water
chemistry, in an ombrogenous mire. The effect of stemflow on the peat
pore water chemistry was clear at the stem base in the peat forest in the
mire, and the peat pore water around the stem base of a tree had its own
chemical properties specific to each species. P. glehnii showed the
highest concentration of salts both in stemflow and peat-pore water,
whereas A. japonica showed the lowest concentrations; however, the
gradient of the chemical environment from the stem base to outside of
the canopy is formed. The peat pore water chemistry under the canopy
was mainly controlled by the chemical processes diluted by the abundant
peat pore water; the stemflow movement in the high water content of the
peat was more slowly because of the flat topography (< 1º). This would
be due to the fact that the chemicals in stemflow would be diluted by the
abundant peat pore water. The spatial heterogeneity of chemical
environment between microsites within forested peatland would be also
contributed indirectly through the control of microorganism activity, and
nutrient regeneration mediated the surface water and the stemflow of the
dominant canopy trees.
Foundation project: This work was partly funded by the Grant in Aid
from JSPS.
The online version is available at http://www.springerlink.com
Tsutomu Iyobe ( )
Research Center for Natural Hazards & Disaster Recovery, Niigata
University, Ikarashi 2, Nishi-ku, Niigata 950-2181, Japan.
E-mail: iyotsuto@hotmail.com;
Tel.:+81-25-262-7051; Fax: +81-25-262-7050
Akira Haraguchi
Faculty of Environmental Engineering, The University of Kitakyushu,
Hibikino 1-1, Wakamatsu, Kitakyushu 808-0135, Japan
Responsible editor: Chai Ruihai
Keywords: Abies sachalinensis; Alnus japonica; Picea glehnii; peat pore
water; stemflow; chemical properties
Introduction
In most forest ecosystems, atmospheric precipitation is chemi-
cally modified by the canopy of trees (Mahendrappa 1974;
Crockford et al. 1996; Chiwa et al. 2003) and the precipitated
solution intercepted by the forest canopy affects the soil chemi-
cal properties in the forest (Bergkvist and Folkeson 1995; Butler
and Likens 1995). Many research efforts have clarified that pre-
cipitation is modified by the tree canopies by washing out the
deposited substances on the canopy, ionic exchange, or chemical
leaching from the leaves (Cappellato and Peters 1995; Houle et
al. 1999; Piirainen et al. 2002). Some chemicals deposited on the
canopies from the atmosphere are washed out by the precipitated
water and these chemicals are enriched in the precipitation
through the canopy. The N components (NO3- + NH4+) in the
precipitation are absorbed by the canopy, and the concentration
of nutrients is usually depressed when precipitation is intercepted
by the canopy (Lovett and Lindberg 1993); however, in the
senescent season, K+ is leached from the canopy to the
precipitation and the nutrients are enriched while precipitated
water is being intercepted by the canopy (Lovett and Lindberg
1993; Houle et al. 1999). The interaction between precipitated
water and the tree canopy is specific for each tree species (Kaul
and Billings 1965; Tajchman et al. 1991; Neary and Gizyn 1994;
Gordon et al. 2000). Chemically modified precipitated water
reaches the soil surface as throughfall and stemflow and affects
the chemical properties of soil. Spatial differences in the
precipitated water chemistry and the flux of precipitated water
lead to the spatial variability of soil chemical environments in the
forest (Potts 1978; Duijsings et al. 1986; Dämmgen and
Zimmerling 2002). The precipitated water also affects the
vegetation pattern of the forest floor (Clement and Wittig 1987)
and soil fauna (Stöckli 1991; Kaneko and Kofuji 2000).
In forest stands, trees can increase the heterogeneity of the
RESEARCH PAPER
Journal of Forestry Research (2010) 21(2): 119−128
120
rainwater input process. When precipitation is intercepted by the
canopy, it is partitioned into throughfall and stemflow compo-
nents as diffuse and point inputs, respectively (Chang and
Matzner, 2000; Liang et al. 2009). Trees can also increase the
heterogeneity of the rainwater infiltration process. Stemflow
water flows into soil at the base of tree stems, hence the effects
of stemflow extend to a much more restricted area than the ef-
fects of throughfall.
Previous studies of soil water dynamics in upland forested
stands have paid little attention to the effect of stemflow, due to
the low ratio of stemflow to total precipitation. However, the
point inflow of stemflow may have major implications for soils
water dynamics. Beven and Germann (1982) indicated that
macropores in soil may be associated with either living or
decayed tree roots and that the structure of macropore systems
derived from roots may be very effective at channeling water
through the soil. Root-induced channels are preferential flow
pathways, and stemflow tends to follow the channels into the soil
(Martinez-Meza and Whitford 1996; Voigt 1960). Liang et al.
(2009) observed that stemflow rapidly moved into soils layers
along pathways around roots. This suggested that stemflow not
only serves as a point source of rainwater on the forest floor, but
also has a high potential to infiltrate multiple soil layers.
In terrestrial ecosystems, stemflow modifies the chemical
properties of soil at the tree base (Jochheim 1984; Förster and
Schimmack 1992). Stemflow is enriched with salts or organic
substances while flowing through the canopy and stems and
carries these chemicals into soils at the stem base. These
substances diffuse or flow from the stem base to the surrounding
soil. In particular, the acid deposition is concentrated in the
stemflow and highly acidified stemflow affects the soil chemistry
(Falkengren-Grerup and Björk 1991; Joslin and Wolfe 1992; Bini
and Bresolin 1998; Kaneko and Kofuji 2000; Matschonat and
Falkengren-Grerup 2000). Many researchers have shown that
stemflow strongly affects the soil chemistry at the stem base and
that a clear gradient of chemical properties appears around the
stem base (Gersper and Holowaychuk 1971; Chang and Matzner
2000). Some researchers have also analyzed the process of
matter flux from the atmosphere to a limnological system via
vegetation and soil (Stevens et al. 1989; Neal et al. 2003).
Peat bogs support a mosaic vegetation types that is largely
determined by differences in water level and nutrient content.
Some area is open and dominated by Sphagnum spp. whereas
others may by forested with single or several tree species
(Ahmad-Shah and Rieley 1989; Haraguchi et al. 2003; Iyobe and
Haraguchi 2008). Stemflow might also induce spatial
heterogeneity of soil solution and of water and element fluxes in
the mire soil. The spatial heterogeneity of soil solution chemistry
and water fluxes caused by stemflow is likely to be influencing
stand-level elemental budgeting.
The evergreen needle-leaved tree P. glehnii Masters and the
deciduous broad-leaved tree A. japonica form wetland forests in
Hokkaido, northern Japan (Tomizawa et al. 1997; Hotes et al.
2001; Haraguchi et al. 2003; Tsuyuzaki and Haraguchi 2009).
The two species have contrast and specific characteristics, i.e., P.
glehnii produce acidic soil environment (Haraguchi et al. 2003)
and A. japonica is nitrogen-fixing plant. Abies sachalinensis
Masters, evergreen needle-leaved, is also abundant spices in
Hokkaido. At a P. glehnii - A. sachalinensis forests on coastal
sand dune in eastern Hokkaido, Nishijima et al. (2003) reported
the boundary between the P. glehnii and A. sachalinensis forests
is determined by the ground-water table depth and the pH of the
soil pore water with groundwater dynamics caused by the dif-
ference of microtopography. If these forest stands established on
the saturated soil on the flat topography, effects of chemicals in
stem flow on soil chemical properties under the tree canopy
would be dependent on the rate of diffusion and flux within the
inundated soil. However, research on the precipitation chemistry
in wetland forests is not so extensive, and information on the
effects of stem flow on the inundated soil chemical environment
is scarce.
In this study, we focused on the effects of stem flow on the
soil chemical environment in a boreal forested peatland. We
determined the chemical properties of stemflow of three common
species as well as peat pore water chemistry, including the
distance and depth profiles of pore water chemistry. The
objectives of this study were (1) to evaluate how stemflow
chemistry affects the soil chemical environment and (2) to
evaluate the species differences in modification of peat chemistry
by stemflow.
Study area and methods
Study area
The study area was in Ochiishi district (43°10–13 N, 145°
28–31 E), Nemuro City, eastern Hokkaido, northern Japan. In
the Ochiishi district, there are some peat mires (5–65 ha) on the
Upper Pleistocene coastal terrace, ca. 50 m a.s.l. More
specifically, the study was carried out at the Cape Ochiishi Mire.
Cape Ochiishi Mire (61.3 ha) is on the Nemuro peninsula and
faces the coastline from a distance of 0.4 km or less. The annual
temperature, rainfall, and humidity from 1961 to 1990 in the
district averaged 5.9°C, 1035 mm, and 80%, respectively
(reports from the meteorological observatory in Nemuro City).
From late spring to mid summer, this area is covered with sea fog
carried by wind originating in the south above the Pacific Ocean.
The fog formation process in this region is described in detail by
Hori (1953).
The peat depth is ca. 2.0–3.0 m in Cape Ochiishi Mire (Iyobe
et al., unpublished data). Volcanic ash has fallen on these mires
several times. In these mires, the uppermost tephra (aerially
transported volcanic ejecta) layer, which was easily detectable in
the cores between a peat stratigraphic depth of 10 to 15 cm,
corresponds with the Me-a (Meakan tephra layer: ca. 500 year
BP, Iyobe and Haraguchi 2005). Cape Ochiishi Mire is on a cape
connected by low-lying land with the mainland and is almost
completely surrounded by the Pacific Ocean.
The dominant plant species in Cape Ochiishi Mire are Sphag-
num spp. and Picea glehnii (Fr. Schm.) Masters, Abies sachalin-
ensis (Fr. Schm.) Masters, Alnus japnica (Thunb.) Steud. The P.
glehni trees grow on the margins of the Sphagnum-dominated
Journal of Forestry Research (2010) 21(2): 119−128
121
mires. The peat surface exhibits a well-developed
hummock-hollow microtopography. The P. glehnii trees in the
transition zone are stunted and distorted. The surrounding area in
Cape Ochiishi Mire is dominated by the A. japnica trees. These
forest canopies are almost monospecific. A. sachalinensis is
established between the P. glehnii and A. japonica trees zone,
which is mixed with P. glehnii trees. The Sphagnum mire is
dominated by Sphagnum fuscum (Schimp.) Klinggr., Empetrum
nigrum L. var. japonicum K. Koch, and Vaccinium oxycoccus L.
In the forest, Lysichiton camtschatcense (L.) Schott, Ledum
palustre L. spp. diversipilosum (Nakai) Hara and Sphagnum spp.
(mainly S. girgensohnii Russ. and S. squarrosum Crome.)
dominate the understory (Haraguchi 1996).
Stemflow chemistry
Stemflow collectors were made of polyethylene tubing (2-m long
and 1-cm diameter) cut longitudinally in half. The tubes
encircled the lower bole of the tree trunks. Stemflow was
collected in 5-L polyethylene bottles. Two individuals (6 to 7 m
high and 20 to 25 cm d.b.h.) of three species, P. glehnii, A.
sachalinensis and A. japonica were selected in the Cape Ochiishi
mire and stemflow collectors were placed on the 6 individuals (3
species × 2 individuals) on 5 June 2001. Stemflow samples were
collected at biweekly from June 15 to December 1, 2001 (total:
13 times). Stemflow samples were transferred to 100 mL
polyethylene bottles after the volume of stemflow was measured
in the field. Samples were stored in dark, cool conditions before
analyses. Electrical conductivity (EC) and pH were measured in the
laboratory within 24 h after sampling with portable EC and pH
meters (TOA Co., Tokyo, Japan). The values for EC were not
corrected for pH. Samples for ion analysis were filtered through
a nitrocellulose membrane filter (0.20 µm, Advantec Toyo Co.,
Tokyo, Japan type DISMIC-25CS) within 24 h after sampling
and stored in a freezer (–18 °C) to reduce microbial activity. The
major cations (NH4+, Na+, K+, Mg2+, Ca2+) and major anions (Cl–,
NO3–, NO2–, SO42–, PO43–) were then analyzed by an ion
chromatograph (Japan Dionex, Tokyo, Japan, model DX-500).
Chemistry of peat pore water
Peat pore water was collected from collectors made of
1–cm–diameter polyethylene pipe fitted with a porous ceramic
cup. The sample collectors were inserted vertically to just below
the ground surface (0 cm depth), 30 cm depth and 60 cm depth at
points just beside the ree stem (N site) and 50 cm from the tree
stem (F site) for each of the individuals with a stemflow collector
(3 species × 2 distances × 3 depths × 2 individuals). We placed
other collectors for peat pore water sampling within the Sphag-
num–dominated community (open site) at just below the ground
surface (0 cm depth), 30 cm depth and 60 cm depth (3 depths × 2
individuals). The water was collected by a suction pump that
reduced pressure to 60 mm Hg in the pipe. To avoid contamina-
tion with stagnant water in the collector, the first 100-ml sample
was discarded, and the subsequent 100-ml sample was collected
in a polyethylene bottle. Samples were stored in dark cool condi-
tions before analysis. Water samplings and chemical measure-
ments of the peat pore water were made biweekly from July 15
to November 22, 2001 (total: 18 times).
EC and pH were measured in the laboratory within 24 h after
sample collection. Concentrations of the major cations (NH4+,
Na+, K+, Mg2+, Ca2+) and major anions (Cl–, NO3–, NO2–, SO42–,
PO43–) were determined as for the stemflow analysis after
filtering through a nitrocellulose membrane filter (0.20 µm).
Statistical analysis
The Wilcoxon signed-rank test was performed on the stemflow
chemical data to evaluate the species differences. Differences in
peat pore water chemistry among three species at each depth
were also tested using the Wilcoxon signed-rank test.
Differences among the three depths of peat pore water for each
species were evaluated using the Friedman test. Principal
component analysis (PCA) by correlation matrix was performed
on the averaged peat chemical data to show the similarities of
chemical properties among various sampling points and water
types, including stemflow and peat pore water.
Results
Stemflow chemistry
The water quantity in stemflow did not show any significant
difference among the three species (Table 1). Proton (H+), Mg2+
and Ca2+ concentrations in stemflow showed significant
differences among the three species in a decreasing order: P.
glehnii > A. sacharinensis > A. japonica. Electrical conductivity
(EC), Na+, K+, Cl– and SO42– concentrations in the stemflow of
conifers (P. glehnii and A. sacharinensis) were significantly
higher than that of A. japonica. PO43– concentration of the
stemflow of A. sacharinensis was significantly higher than was
the case with the other two species. NH4+ and NO3–
concentrations in the stemflow did not show significant
differences among the species. NO2- was always below the
detection limit (0.01 µmolc•L-1).
Species difference of peat pore water chemistry
At the surface of the peat (0 cm depth), EC at the stem base of P.
glehnii was significantly higher than in the case of A. japonica
(Fig. 1). H+ concentration of the peat pore water at 0 cm depth at
the stem base of the conifers was significantly higher than that of
A. japonica. NH4+, Mg2+ and Cl– concentrations at the 0 cm
depth at the stem base of P. glehnii were significantly higher than
were those of the other two species. Na+, K+, Ca2+, NO3–, PO43–
and SO42– concentrations of the peat surface at the stem base did
not show significant differences among the three species. NO2-
was always below the detection limit (0.01 µmolc•L-1).
EC and the Cl– concentration at the 30 cm depth at the stem
base showed a significant decreasing order: P. glehnii > A. sa-
charinensis > A. japonica (Fig. 1). Na+ concentration at the 30
cm depth at the stem base of the conifers was significantly higher
Journal of Forestry Research (2010) 21(2): 119−128
122
than that of A. japonica. NH4+, Mg2+ and SO42– concentrations at
the 30 cm depth at the stem base of P. glehnii were significantly
higher than were those of the other two species. Ca2+ concentra-
tion at the 30 cm depth at the stem base of P. glehnii was signifi-
cantly higher than in the case of A. sacharinensis. K+ concentra-
tion at the 30 cm depth at the stem base of A. sacharinensis was
significantly higher than for the other two species. PO43– concen-
tration at the 30 cm depth at the stem base of P. glehnii was sig-
nificantly lower than in the case of A. japonica. H+ and NO3–
concentrations at 30 cm depth at the stem base did not show
significant differences among the three species.
Table 1. Chemical property (mean and SD) of stem flow of Picea glehnii (PG), Abies sacharinensis (AS) and Alnus japonica (AJ) in Cape Ochiishi
Mire, eastern Hokkaido, Japan.. Data were collected 13 times from 15 July to 1 December 2001 at regular intervals. Significance level; ** p < 0.01, * p < 0.05,
NS not significant p > 0.05. SD: standard deviation, EC: electric conductivity
Chemical property
Water EC H+ NH4+ Na+ K+ Mg2+ Ca2+ Cl- NO3- PO43- SO42-
Species
mL ·interval-1 mS·m-1 molc·L-1 molc ·L-1 molc ·L-1 molc ·L-1 m olc·L-1 mol c·L-1 mol c·L-1 mol c·L-1 mol c·L-1 mol c·L-1
P. glehnii Mean+ 3695± 31.8± 72.6± 10.74± 1471.36± 284.34± 431.4± 458.98± 2608.15± 0.17± 0.06± 298.08±
SD 6634 15.62 44.4 14.16 810.96 126.6 285.74 300.51 1670.51 0.38 0.21 175.52
A. sacharinensi Mean± 2624± 33.03± 28.85± 17.01± 1054.84± 235.06± 201.17± 163.88± 1630.57± 0.06± 47.7± 194.1±
s SD 3924 22.95 23.82 30.02 708.12 155.47 188.6 129.21 1279.89 0.13 56.49 162.69
A.. japonica Mean± 2373± 7.25± 21.45± 14.21± 375.32± 48.94± 79.22± 44.32± 515.93± D.L.* D.L. 53.52±
SD 1627 5.8 9.83 9.83 313.7 33.97 76.8 37.5 514.56 41.62
PG - AS NS NS ** NS NS NS * ** NS NS * NS
PG - AJ NS ** ** NS ** ** ** ** ** NS NS **
Significance
level
(Wilcoxson
signed-rank
test) AS - AJ NS * * NS ** ** ** ** ** NS * **
*: below the detection limit (0.01 µmolc·L-1)
EC and the NH4+, Mg2+ and Cl– concentrations at the 60 cm
depth at the stem base showed a significant decreasing order: P.
glehnii > A. sacharinensis > A. japonica (Fig. 1). Na+ and Ca2+
concentrations at the 60 cm depth at the stem base of the conifers
were significantly higher than those of A. japonica. H+ and SO42–
concentrations at the 60 cm depth at the stem base of P. glehnii
were significantly higher than with the other two species. NO3–
concentration at the 60 cm depth at the stem base of P. glehnii
was significantly higher than in the case of A. sacharinensis.
PO43– concentration at the 60 cm depth at the stem base of A.
sacharinensis was significantly lower than that for A. japonica.
K+ concentrations at 60 cm depth at the stem base did not show
significant differences among the three species.
Vertical profile of peat pore water chemistry
Electrical conductivity (EC) of the peat pore water significantly
increased from the top to the 60 cm depth for all the three species
(Fig. 1). H+ concentration significantly decreased from the top (0
cm) to the 60 cm depth for P. glehnii, A. sacharinensis and A.
japonica.
In the vicinity of the two coniferous species P. glehnii and A.
sachalinensis, NH4+, Na+, Mg2+ and Ca2+ concentrations signifi-
cantly increased from the top to the 60 cm depth, whereas K+ did
not show significant difference among depths for these two spe-
cies (Fig. 1). In the vicinity of A. japonica, Na+ and Mg2+
showed significant differences among depths and the concentra-
tion at 30 cm depth was the minimum. Ca2+ also showed signifi-
cant differences among depths for A. japonica and the concentra-
tion at 30 cm depth was the maximum. NH4+ and K+ did not
show significant differences among depths in the vicinity of A.
japonica.
Among anions, Cl– showed significant differences between
depths for the three species, and the Cl– concentration showed a
tendency to increase from the top to the 60 cm depth, with the
exception of A. japonica, which showed a minimum at 30 cm
depth. PO43– concentration showed significant differences among
depths only for A. japonica, showing a maximum concentration
at 30 cm depth (Fig. 1). NO3– and SO42– concentration did not
show significant differences among depths for the three species.
Difference of peat pore water chemistry between under canopy
and open site
EC and H+, Na+, K+, Mg2+, Ca2+ concentrations of the peat pore
water at the stem base (either or both the N and F sites) were
significantly higher than at the corresponding depth in the open
sites, for all species and for all depths of the peat (Table 2). NH4+
concentrations at the stem base (either or both the N and F sites)
of P. glehnii were significantly higher than results at the corre-
sponding depth in the open sites for all depths of the peat,
whereas the difference was significant at 30 cm depth for A.
sacharinensis (stem base > open site). The concentration of NH4+
at the stem base of A. japonica (either or both the N and F sites)
showed significant difference from the open site at all the depths;
however, the difference did not show a consistent trend between
open sites and stem base. Cl– concentrations at the stem base
(either or both the N and F sites) were significantly higher than at
the open site for P. glehnii at 30 cm and 60 cm depths, for A.
sacharinensis at 60 cm, and for A. japonica for all depths. NO3–
concentration did not show significant difference between the
stem base and the open site, for all species and all depths. PO43–
Journal of Forestry Research (2010) 21(2): 119−128
123
concentrations at the stem base (either or both the N and F sites)
of A. japonica were significantly higher than at the open site at
30 cm and 60 cm depths. SO42– concentrations at the stem base
were significantly higher than at the open site, except for A. sa-
charinensis at 0 cm depth (open site > stem base) and for A. ja-
ponica at 0 cm (open site > stem base) and 30 cm depths.
Peat pore water chemistry along the distance from the tree stem
base
Differences of peat pore water chemistry between stem base (N
site) and 50 cm from the stem base (F site) of P. glehnii showed
significant differences in Ca2+ at 0 cm depth (N>F), in SO42– at
30 cm and 60 cm depths (N
at 60 cm depth (Table 2). For A. japonica, EC of the peat pore
water showed significant differences between N and F sites
(N>F) at 0, 30 and 60 cm depths. Peat pore water showed sig-
nificant differences between N and F sites in Cl– (N
Ordination by the principal component analysis (PCA)
Eigenvalues of the first and second principal components calcu-
lated using a correlation matrix were 5.41 and 1.57, respectively,
and their proportions were 60.1 % and 17.5 %, respectively.
Stemflow chemistry of the three species was distributed on the
first principal component axis (Fig. 2a). Stemflow of P. glehnii
had the highest and that of A. japonica had the lowest score for
the first principal component. Peat pore water chemistry in the
vicinity of the three species was distributed on the first principal
component axis, and higher scores resulted than that at the open
site. Among the peat pore water results near the three species, P.
glehnii had the highest and A. japonica had the lowest score on
the first principal component axis. Although there was not dis-
cernible difference at various depths for the peat pore water in
the open site, the peat pore water at the three depths in close
proximity to the three species distributed on the second principal
component axis. Peat pore water at the peat surface had the low-
est and that at the 60 cm depth had the highest score on the sec-
ond principal component axis.
H+
0 20 40 60 80 100
µmolc L-1
Picea***
Abies***
Alnus**
0
30
60
D
ep
th
(
cm
) a
b
a
a
a
a
a
b
b
EC
0 10 20 30
mS m-1
Picea***
Abies**
Alnus***
0
30
60
D
ep
th
(
cm
)
c
a
ab
a
a
b
b
b
c
NH4+
0 20 40 60 80 100
mmolc L-1
0
30
60
D
ep
th
(
cm
) a
a
b
b
b
b
b
c
a Picea*
Abies**
AlnusNS
Mg2+
0 200 400 600
mmolc L-1
Picea***
Abies***
Alnus**
0
30
60
D
ep
th
(
cm
)
c
b
b
bb
b
a
a
a
Ca2+
0 50 100 150 200
mmolc L-1
Picea***
Abies***
Alnus*
0
30
60
D
ep
th
(
cm
)
a
a
a
a
ab
a
a
b
b
0
30
60
D
ep
th
(
cm
)
Na+
0 500 1000 1500 2000
mmolc L-1
a
b
a
a
a
a
a
a
b
Picea***
Abies***
Alnus**
K+
0 20 40 60 80
mmolc L-1
0
30
60
D
ep
th
(
cm
)
a
a
a
a
a
a
a
a
b
PiceaNS
AbiesNS
AlnusNS
PO43-
0 5 10 15
mmolc L-1
PiceaNS
AbiesNS
Alnus**
0
30
60
D
ep
th
(
cm
) a a
a
a
ab
ab
a
b
b
SO42-
0 50 100 150 200
mmolc L-1
PiceaNS
AbiesNS
AlnusNS
0
30
60
D
ep
th
(
cm
)
b
a
a
a
a
a
b
b
b
NO3-
0 0.2 0.4 0.6 0.8
mmolc L-1
0
30
60
D
ep
th
(
cm
) a
ab
a
a
a
a
a
a
b Picea
NS
AbiesNS
AlnusNS
Cl-
0 1000 2000 3000
mmolc L-1
Picea***
Abies***
Alnus***
0
30
60
D
ep
th
(
cm
) a
b
b
a
bc
a
b
c
D
ep
th
(
cm
)
D
ep
th
(
cm
)
D
ep
th
(
cm
)
D
ep
th
(
cm
)
D
ep
th
(
cm
)
D
ep
th
(
cm
)
D
ep
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Fig. 1 Means (bars) and SD (error bars) of the electric conductivity (EC), ion concentrations of proton, ammonium, sodium, magnesium,
potassium, calcium, chloride ion concentrations, phosphate, nitrate and sulfate in peat pore water at surface (0 cm), 30 cm and 60 cm depths
from the peat surface around Picea glehnii (closed bars), Abies sacharinensis (hatched bars) and Alnus japonica (open bars) in the forest mire in
Cape Ochiishi, eastern Hokkaido, Japan. Water samplings and chemical measurements of the peat pore water were made 18 times from 15 July to 22
November 2001 at regular intervals. Means sharing the same alphabetical letter does not significantly different between species at each depth by
Wilcoxon teat at p < 0.05 (n=18). Significance level by Friedman test among three depth for each species are; *** p < 0.001, ** p < 0.01, * p < 0.05, NS
p > 0.05.
Journal of Forestry Research (2010) 21(2): 119−128
124
Table 2 Chemical property (mean and SD) of peat pore water at three depths (0 cm, 30 cm, 60 cm) under the canopy of Picea glehnii, Abies
sacharinensis and Alnus japonica in the Cape Ochiishi Mire, eastern Hokkaido, Japan. Water samplings and chemical measurements of the peat
pore water were made 18 times from 15 July to 22 November 2001 at regular intervals. N and F sites denote 0 cm and 50 cm from each stem base,
respectively. Open site is in the Sphagnum community outside the forest. Significance level; *** p < 0.001, ** p < 0.01, * p < 0.05, NS not significant p >
0.05. SD: standard deviation, EC: electric conductivity
Species Chemical property
Picea glehnii EC H+ NH4+ Na+ K+ Mg2+ Ca2+ Cl- NO3- PO43- SO42-
mS m-1 mol c·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1
0 cm
N site Mean + 17.15+ 81.02+ 39.17+ 813.15+ 38.54+ 195.81+ 107.4+ 1156.28+ 0.11+ D.L.* 26.96+
SD 1.66 20.79 19.79 293.13 10.72 56.67 24.1 442.89 0.3 27.33
F site Mean 16.47+ 72.24+ 26.03+ 849.67+ 50.28+ 198.44+ 95.72+ 1162.99+ 0.04+ 0.63+ 32.6+
SD 3.3 26.7 26.32 285.23 38.21 65.38 31.64 496.06 0.16 1.75 28.88
Open site Mean 7.6+ 18.21+ 17.48+ 401.87+ 12.22+ 84.89+ 50.22+ 532.38+ 0.51+ D.L 22.3+
SD 1.51 6.24 10.5 123.33 5.64 31.11 17.91 211.74 0.79 16.85
N - F NS NS NS NS NS NS * NS NS NS NS
N- Open * * ** ** ** ** ** NS NS NS **
Significance
level
(Wilcoxson
signed-rank
test) F - Open * *** NS ** *** *** ** NS NS NS **
30 cm
N site Mean+ 19.66+ 27.89+ 47.61+ 996.52+ 56.39+ 219.1+ 130.1+ 1636.11+ 0.47+ D.L. 28.53+
SD 1.92 11.24 45.48 437.62 30.14 78.96 40.51 551.15 0.69 19.42
F site Mean+ 19.73+ 28.44+ 33.44+ 1146.78+ 39.07+ 287.36+ 151.76+ 1741.51+ 0.07+ 0.03+ 102.97+
SD 2.12 11.3 21.51 391.23 13.81 103.1 93.69 631.35 0.2 0.14 96.02
Open site Mean+ 8.04+ 2.17+ 16.9+ 432.47+ 9.36+ 82.3+ 60.34+ 519.34+ 0.28+ D.L 8.93+
SD 1.33 1.37 13.29 164.99 9.9 32.51 14.81 250.09 0.47 8.58
N - F NS NS NS NS NS NS NS NS NS NS **
N - Open ** *** * ** ** ** ** ** NS NS **
Significance
level
(Wilcoxson
signed-rank
test) F - Open ** *** NS ** ** ** ** * NS NS **
60 cm
N site Mean+ 20.06+ 21.05+ 42.25+ 1089.7+ 37.42+ 285.5+ 139.15+ 1761+ 0.41+ D.L 42.78+
SD 1.72 11.73 31.77 435.7 19.8 102.81 39.86 648.24 0.76 58.85
F site Mean+ 22.04+ 24.22+ 62.08+ 1114.56+ 44.08+ 327.85+ 142.82v 1805.25+ 0.08+ D.L 131.37v
SD 1.51 13.21 94.58 490.72 13.03 118.37 36.98 635.91 0.24 140.8
Open site Mean+ 8.86+ 4.21+ 17.56+ 411.5+ 14.52+ 77.34+ 61.1+ 510.86+ 0.29+ 0.15+ 4.27+
SD 1.09 2.04 7.32 78.22 21.26 16.08 19.23 116.17 0.62 0.48 3.61
N - F NS * NS NS *** NS NS NS NS NS *
N - Open ** *** ** *** ** *** *** ** NS NS ***
Significance
level
(Wilcoxson
signed-rank
test) F - Open ** *** ** ** ** *** *** ** NS NS ***
Species Chemical property
Abies sacharinensis EC H+ NH4+ Na+ K+ Mg2+ Ca2+ Cl- NO3- PO43- SO42-
mS m-1 molc·L -1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1
0 cm
N site Mean+ 16.25+ 72.4+ 14.69+ 843.48+ 48.09+ 163.18+ 96.69+ 1047.28+ D.L 3.57+ 15.66+
SD 1.28 20.81 17.76 135.48 12.11 52.55 25.33 350.35 11.61 14.22
F site Mean+ 15.67+ 65.77+ 14.12+ 821.57+ 38.93+ 156.05+ 100.35+ 1015.45+ D.L D.L 19.76+
SD 2.46 14.87 17.12 171.32 9.49 51.61 24.56 344.24 17.62
Open site Mean+ 7.6+ 18.21+ 17.48+ 401.87+ 12.22+ 84.89+ 50.22+ 532.38+ 0.51+ D.L 22.3+
SD 1.51 6.24 10.5 123.33 5.64 31.11 17.91 211.74 0.79 16.85
N - F NS NS NS NS NS NS NS NS NS NS NS
N- Open * ** NS ** ** ** ** NS NS NS **
Significance
level
(Wilcoxson
signed-rank
test) F - Open * *** NS *** NS *** ** NS NS NS **
30 cm
N site Mean+ 16.03+ 40.18+ 13.76+ 780.62+ 49.12+ 139.25+ 99.17+ 917.03+ D.L 1.43+ 16.99+
SD 1.23 11.65 17.07 113.84 12.21 28.13 20.18 236.58 3.51 13.01
F site Mean+ 17.72+ 18.8+ 20.21+ 941.43+ 65.96+ 197.28+ 122.47+ 1314+ 0.02+ 1.23+ 21.77+
Journal of Forestry Research (2010) 21(2): 119−128
125
Continue Table 2
SD 2.07 13.53 20.64 174.11 39.58 44.06 30.18 393.84 0.09 3.2 31.46
Open site Mean+ 8.04+ 2.17+ 16.9+ 432.47+ 9.36+ 82.3+ 60.34+ 519.34+ 0.28+ D.L. 8.93+
SD 1.33 1.37 13.29 164.99 9.9 32.51 14.81 250.09 0.47 8.58
N - F ** ** NS ** NS ** * ** NS NS *
N - Open ** *** NS *** ** ** ** NS NS NS **
Significance
level
(Wilcoxson
signed-rank
test) F - Open ** *** NS *** ** ** ** NS NS NS **
60 cm
N site Mean+ 18.88+ 11.77+ 24.53+ 943.31+ 45.21+ 221.23+ 139.33+ 1265.23+ 0.01+ 0.08+ 12.91+
SD 1.98 8.42 25.92 179.35 14.04 52.94 33.37 331.98 0.06 0.23 13.59
F site Mean+ 18.43+ 14.03+ 34.87+ 998.62+ 39.05+ 222.05+ 132.2+ 1329.62+ 0.03+ 0.9+ 12.02+
SD 1.98 6.88 17.27 145.71 31.02 42.87 29.13 329.86 0.12 3.61 12.18
Open site Mean+ 8.86+ 4.21+ 17.56+ 411.5+ 14.52+ 77.34+ 61.1+ 510.86+ 0.29+ 0.15+ 4.27+
SD 1.09 2.04 7.32 78.22 21.26 16.08 19.23 116.17 0.62 0.48 3.61
N - F NS NS NS * ** NS *** NS NS NS NS
N - Open ** *** NS *** *** *** *** ** NS NS ***
Significance
level
(Wilcoxson
signed-rank
test) F - Open ** *** *** *** *** *** *** * NS NS ***
Species Chemical property
Alnus japonica EC H+ NH4+ Na+ K+ Mg2+ Ca2+ Cl- NO3- PO43- SO42-
mS m-1 mol c·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1 mmolc·L-1
0 cm
N site Mean+ 15.65+ 37.35+ 15.67+ 773.37+ 48.86+ 155.39+ 100.14+ 875.34+ 0.03+ D.L 8.02+
SD 1 8.77 17.35 104.74 12.34 21.45 16.32 269.87 0.13 12.77
F site Mean+ 14.9+ 33.39+ 22.7+ 793.91+ 49.36+ 159.04+ 104.5+ 915.45+ D.L. 0.04+ 9.49+
SD 1.23 13.3 19.31 111.68 19.24 25.52 18.31 254.44 0.16 25.42
Open site Mean+ 7.6+ 18.21+ 17.48+ 401.87+ 12.22+ 84.89+ 50.22+ 532.38+ 0.51+ D.L. 22.3+
SD 1.51 6.24 10.5 123.33 5.64 31.11 17.91 211.74 0.79 16.85
N - F * NS NS NS NS NS NS * NS NS NS
N- Open * ** *** ** *** NS ** ** NS NS ***
Significance
level
(Wilcoxson
signed-rank
test) F - Open * ** NS ** *** * ** NS NS NS **
30 cm
N site Mean+ 15.68+ 24.61+ 15.39+ 771.6+ 38.98+ 167.82+ 121.21+ 869.11+ D.L 1.47+ 3.39+
SD 1.4 8.34 20.6 248.7 26.19 68.21 42.36 330.43 3.06 4.1
F site Mean+ 14.56+ 27.78+ 14.97+ 718.65+ 39.07+ 139.87+ 104.02+ 790.85+ 0.06+ 6.17+ 27.61+
SD 1.19 18.55 17.95 204.66 19.7 37.06 24.03 280.06 0.17 17.7 30.92
Open site Mean+ 8.04+ 2.17+ 16.9+ 432.47+ 9.36+ 82.3+ 60.34+ 519.34+ 0.28+ D.L 8.93+
SD 1.33 1.37 13.29 164.99 9.9 32.51 14.81 250.09 0.47 8.58
N - F ** NS NS NS NS * * NS NS NS *
N - Open ** *** *** *** *** ** ** NS NS * **
Significance
level
(Wilcoxson
signed-rank
test) F - Open ** *** NS *** *** ** ** * NS * **
60 cm
N site Mean+ 18.11+ 13.19+ 23.27+ 803.2+ 29.94+ 168.89+ 103.67+ 924.11+ 0.11+ D.L 19+
SD 2.51 7.29 22.64 247.97 10.41 53.08 29.57 346.69 0.47 36.96
F site Mean+ 16.2+ 13.33+ 14.81+ 812.21+ 47.4+ 173.29+ 112.14+ 929.34+ 0.28+ 3.27+ 11.41+
SD 2.32 8.52 21.85 223.91 37.15 57.12 31.83 326.2 0.64 5.51 21.76
Open site Mean+ 8.86+ 4.21+ 17.56+ 411.5+ 14.52+ 77.34+ 61.1+ 510.86+ 0.29+ 0.15+ 4.27+
SD 1.09 2.04 7.32 78.22 21.26 16.08 19.23 116.17 0.62 0.48 3.61
N - F ** NS NS NS NS NS NS NS NS * NS
N - Open ** *** *** *** *** *** *** ** NS * ***
Significance
level
(Wilcoxson
signed-rank
test) F - Open ** *** *** *** *** *** *** NS NS NS ***
*: below the detection limit (0.01µmolc•L-1).
Most of the ion species except for NH4+ and NO3– concentra-
tions had positive loadings for the first principal component,
whereas NO3– concentration had negative loadings for the first
principal component (Fig.2b). NH4+ and NO3– concentrations
had positive loadings for the second principal component,
whereas H+ concentration had negative loading for the second
Journal of Forestry Research (2010) 21(2): 119−128
126
principal component.
Fig. 2 (a) Ordination by principal component analysis (PCA) using
first and second principal component scores at each sampling point.
N and F denote the distance from the stem base of 0 cm (N) and 50
cm (F), respectively. Open denotes the open site outside the forest.
The following number of 0, 30, and 60 denote the depth of the
sampling point in cm. Closed square: peat pore water at the base of
Picea glehnii, Hatched square: peat pore water at the base of Abies
sacharinensis, Open square: peat pore water at the base of Alnus
japonica, Open triangle: peat pore water at the open site, Closed
circle: stemflow of Picea glehnii, Hatched circle: stemflow of Abies
sacharinensis, Open circle: stemflow of Alnus japonica, and (b)
loadings on the first and second principal component scores of each
chemical parameter used in the PCA.
Discussion
The bulk deposition in the Cape Ochiishi Mire was 637 mm from
June 15 to December 1, 2001. The distribution ratio of the
stemflow to the bulk deposition was estimated to be 0.19%,
0.12%, and 0.57%, respectively, in the Picea glehnii, Abies
sacharinensis, and Alnus japonica. This result was representative
of forest types. Capturing ability of precipitation was greater in
coniferous canopy due to the higher interception within the
canopy and to the conifer morphology. The lower volume
observed in the coniferous stand was probably due to branch
disposition and greater trunk surface roughness.
Three mire species were distinguished using principal compo-
nent analysis of the chemical composition of stemflow (Fig. 2).
Most ion concentrations including sea salt components had posi-
tive loadings for the first principal component, and hence the
stemflow of P. glehnii was affected by the highest salt loading
and that of the A. japonica was affected by the lowest salt load-
ing to the stemflow. The differences in the salt loadings among
the species would be due to the differences in the canopy struc-
ture. Coniferous species canopies usually have a higher ability to
capture atmospheric deposition than hardwood, and as a conse-
quence throughfall and stemflow of conifers usually contain
higher concentrations of salts and H+ (Kaul and Billings 1965;
Mahendrappa 1974; Neary and Gizyn 1994; Bergkvist and
Folkeson 1995). In Cape Ochiishi Mire, the H+ concentrations in
the stemflow of P. glehnii and A. sacharinensis were higher than
in A. japonica trees is in good agreement with these published
results.
Although the differences in peat pore water chemistry among
the three species was not so clearly separated by PCA, the
distribution order of the peat pore water of the three species
matched the order of the stem flow of the three species: P.
glehnii had the highest and A. japonica had the lowest score on
the first principal component axis for both the stem flow and peat
pore water (Fig. 2). Chemical environment in the peat soils were
different between coniferous and broadleaf forests, and a part of
the difference could be explained by the chemical properties of
stemflow. Tsuyuzaki and Haraguchi (2009) demonstrated that the
chemical difference in the peat soil environments between P.
glehnii and A. japonica affects both the seed germination and
seed survival and consequence an abrupt boundary is developed
between them. Therefore, the maintenance of chemical
environments could be important on the stability of the
communities. Our study shows that stemflow contribute the
maintenance of communities because stemflow provide one of
components to the chemical difference in the peat soil between
coniferous and broadleaf forests.
On the other hand, the differences along the gradient from the
tree base to the site 50 cm from the base (comparison between N
and F sites) was significant only for limited parameters and the
tendency was not consistent (Table 2). An unclear gradient of
chemical parameters from the stem base in the peat soil stands
was the common observation in the three species. Gesper (1970)
who investigated radioisotope accumulation in soils around
beech trunks, speculated that stemflow water on the slope
(ca.11º) was likely to flow vertically and also contributed sig-
nificantly to internal downslope flow. In Cape Ochiishi Mire, the
stemflow movement in the high water content of the peat could
be more slow, because three species in the mire established on
the flat topography (< ca. 1º). This would be due to the fact that
the chemicals in stem flow would be diluted by the abundant
peat pore water. Moreover, proton that was produced by dissolu-
tion from humic substances (Haraguchi et al. 2003; Nishijima
and Nakata 2004) and ion exchange with base cations in the
precipitation (Gorham 1956) was dominant buffer reactions in
the peat soil. The significant differences in the chemical envi-
ronments particularly soil pH between stem base and distal areas
appeared interspecies difference rather than the surface soil. This
Journal of Forestry Research (2010) 21(2): 119−128
127
result indicated that effect of stemflow not only determined the
soil close to the stem but also the soil extended the area under the
canopy. Thus, it is presumed that stemflow provide the best site
both close to the site and under the canopies and play an impor-
tant role in promoting the natural regeneration.
Peat pore water at the peat surface had the lowest and that at
the 60 cm depth had almost the highest score on the second
principal component axis. This high score for the second
principal component was due to the high concentrations of NO3-
and NH4+ and the low concentration of H+. Accumulation of
NO3- and NH4+ under the high pH condition in the subsurface of
the peat layer would be due to the decomposition of organic
substances in the subsurface of the peat layer. The spatial
variability of the peat soil chemistry was affected by stemflow
chemistry, would be an important factor in the decomposition of
peat within the peatland forest.
Thus we can conclude that the effects of stemflow on the peat
pore water chemistry are clearly seen only at the stem base in the
peat forest in the mire and the peat pore water around the stem
base of a tree has particular chemical properties specific to each
species; however, the gradient of the chemical environment from
the stem base to the area beyond the canopy was not significant
in the peatland forest. The peat pore water chemistry under the
canopy would be mainly controlled by the chemical processes
within the peat. The spatial heterogeneity of chemical
environment between microsites within peatland forests would
be contributed indirectly through the control of microorganism
activity, nutrient regeneration and the understory vegetation
mediated the surface water and the stemflow of the dominant
canopy trees. In inundated soil in peatland, the chemical
environments of the peat soil around the tree stand appear
species specificity. Therefore, the maintenance of chemical
environment which focus on species-specific needs for the
conservation of the peatland forests, it means that the
hydrological change such as large fluctuation of groundwater
level diminishes.
Acknowledgements
This work was partly funded by the Grant in Aid from JSPS.
Chemical analysis was made in the Instrumentation Centre of
The University of Kitakyushu.
References
Ahmad-Shah A, Rieley JO. 1989. Influence of tree canopies on the quantity of
water and amount of chemical elements reaching the peat surface of a basin
mire in the midlands of England. Journal of Ecology, 77: 357–370.
Bergkvist B, Folkeson L. 1995. The influence of tree species on acid
deposition, proton budgets and element fluxes in south Swedish forest
ecosystems. Ecological Bulletins, 44: 90–99.
Beven K, Germann P. 1982. Macropores and water flow in soils. Water
Resources Research, 18: 1311–1325.
Bini C, Bresolin F. 1998. Soil acidification by acid rain in forest ecosystems:
A case study in northern Italy. The Science of Total Environment, 222:
1–15.
Butler TJ, Likens GE. 1995. A direct comparison of throughfall plus stem
flow to estimates of dry and total deposition for sulfur and nitrogen. At-
mospheric Environment, 29: 1253–1265.
Cappellato R, Peters NE. 1995. Dry deposition and canopy leaching rates in
deciduous and coniferous forests of the Georgia Piedmont: an assessment
of a regression model. Journal of Hydrology, 169: 131–150.
Chang SC, Matzner E. 2000. The effect of beech stemflow on spatial patterns
of soil solution chemistry and seepage fluxes in a mixed beech/oak stand.
Hydrological Processes, 14: 135–144.
Chiwa M, Kim DH, Sakugawa H. 2003. Rainfall, stemflow, and throughfall
chemistry at urban– and mountain-facing sites at Mt. Gokurakuji,
Hiroshima, western Japan. Water, Air, and Soil Pollution, 146: 93–109.
Clement M, Wittig R. 1987. Heavy metal content of the moss Mnium hornum
growing in the stem flow area of Fagus sylvatica. Acta Oecologica, 8:
257–264.
Crockford RH, Richardson DP, Sageman R. 1996. Chemistry of rainfall,
throughfall and stemflow in a eucalypt forest and a pine plantation in
south-eastern Australia: 3. stemflow and total inputs. Hydrological
Processes, 10: 25–42.
Dämmgen U, Zimmerling R. 2002. Vertical fluxes of air-borne acidifying and
eutrophying species in the Schorheide Nature Researve in Brandenburg,
Germany. Journal of Applied Botany, 76: 190–202.
Duijsings JJHM, Verstraten JM, Bouten W. 1986. Spatial variability in
nutrient deposition under an oak/beech canopy. Zeitschrift für
Pflanzenernährung und Bodenkunde, 149: 718–727.
Falkengren-Grerup U, Björk L. 1991. Reversibility of stemflow-induced soil
acidification in Swedish beech forest. Environmental Pollution, 74: 31–37.
Förster H, Schimmack W. 1992. Influence of the stemflow on the depth
distribution of radiocesium in the soil under a beech stand.
Naturwissenschaften, 79: 23–24.
Gersper PL, Holowaychuk N. 1971. Effects of stem flow from forest canopy
trees on chemical properties of soils. Ecology, 52: 691–702.
Gordon AM, Chourmouzis C, Gordon AG. 2000. Nutrient inputs in litterfall
and rainwater fluxes in 27-year old red, black and while spruce plantations
in Central Ontario, Canada. Forest Ecology and Management, 138: 65–78.
Gorham E. 1956. The ionic composition of some bogs and fen waters in the
English Lake District. Journal of Ecology, 44: 142–152.
Haraguchi A. 1996. Effect of pH on photosynthesis of five Sphagnum species
in mires in Ochiishi, northern Japan. Wetlands, 16: 10–14.
Haraguchi A, Iyobe T, Nishijima H, Tomizawa H. 2003. Acid and sea-salt
accumulation in coastal peat mires of a Picea glehnii forest in Ochiishi,
eastern Hokkaido, Japan. Wetlands, 23: 229–235.
Hori T. 1953. Studies on Fogs. The Institute of Low Temperature Science,
Hokkaido Univ, Sapporo: 399pp.
Hotes S, Poschlod P, Sakai H, Inoue T. 2001. Vegetation, hydrology, and
development of a coastal mire in Hokkaido, Japan, affected by flooding and
tephra deposition. Canadian Journal of Botany, 79: 341–361.
Houle D, Ouimet R, Paquin R, Laflamme J-G. 1999. Interactions of
atmospheric deposition with a mixed hardwood and a coniferous forest
canopy at the Lake Clair Watershed (Duchesnay, Quebec). Canadian
Journal of Forest Research, 29: 1944–1957.
Iyobe T, Haraguchi A. 2008. Ion flux from precipitation to peat soil in spruce
forest – Sphagnum bog communities in Ochiishi district, eastern Hokkaido,
Japan. Limnology, 9:89–99.
Jochheim H. 1986. Einfluß des Stammablaufwassers auf den
Pflanzenbewuchs und den chemischen Bodenzustand im Oberboden von
Altbuchenbeständen. Verhandlungen der Gesellschaft für Ökologie, 14:
Journal of Forestry Research (2010) 21(2): 119−128
128
299–307.
Joslin JD, Wolfe MH. 1992. Red spruce soil solution chemistry and root
distribution across a cloud water deposition gradient. Canadian Journal of
Forest Research, 22: 893–904.
Kaneko N, Kofuji R. 2000. Effects of soil pH gradient caused by stemflow
acidification on soil microarthropod community structure in a Japanese red
cedar plantation: an evaluation of ecological risk on decomposition.
Journal of Forest Research, 5: 157–162.
Kaul ON, Billings WD. 1965. Cation content of stemflow in some forest trees
in North Carolina. Indian Forester, 91: 367–370.
Liang W.-L, Kosugi K, Mizuyama T. 2009. A three-dimensional model of the
effect of stemflow on soil water dynamics around a tree on a hillslope.
Journal of Hydrology, 366: 62–75.
Lovett G.M, Lindberg SE. 1993. Atmospheric deposition and canopy
interactions of nitrogen in forests. Canadian Journal of Forest Research,
23: 1603–1616.
Mahendrappa MK. 1974. Chemical composition of stemflow from some
eastern Canadian tree species. Canadian Journal of Forest Research, 4:
1–7.
Martinez-Meza E, Whitford W.G. 1996. Stemflow, throughfall and
channelization of stemflow by roots in three Chihuahuan desert shrubs.
Journal of Arid Environments, 32: 271–287.
Matschonat G., Falkengren-Grerup U. 2000. Recovery of soil pH,
cation-exchange capacity and the saturation of exchange sites from
stemflow-induced soil acidification in three Swedish beech (Fagus sylvatica
L.). Scandinavian Journal of Forest Research, 15: 39–48.
Neal C, Reynolds B, Neal M, Hughes S, Wickham H, Hill L, Rowland P,
Pugh B. 2003. Soluble reactive phosphorus levels in rainfall, cloud water,
throughfall, stemflow, soil waters, stream waters and groundwaters for the
Upper River Severn area, Plynlimon, mid Wales. The Science of the Total
Environment, 314–316: 99–120.
Neary AJ, Gizyn WI. 1994. Throughfall and stemflow chemistry under
deciduous and coniferous forest canopies in south-central Ontario.
Canadian Journal of Forest Research, 24: 1089–1100.
Nishijima H, Iyobe T, Nishio F, Tomizawa H, Nakata M, Haraguchi A. 2003.
Site selectivity of Picea glehnii forest on Syunkunitai sand spit, north
eastern Japan. Wetlands, 23: 406–415.
Nishijima H, Nakata M. 2004. Relationship between plant cover type and soil
properties on Syunkunitai coastal sand dune in eastern Hokkaido.
Ecological Research, 19: 581–591.
Piirainen S, Finér L, Starr M. 2002. Deposition and leaching of sulphate and
base cations in a mixed boreal forest in eastern Finland. Water, Air, and Soil
Pollution, 131: 185–204.
Potts M.J. 1978. The pattern of deposition of air-borne salt of marine origin
under a forest canopy. Plant and Soil, 50: 233–236.
Stevens PA, Hornung M, Hughes S. 1989. Solute concentrations, fluxes and
major nutrient cycles in a mature sitka-spruce plantation in Beddgelert
forest, North Wales. Forest Ecology and Management, 27: 1–20.
Stöckli H. 1991. Influence of stemflow upon the decompositing system in two
beech stands. Revue décologie et de biologie du sol, 28: 265–286.
Tajchman SJ, Keys RN, Kosuri SR. 1991. Comparison of pH, sulfate and
nitrate in throughfall and stemflow in yellow-poplar and oak stands in
north-central West Virginia. Forest Ecology and Management, 40: 137–144.
Tomizawa H, Oikawa M, Nishio F, Haraguchi A. 1997. Chemical
modification of rain water by Alnus japonica wetland forest in Kiritapp
Mire, Eastern Hokkaido, Japan. Journal of Forest Research, 2: 189–192.
Tsuyuzaki S, Haraguchi A. 2009. Maintenance of an abrupt boundary between
needle-leaved and broad leaved forests in a wetland near coast. Journal of
Forestry Research, 20: 91–98.
Voigt GK. 1960. Distribution of rainfall under forest stands. Forest Science, 6:
2–10.