ORIGINAL ARTICLE
Twenty-three years of stand dynamics in an old-growthChamaecyparis forest in central Japan
Michinari Matsushita • Daisuke Hoshino •
Shin-Ichi Yamamoto • Naoyuki Nishimura
Received: 25 June 2012 / Accepted: 26 January 2013 / Published online: 6 March 2013
� The Japanese Forest Society and Springer Japan 2013
Abstract Structures and dynamics of old-growth conif-
erous stands are affected by several types of disturbances
including typhoons. We report the forest dynamics of four
old-growth Chamaecyparis stands in central Japan that
differ in the disturbance history of typhoons over a period
of 23 years. The stem number, basal area and mortality
were examined. In a predominant stand of C. obtusa (Sieb.
et Zucc.) Endl., 24 % of the C. obtusa canopy trees died,
mainly as a result of the severe damage of a strong typhoon
that caused a single tree-fall gap and the following gap
enlargements. In this stand, the total basal area decreased to
76.5 % of the initial value, although the mortality declined
in recent years. In contrast, the other three stands decreased
only slightly in the stem numbers (0.0–5.6 %) and
increased in the basal areas of C. obtusa canopy trees. It is
confirmed that the stand-level ingrowths of 300-year-old
C. obtusa canopy trees could contribute to the increase in
the stock of each stand. Our results support an idea that the
dynamics of old-growth Chamaecyparis forests were
greatly affected by typhoons. The stand structures will be
gradually changed (with the processes of gap dynamics)
and C. obtusa will continue to be dominant, potentially
over hundreds of years.
Keywords Forest dynamics � Gap formation �Long-term study � Natural disturbance � Stem growth
Introduction
Stand structures and dynamics are often affected by several
types of disturbances (e.g., typhoons) that vary in intensity
and frequency (Yamamoto et al. 2011; Torimaru et al.
2012). In many old-growth coniferous forests in East Asia,
gap creation by typhoons is one of the major disturbance
regimes (e.g., Asai et al. 2003). Because such disturbances
occur infrequently and stochastically (Yamamoto et al.
2011; Torimaru et al. 2012), their effects on dynamics of
old-growth forests are often complex (Parish and Antos
2004), and therefore long-term monitoring studies are
important.
The genus Chamaecyparis is distributed around the
Pacific basin and along the eastern coast of North America
(Farjon 2005). Most Chamaecyparis species are known as
long-lived with slow growth rates. For example, the max-
imum age of C. formosensis, a species in Taiwan, exceeds
ca. 3,000 years old (Zobel 1998). In North America,
C. nootkatensis is a very long-lived stress tolerator found in
a wide variety of habitats, which appears to need some
opening of the stand to reach the canopy (Antos and Zobel
1986; Antos et al. 2005; Parish and Antos 2006). Because
Nomenclature: Ohwi and Kitagawa (1992).
M. Matsushita (&)
Laboratory of Forest Sciences, Department of Biological
Environment, Akita Prefectural University, Akita 010-0195,
Japan
e-mail: [email protected]
D. Hoshino
Tohoku Research Center, Forestry and Forest Products Research
Institute, 92-25, Morioka, Iwate 020-0123, Japan
S.-I. Yamamoto
Laboratory of Forest Plant Ecology, Graduate School
of Bioagricultural Sciences, Nagoya University, Chikusa,
Nagoya 464-8601, Japan
N. Nishimura
Environmental Sciences Laboratory, Faculty of Social
and Information Studies, Gunma University, 4-2 Aramaki,
Maebashi, Gunma 371-8510, Japan
123
J For Res (2014) 19:134–142
DOI 10.1007/s10310-013-0398-x
of their longevities and slow growth, the regeneration
processes of Chamaecyparis species have not fully been
understood.
Two Chamaecyparis species, C. obtusa (Sieb. et Zucc.)
Endl. and C. pisifera (Sieb. et Zucc.) Endl. are indigenous
to Japan. The rates of exploitation of Chamaecyparis have
been very high in the past, since the lumber of Chamae-
cyparis spp., especially C. obtusa, has been used prefer-
entially to build feudal architectures (e.g., castles) because
of its excellent quality (Nishioka and Obara 1975; Ito
2000). According to IUCN 2010, C. obtusa is classified as
‘‘Near Threatened’’, and the remaining natural C. obtusa
forests are small and fragmented (Maeda 1951; Maeda and
Yoshioka 1952; Matsumoto et al. 2010). Therefore, their
conservation has been an important and urgent issue
(Yamamoto 1998; Tsumura et al. 2007). Currently, Kiso
district in central Japan is the core habitat over the entire
geographical distribution of natural C. obtusa populations
(Matsumoto et al. 2010); old-growth Chamaecyparis for-
ests are rare and thus very precious because of both
their ecological and commercial value (Yokouchi 1970;
Yamamoto 1998).
Akaswa Forest Reserve in the Kiso district is an excel-
lent representative of Chamaecyparis forests, characterized
by the pronounced dominance of C. obtusa (Nagano
Regional Forest Office 1985; Yamamoto 1993a; Hoshino
et al. 2001). Most Chamaecyparis canopy trees reach
around 300 years old (Hoshino et al. 2001; Asai et al.
2003). To understand the current condition and long-term
trends in Chamaecyparis forests, we began to examine the
dynamics of typical old-growth stands in this Reserve in
1985. In this paper, we report the dynamics of old-growth
C. obtusa stands over 23 years: i.e., the changes in the stem
number, basal area and mortality patterns. Since little
information is available on old-growth Chamaecyparis
forests, our report provides useful information about Cha-
maecyparis regeneration.
Here, specific questions were: (1) are the number and
basal area of C. obtusa canopy trees decreasing or
increasing; (2) how much are C. obtusa trees growing in
diameter; and (3) what, if any, are the mortality patterns of
C. obtusa trees? We summarized the structure and
dynamics of each stand focusing on the role of typhoon
disturbances, and then discussed long-term trends and
future states of old-growth Chamaecyparis forests.
Materials and methods
Study area
The Akasawa Forest Reserve (35�4305700N, 137�3705000E;
1,046 ha; 1,080–1,558 m a.s.l.) is located in the Kiso
district of Nagano Prefecture, central Honshu, Japan
(Nagano Regional Forest Office 1985). Annual precipita-
tion is ca. 2,500 mm and snow accumulation is 50–100 cm.
The mean annual temperature is 7.8 �C at 1,113 m a.s.l.;
the mean monthly maximum and minimum temperatures
are 14.3 �C in August and -11.8 �C in February, respec-
tively. The reserve is on an elevated peneplain with a
gentle slope. The geology is dominated by acidic igneous
rocks, including granite, granite porphyry and rhyolite.
Soils are mainly dry or wet podzols, although brown forest
soils appear on hillsides or along streams (Nagano Regio-
nal Forest Office 1985; Yamamoto 1993a).
Old-growth Chamaecyparis stands in this Reserve, like
the other stands in the Kiso and neighboring districts, have
become established after heavy cutting during 1688–1703
(Nagano Regional Forest Office 1985). Since that time, most
stands have been protected from heavy cutting, but selection
cutting mainly for hardwoods has been undertaken.
In the reserve, C. obtusa mainly dominates the over-
story, and C. pisifera frequently co-dominates on the lower
slopes or along the streams (Nagano Regional Forest Office
1985; Hoshino et al. 2001). Other coniferous trees like
Thujopsis dolabrata (L.f.) Sieb. et Zucc. and Thuja
standishii (Gordon) Carriere, and some hardwood trees
such as Magnolia obovata Thunb. and Quercus mongolica
Fisch. ex Ledeb. var. grosseserrata (Blume) Rehder et
E.H.Wilson, occasionally co-occur. The understory layer in
this Reserve is characterized by dense cover of the saplings
of T. dolabrata, occasionally with small trees of some
broadleaved species (Yamamoto and Suto 1994; Hoshino
et al. 2001, 2003; Matsushita et al. 2010). T. dolabrata is a
highly shade-tolerant species which can reproduce by
layering under closed canopy conditions (Yamamoto and
Suto 1994); meanwhile, the seedlings of C. obtusa estab-
lished on exposed mineral soil beneath tree-fall gaps
(Yamamoto 1988, 1993b), and there are no or very few
saplings of C. obtusa (Yamamoto and Suto 1994; Hoshino
et al. 2001, 2003).
Study plot and field methods
Four representative old-growth stands were chosen for this
study (Table 1). The canopy of Stand O consisted of only
C. obtusa, whereas Stands S and H had a few other species
in the canopy but were dominated by C. obtusa. However,
several species co-existed in the canopy of Stand K. While
there were no records of selection cutting in Stand H,
Stands O, S and K underwent selection cutting during the
period 1915–1947, mainly to remove ‘‘dead’’ canopy trees
(Nagano Regional Forest Office 1985). When C. obtusa
trees have died naturally (e.g., as the result of typhoons),
the dead trees have been (but not frequently) removed from
the reserve by forest managers.
J For Res (2014) 19:134–142 135
123
There is no evidence of forest fires, but typhoons are a
major natural disturbance. The Isewan typhoon in 1959
caused severe damage to several stands in the reserve
(Nagano Regional Forest Office 1985), and, especially,
Stand K was severely damaged by the typhoon (Yamamoto
1993a). Stand O was affected by another strong typhoon in
1991. At these times, dead trees were removed from these
stands, and stumps were left.
In September–October 1985, a 0.2-ha plot (40 9 50 m)
was laid out in a representative location within each forest
stand. In each plot, canopy trees [stem diameter at breast
height (DBH) C20 cm] were tagged, and their species
name, living status, location, and DBH were recorded.
Re-censuses were conducted from then until 2008 at ca.
5-year intervals (specific census years are shown in
Table 2). The condition of dead trees was classified as
standing dead, uprooted, stem broken or stump.
Data analysis
The annual rate of stem mortality was calculated as:
100 � lnNi � lnNsð Þ=T ;
where Ni is the initial number of living stems and Ns is the
number of the surviving stems at each subsequent census,
and T is the time interval (years) between the censuses.
Basal area (BA) of each stem ([20 cm DBH) was cal-
culated as: p 9 (DBH/2)2. Then, the stand-level BA was
defined as the sum of the BAs of all stems ([20 cm DBH)
within each plot. As an index of ingrowth, the increment in
the stand BA of surviving trees (excluding recruited trees)
was defined as follows:
Growth of stand BA ¼ ln BAf � ln BAið Þ=T;
where BAf and BAi are the stand BAs at the final and initial
census, respectively, and T is the time interval (years)
between the final and initial censuses.
As an index of the growth of each individual stem, the
increment in the DBH of surviving trees was defined as the
absolute growth rate (AGR), as follows:
AGR ¼ DBHf � DBHið Þ=T;
where DBHf and DBHi are the DBHs of each stem at the
final and initial census, respectively, and T is the time
interval (years) between the final and initial censuses.
The 95 % confidence intervals for these parameters
(mortality, growth of stand BA, and AGR), were estimated
by the standard bootstrapping method, with the populations
resampled 1,000 times from the original data. We used the
bias-corrected percentile method to determine confidence
intervals (Manly 2007). Statistical analyses were performed,
by using R 2.8.2 (R Development Core Team 2008).
Results
Changes in the stem density and mortality
Over the 23-year study period, the numbers of coniferous
canopy trees decreased from 75 to 57 in Stand O, from 67
to 65 in Stand S, and from 85 to 76 in Stand H, but
increased from 85 to 96 in Stand K (Table 2). When tar-
geting only C. obtusa, whereas there was a slight increase
in the number from 15 to 16 in Stand K, the numbers
decreased in the other three stands.
The mortality rate of canopy trees was highest in Stand
O, whereas it was very low in Stands S and K (Fig. 1). In
Stand O, the mortality rate of C. obtusa canopy trees was
higher in the periods 1990–1998 and 1998–2003, and then
decreased in the period 2003–2008 (Table 2). During the
23 years, stem recruitment events were observed only for
Stand K, where 1 C. obtusa tree and 12 hardwood trees
(mainly of M. obovata) were recruited, while there was no
recruitment stems in Stands O, S and H.
Table 1 Attributes of the four old-growth Chamaecyparis stands in the Akasawa Forest Reserve
Stand H Stand S Stand O Stand K
Topographic position Middle slope Middle slope Upper slope Lower slope
Slope direction E SW NW SW
Slope inclination (�) 0–5 0–5 0–5 5–10
Stand height (m) 33–34 35 35–36 25–26
Dominant canopy tree
species
Chamaecyparis obtusa, Thujopsis
dolabrata, Magnolia obovata
C. obtusa, Thujopsis
dolabrata
C. obtusa C. obtusa, C. pisifera;
hardwood trees
Dominant understory
species
Thujopsis dolabrata T. dolabrata Deciduous broad-
leaved shrubs
C. pisifera
History of major
disturbance
None Selection cutting
during 1915–1947
Selection cutting
during 1915–1947
Strong typhoon in
1991
Selection cutting during
1915–1947
Isewan typhoon in 1957
136 J For Res (2014) 19:134–142
123
Ta
ble
2T
wen
ty-t
hre
e-y
ear
chan
ge
inth
en
um
ber
and
BS
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can
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ur
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Ch
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Nu
mb
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fst
emB
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area
(m2
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19
85
19
90
–1
99
3a
19
98
–2
00
0b
20
03
20
08
19
85
19
90
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99
3a
19
98
–2
00
0b
20
03
20
08
Sta
nd
O
To
talc
75
74
0.2
7(0
.00
–0
.82
)
66
1.4
3(0
.52
–2
.42
)
60
1.9
1(0
.62
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.65
)
57
1.0
3(0
.00
–2
.48
)
17
.31
7.1
(0.4
)1
5.0
(2.3
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3.9
(1.1
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3.3
(0.7
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Sta
nd
S
Ch
am
aec
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ris
ob
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66
64
0.6
3(0
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ot
test
ed)
64
0.0
0(n
ot
test
ed)
64
0.0
0(n
ot
test
ed)
14
.41
4.4
(0.4
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4.5
(0.0
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4.9
(0.0
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5.0
(0.0
)
Th
ujo
psi
sd
ola
bra
ta1
1 0.0
0(n
ot
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1 0.0
0(n
ot
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1 0.0
0(n
ot
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ed)
1 0.0
0(n
ot
test
ed)
0.3
0.3
(0.0
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0.3
(0.0
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.3(0
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To
tal
67
65
0.6
1(0
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65
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ot
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ot
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ed)
14
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4.6
(0.4
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Sta
nd
H
Ch
am
aec
ypa
ris
ob
tusa
71
70
0.2
8(0
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67
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4(0
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)
67
0.0
0(n
ot
test
ed)
67
0.0
0(n
ot
test
ed)
15
.71
5.9
(0.1
)1
5.9
(0.5
)1
6.0
(0.1
)1
6.5
(0.0
)
Th
ujo
psi
sd
ola
bra
ta1
31
2
1.6
0(0
.00
–5
.30
)
11
0.8
7(0
.00
–2
.88
)
10
3.1
8(0
.00
–1
0.6
0)
9 2.1
0(0
.00
–7
.13
)
1.4
1.4
(0.0
)1
.3(0
.1)
1.2
(0.1
)1
.1(0
.1)
Oth
ers
11
10
00
.10
.10
.10
.10
.0
To
tal
85
83
0.4
8(0
.00
–1
.21
)
79
0.4
9(0
.12
–1
.01
)
77
0.4
9(0
.85
–2
.18
)
76
0.2
6(0
.00
–0
.80
)
17
.21
7.3
(0.1
)1
7.3
(0.5
)1
7.2
(0.2
)1
7.6
(0.1
)
Sta
nd
K
Ch
am
aec
ypa
ris
ob
tusa
15
15
0.0
0(n
ot
test
ed)
16
0.0
0(n
ot
test
ed)
16
0.0
0(n
ot
test
ed)
16
0.0
0(n
ot
test
ed)
1.7
1.7
(0.0
)1
.9(0
.0)
2.0
(0.0
)2
.1(0
.0)
Ch
am
aec
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ris
pis
ifer
a3
33
3
0.0
0(n
ot
test
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33
0.0
0(n
ot
test
ed)
33
0.0
0(n
ot
test
ed)
32
0.6
2(0
.00
–1
.90
)
4.1
4.3
(0.0
)4
.6(0
.0)
4.7
(0.0
)4
.8(0
.1)
Oth
ers
37
42
49
48
48
1.8
2.4
2.9
3.1
3.5
To
tal
85
90
0.0
0(n
ot
test
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98
0.0
0(n
ot
test
ed)
97
0.2
1(0
.00
–0
.62
)
96
0.4
2(0
.00
–1
.06
)
7.6
8.4
(0.0
)9
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.0)
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(0.0
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0.4
(0.1
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J For Res (2014) 19:134–142 137
123
Growth trends at the stand and stem levels
The total BA decreased in Stand O, and did not change (or
slightly increased) in Stands S and H (Table 2). However,
the stand total BA consistently increased in Stand K. The
stand-level BA of C. obtusa showed similar trends with the
stand total BA.
During the 23 years, the growth of stand total BA was
highest in Stand K, and that was gradually higher in Stand
H than Stands S and O (Fig. 2, lower). On the other hand,
the growth in the stand-level BA of C. obtusa was lowest in
Stand K (Fig. 2, upper), while the corresponding trends for
the other stands were similar with those stand total BAs.
At individual-stem level, the AGR was higher in Stand
K than the other stands, but the rates were relatively similar
among Stands O, S, and H (Fig. 3). C. obtusa trees in Stand
K showed greatest absolute diameter growth rates (Fig. 3,
upper), although the DBH distribution in Stand K was
smaller than the other three stands (Fig. 4).
Size distribution and state of mortality
The DBH class distributions of canopy trees in 1985 varied
between the stands (Fig. 4). The dominant DBH class was
around 50–60 cm in Stands O, S, and H, whereas smaller
trees were more abundant in Stand K. During the study
period, deaths of canopy trees were observed in all DBH
classes in Stand O. In the other stands, just one canopy
trees died in Stand S, and the trees belonging to smaller
DBH classes died in Stands H and K.
With respect to the dead trees, the classifications of deaths
were mainly recorded as ‘‘standing-dead’’ or ‘‘stump’’
(Table 3).
Spatial patterns of stem mortality
The spatial pattern of the dead canopy trees in Stand O showed
a clumped distribution (Fig. 5). In this stand, canopy trees’
death spread out from the single tree with the broken trunk that
was injured by a severe typhoon in 1991 (Figs. 5, 6).
Fig. 1 Mortality of canopy trees over the 23-year study period in four
old-growth Chamaecyparis stands in the Akasawa Forest Reserve.
Error bar 95 % confidence interval
Fig. 2 Growth rates of BA of surviving canopy trees over the 23-year
study period in four old-growth Chamaecyparis stands in the
Akasawa Forest Reserve. Error bar 95 % confidence interval
138 J For Res (2014) 19:134–142
123
Discussion
Our long-term monitoring study provides precious infor-
mation on the dynamics of old-growth Chamaecyparis
forests. Here, we first summarize the characteristics of each
stand, and then discuss the trends in old-growth Chamae-
cyparis forests over time.
In Stand O, the largest decrease in the stem number and BA
resulted from the sudden death of the single tree injured by a
typhoon in 1991: thereafter, the surrounding canopy trees
began to die. This phenomenon is a typical gap enlargement
process, originating from a single tree falling to form a gap
(Foster and Reiners 1986; Liu and Hytteborn 1991; Yamam-
oto 2000). Such single tree-fall gaps often enlarge as a result of
the trees surrounding the original gap subsequently falling
(McCarthy 2001). Therefore, Stand O may be in the process of
becoming a ‘‘gap-dynamic’’ stand (Watt 1947), or progressing
towards a shifting steady-state (Bormann and Likens 1981).
In contrast to Stand O, Stand S was not severely affected
by the 1991 typhoon, possibly because the stand was not
located on an upper slope, and thus the trees in the stand
could escape from strong winds. In Stand S, very few
canopy trees died, and deaths only occurred during
1985–1990, i.e., before the 1991 strong typhoon.
In Stand K, it has been reported that the Isewan typhoon
in 1959 caused severe damage to this stand (Nagano
Regional Forest Office 1985). During 1985–2008, no
C. obtusa canopy stems died, but stem recruitments were
observed, and, consequently, the stand BA continued to
increase. This suggests that Stand K can be considered in a
state of recovery from the severe disturbance of the
typhoon (Yamamoto 1993a).
In Stand H, the only death of C. obtusa trees was
recorded during 1985–2000, and the BA of C. obtusa trees
gradually increased. Stand H has been considered to be a
well-stocked stand within this reserve (Yamamoto 1993a).
Historically, there has been no record of selection cutting
on this stand (Nagano Regional Forest Office 1985), indi-
cating that Stand H can be considered to have developed
without human disturbances.
Natural disturbance is a major factor affecting the
structure and dynamics of forests (White 1979; Pickett and
White 1985). The stands in this Reserve are no exception;
several typhoons, varying in intensity, have passed during
the last several decades (weather data available online from
Japan Meteorological Agency, http://www.jma.go.jp/jma/
menu/report.html/). So, it is likely that canopy gap cre-
ations and enlargements, owing to natural disturbances like
typhoons, will occur in the future in stands in this Reserve.
Chamaecyparis forests that would experience such distur-
bances may show a stand regeneration pattern relating with
canopy-gap dynamics; Stand K is considered to be in a
state of recovery (low mortality with high AGR), while
Stand O is in a state of a decay (high mortality with low
AGR).
For stands resembling Stands S and H, with a T. do-
labrata understory, Hoshino et al. (2001, 2002, 2003)
suggested that the occurrence of abundant growth of
T. dolabrata does not mean that the species monopolizes
every gap that occurs; certain deciduous broadleaved trees
and a few Chamaecyparis trees occupy these gaps. This
suggests that such stands become more diverse, with a
shifting mosaic of developmental stages (Bormann and
Likens 1981) through gap formations (Asai et al. 2003). It
could be proposed that the stands in Akasawa Forest
Reserve, probably like other old-growth Chamaecyparis
forests in Kiso district, are still in the process of develop-
mental stages.
Some Chamaecyparis species, like C. nootkatensis in
North America, are very long-lived (ca. 1,000 years old)
and show great dominance in several stands (Parish and
Fig. 3 The absolute growth rate of surviving canopy trees over the
23-year study period in four old-growth Chamaecyparis stands in the
Akasawa Forest Reserve. Each box indicates the middle 50 % of
the observed data, while the lower, middle and upper lines for each
box represent the 25, 50, and 75th percentiles, respectively. The lower
and upper bars represent the range equivalent to 1.5 times the extent
of the boxes. Outliers are plotted as empty circles. Different letters
indicate significant differences (P \ 0.05)
J For Res (2014) 19:134–142 139
123
Fig. 4 DBH class distributions
of canopy trees in four old-
growth Chamaecyparis stands
in the Akasawa Forest Reserve
in 1985
Fig. 5 Spatial distribution of
dead and recruited canopy trees
in four old-growth
Chamaecyparis stands in the
Akasawa Forest Reserve. Arrow
indicates the dead canopy tree
with a broken trunk caused by a
strong typhoon, TY19, which
passed along the Japanese
archipelago on 27–28
September 1991
140 J For Res (2014) 19:134–142
123
Antos 2004, 2006). In old-growth forests, C. nootkatensis
often has few individuals of intermediate sizes (Antos and
Zobel 1986), and its growth rates are often very slow since
their recruitment depends on gap creations on a suitable
scale (Antos et al. 2005). Based both on the previous
findings and on our present results, it can be concluded that
the dynamics and regeneration patterns of old-growth
Chamaecyparis forests are greatly affected by natural dis-
turbance regimes, such as typhoons.
Our result showed that the stand-level ingrowth of
300-year-old C. obtusa canopy trees could contribute to the
stock of each stand. This suggested that C. obtusa will
continue to maintain its dominance potentially over hun-
dreds of years (Oda et al. 1997; Kawada 1933), and that
Chamaecyparis forests will turn to the state of diverse and
late successional forests if the present conservation suc-
cessfully prevails.
Acknowledgments We thank the Kiso (Agematsu) District Forest
Office and the Chubu (Nagano) Regional Forest Office for permitting
this study in Akasawa Forest Reserve. We also thank the many
researchers and students who helped our work, including Masao
Katayama, Yojiro Matsuura, Yasuhide Nagayama, Shigeharu
Kogushi, Keita Hamaguchi, Yasuko Moriyama, Nobuhiro Tomaru,
Yuri Nakao, Takahiro Asai, Kouji Kurita, Hajime Kobayashi, Yasu-
hiro Koyama, Daisuke Kabeya and Tomoyuki Saitoh. This work was
supported partly by the Nagoya University Global COE (Center of
Excellence) Program ‘‘From Earth System Science to Basic and
Clinical Environmental Studies (GCOE-BCES) of the Ministry of
Education, Culture, Sports, Science and Technology (MEXT) of
Japan.
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