An introduction to theAya Research Site, a Long-Term Ecological … · 2011-04-04 · dophytes, 5...

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Bull. Kitakyushu Mus. Nat. Hist., 18: 157-180. March 31, 1999 An introduction to the Aya Research Site, a Long-Term Ecological Research site, in a warm temperate evergreen broad-leaved forest ecosystem in southwestern Japan: Research topics and design Tamotsu Sato1*, Yohsuke Kominami1, Satoshi Saito1, Kaoru Niiyama1, Tohru Manabe2, Hiroyuki Tanouchi3, Naohiko Noma4 and Shin-Ichi Yamamoto5 'Warm Temperate Forest Laboratory, Kyushu Research Center, Forestry and Forest Products Research Institute (FFPRI), 4-11-16 Kurokami, Kumamoto 860-0862, Japan. 2Kitakyushu Museum and Institute of Natural History, 3-6-1 Nishihonmachi, Yahatahigashi-ku, Kitakyushu 805-0061, Japan Silviculture Laboratory, Hokkaido Research Center, Forestry and Forest Products Research Institute (FFPRI), 7 Hitsujigaoka, Toyohira, Sapporo, Hokkaido 062-0045, Japan. ^Department of Ecosystem Studies, School of Environmental Science, The University of Shiga Prefecture, 2500 Hassaka-cho, Hikone, Shiga 522-8533, Japan 5Laboratory of Forest Ecology and Physiology, School of Agricultural Sciences, Nagoya University, Furo-cho, Chikusa-ku, Nagoya 464-0814, Japan (Received October 14, 1998) Abstract The Aya Research Site (ARS), a long-term ecological research site, was estab lished in 1989 in a warm temperate evergreen broad-leaved forest, southwestern Japan. Distylium racemosum, Persea thunbergii and Quercus spp. predominate in the ARS, together with a few deciduous broad-leaved species. To quantify ecological parameters of tree populadon and forest dynamics, tree censuses (dbh > 5 cm) and sapling censuses (dbh <5 cm) were conducted in a 4-ha plot in the ARS. Seedling censuses have also been conducted monthly since 1991. In September 1993, a strong typhoon caused uprootings and stem breakages within the ARS. After the typhoon disturbance, tree densities and basal area decreased and the emergence of pioneer species (e.g., Mallotus japonica, Idesia polycarpa and Zanthoxylum alianthoides) were stimulated beneath canopy gaps. This typhoon is considered to have had a major impact on the forest dynamics in the ARS, and long-term observation is needed to evaluate many basic ecological parameters. Key Words LTER Site, Warm Temperate Zone, Evergreen Broad-leaved Forest, Aya, Southwestern Japan * Corresponding author

Transcript of An introduction to theAya Research Site, a Long-Term Ecological … · 2011-04-04 · dophytes, 5...

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Bull. Kitakyushu Mus. Nat. Hist., 18: 157-180. March 31, 1999

An introduction to the Aya Research Site, a Long-Term

Ecological Research site, in a warm temperate

evergreen broad-leaved forest ecosystem in

southwesternJapan: Research

topics and design

Tamotsu Sato1*, Yohsuke Kominami1, Satoshi Saito1, Kaoru Niiyama1, TohruManabe2, Hiroyuki Tanouchi3, Naohiko Noma4 and Shin-Ichi Yamamoto5

'Warm Temperate Forest Laboratory, Kyushu Research Center, Forestry and Forest ProductsResearch Institute (FFPRI), 4-11-16 Kurokami, Kumamoto 860-0862, Japan.

2Kitakyushu Museum and Institute of Natural History, 3-6-1 Nishihonmachi, Yahatahigashi-ku,Kitakyushu 805-0061, Japan

Silviculture Laboratory, Hokkaido Research Center, Forestry and Forest Products ResearchInstitute (FFPRI), 7 Hitsujigaoka, Toyohira, Sapporo, Hokkaido 062-0045, Japan.

^Department of Ecosystem Studies, School of Environmental Science, The University ofShigaPrefecture, 2500 Hassaka-cho, Hikone, Shiga 522-8533, Japan

5Laboratory ofForest Ecology and Physiology, School ofAgricultural Sciences,Nagoya University, Furo-cho, Chikusa-ku, Nagoya 464-0814, Japan

(Received October 14, 1998)

Abstract The AyaResearch Site (ARS), a long-term ecological research site, was established in 1989 in a warm temperate evergreen broad-leaved forest, southwesternJapan. Distylium racemosum, Persea thunbergii and Quercus spp. predominate in the ARS,together with a fewdeciduous broad-leaved species. To quantify ecological parametersof tree populadon and forest dynamics, tree censuses (dbh > 5 cm) and sapling censuses(dbh <5 cm) were conducted in a 4-ha plot in the ARS. Seedling censuses have alsobeen conducted monthly since 1991. In September 1993, a strong typhoon causeduprootings and stem breakages within the ARS. After the typhoon disturbance, treedensities and basal area decreased and the emergence of pioneer species (e.g., Mallotusjaponica, Idesia polycarpa and Zanthoxylum alianthoides) were stimulated beneath canopygaps. This typhoon is considered to have had a major impact on the forest dynamicsin the ARS, and long-term observation is needed to evaluate many basic ecologicalparameters.

Key Words LTER Site, Warm Temperate Zone, Evergreen Broad-leaved Forest, Aya,Southwestern Japan

* Corresponding author

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158 Tamotsu Sato el al.

Introduction

Long-term ecological research (LTER) programs have been set up in manycountries over the past 20 years. Various large-scale studies have been conducted atthese LTER sites to clarify the structure and dynamics of ecosystems occurring at onesite and to compare these patterns with those observed at other sites. Researchat the Hubbard Brook Experimental Forest, a famous LTER site in the U.S., hasprovided much information on vegetation structure and nutrient flux (Bormann andLikens, 1979; Likens and Bormann, 1995). Valuable data on stand productivity andforest-stream interactions have been obtained from the H.J. Andrews ExperimentalForest, another U. S. LTER site (e.g., Long, 1982; Swanson et aL, 1982). Researchdata from these LTER sites have been used to develop new forest resource management policies (Likens, 1989; Hartshorn, 1990; Franklin, 1992). Results fromthese forest sites have also been used in the application of ecological science to theidentification and solution of critical social issues (Franklin, 1994; McDade et aL,

1994;Condit, 1998).

The importance of a global network among LTER sites has been emphasized bymany researchers (Nottrott and Montonen, 1994; Clark and Hamburg, 1994).The exchange of scientists and comparison of data between LTER sites may providethe key to the solution of many social and environmental issues such as globalwarming and acid rain.

In this paper, we present basic information about the Aya Research Site with thehope of promoting collaboration with other LTER researchers.

The reasons for selecting an evergreen broad-leaved forest as an LTER site

Evergreen broad-leaved forests (EBLF) are widely distributed throughout thewarm temperate zone of East Asia and are dominated by genera of Quercus andCaslanopsis with associated rich evergreen tree species such as Persea, Cinnamomum,Ilex, Symplocos and Camellia (Kira, 1991). These forests are found in areas with ahumid, virtually frost-free climate (Ovington, 1983), and are distinguished fromevergreen sclerophyllous forests by leaf morphology, physiognomy, structure andclimate pattern (Ovington, 1983;Satoo, 1983; Hattori, 1985; Song, 1995).

In Japan, geographical distribution ofEBLFare found in the lowlands ofKyushu,Shikoku, the southern half of Honshu and many islands in the northern half of theOkinawa Islands (Kira et aL, 1978; Satoo, 1983). As civilization in Japan spreadfrom the southwest, natural forest was converted to manmade secondary forests

(e.g., coniferous plantations and coppices) and agricultural areas (Prow, 1983;Satoo, 1983). Nowadays, old-growth EBLFonly remain at restricted areas such asBuddhist temples, Shinto shrines and National Parks.

Between 1966 and 1973, integrated ecosystem research on EBLFwas conducted

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An LTERSite in an Evergreen Broad-LeavedForest 159

in the Minamata JIBP (Japan International Biological Programs) Special ResearchArea (Kira et aL, 1978). The research programs at Minamata focused on production ecology; consequently, the JIBP study plots were set up in secondary forest(Kira et aL, 1978). Many phytosociology studies have been conducted in EBLFecosystems, especially in southwestern Kyushu (Suzuki, 1951; Kitazawa et aL, 1959;Sako, 1966; Miyawaki, 1981). However, there have been few studies on forest

dynamics (Naka and Yoneda, 1984;Yamamoto, 1992; Takyu and Ohsawa, 1997) oron primary production (Kimura, 1960) in old-growth EBLF ecosystems because ofthe scarcity of old-growth forests.

Many large LTER plots have been established over the past 10 years in Japan in

order to clarify the structure and dynamics of population and community, especially

in cool temperate forest ecosystems (e.g., Masaki etaL, 1992; Yamamoto etaL, 1995;Hoshizaki etaL, 1997). However, there is a lack of data on forest dynamics in EBLF

ecosystems in Japan. LTER programs conducted in EBLF could provide useful datafor establishing management policies not only in EBLF ecosystems but also in otherecosystems such as those in a cool temperate zone.

Site description

Aya Town is located in MiyazakiPrefecture about 860 km west ofTokyo (Fig. 1).A part of the EBLF at Aya Town is comprised of well-preserved old-growth stage.This forest is under the management of the Kumamoto Regional Forestry Office, theForestry Agency. In 1989, we surveyed Compartment No. 93 of the Takeno NationalForest (ca. 109 ha) and found no evidence ofartificial disturbances. After a preliminary survey, Compartment No. 93 was then registered as an experimental compartment and later became known as the Aya Research Site (ARS). The ARS is situatedon the north slope of a broad ridge running east from Mt. Ohmori (32°04'N,131°09'E). In 1989, we set up a 4-ha plot (200 m x 200 m) on a steep north tonorthwest-facing slope within the ARS (Figs. 2 and 3). Elevation of the plot rangesfrom 380 m to 520 m above sea level.

The dominant tree species in the ARS are Distylium racemosum, Persea thunbergii,Quercus acuta, Quercus salicina and Quercus gilva. A few deciduous broad-leavedspecies such as Cornus controversa and Carpinus tschonoskii and an evergreen coniferousspecies, Podocarpus macrophyllus, are mixed with the evergreen broad-leaved species.The ARS belongs to a transitional zone between Distylio-Quercetum salicinae andLasiantho-Quercetum gilvae associations (Miyawaki, 1981). We enumerated 206

species of higher plants belonging to 78 families in the ARS, of which 20 were pteri-dophytes, 5 were gymnosperms, 24 were monocotyledons and 157were dicotyledons(Appendix 1).

The area around the ARS has a monsoon climate (Fig. 4). The annual meantemperature in the ARS at 495 m above sea level for the period 1994 to 1998 was

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160 Tamotsu Sato et al.

130° E 132° E,

130" E 140' E

32° N

~-

45* N

0 500 km

i 1

-35" N

TokyoJ)

128°!E

ff (^ Kitakyushu \

34° N

East China Sea

150 km

Fig. 1. Location of Aya Town. Mivazaki Prefecture.

Fig. 2. Aya Research Site in 1991, two years before the 1993 typhoondisturbance (photo by H. Tanouchi).

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An L.TER Site in an Evergreen Broad-Leaved Forest 161

"X

Fig. 3. Interior of the 4-ha plot in the Aya Research Site (photo by T. Sato).

14.2°C, with a maximum difference in mean monthly temperature of 20.0°C betweenAugust (24.0°C) and January (4.0°C). Snowfall is rare around the ARS. Thewarmth index was 111, within the range for a warm temperate EBLF class (Kira,1949). Annual precipitation at the Ayakita Prefectural Observatory, 294 m abovesea level, is 3,070mm (Miya/.aki Local Meteorological Observatory, 1951-

1997), and about one third of total rainfall isconcentrated in rainy season, from Juneto July.

According to the Soil Survey Report of the Takeno National Forest (KumamotoRegional FORESTRY Office, 1963), the bedrock around the ARS is shale and sand

stone of Mesozoic formation. A moderately moist Brown Forest soil type, classifiedas Dystric-Cambisol by FAO-UNESCO and as Dystrochrept by the U. S. SoilTaxonomy,is dominant in the ARS. Ohnuki etal. (1995) classified the microtopography of theplot using Tamura's criteria (1987). Figure5 shows the pattern of microtopograhy,with an upper sideslope being the most dominant type in the plot.

Research topics and goals

We are conducting the following research topics:— Population and community dynamics of forests;— Regeneration strategies of tree species;— Response of forest ecosystems to natural disturbance;

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162

0

ov.

3

COv.

Q)Q.

E

cCOo

30 -

c

2 20

10 -

Tamotsu Sato et al.

Aya Research Site (495 m)

14.2 °C 3,070mm[1994-1998] [1951-1997]

500

300

- 80

- 60

- 40

- 20

/^//W/iJ L J L

JFMAMJJASOND

Fig. 4. Climatic diagram of the Aya Research Site. The lowermost data is the monthly meantemperature scaled on the left axis. The uppermost line is the monthly precipitation scaled onthe right axis. The hatched areas below the X axis indicate months in which the minimumtemperature was below 0°C.

EE,co

100 %

ua>

a.

Nutrient cycle in forest ecosystems; andMonitoring of micro-climate conditions (e.g., light conditions and soil temperature) .

The goals for our long-term research projects include:Developing a standard measuring procedure for LTER studies in Japan;Clarifying the ecological traits of an EBLF; andProviding a scientific data for sustainable forest management and criteria ofenvironmental issues at both regional and global levels.

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An LTER Site in an Evergreen Broad-Leaved Forest 163

Research design

Ten-meter grids were marked out in the 4-ha plot in the ARS, and each intersecting point was marked with a numbered yellow plastic tag. Intersecting points wereused as base points for measuring tree location. A 1.2-ha core plot (120 m x 100 m)was established within the 4-ha plot to study saplings, seedlings, seed rain and Htterfall(Figs. 5 and 6). The 1.2-ha core plot contained most of the microtopographical

types of the 4-ha plot (Fig. 5).

500 m

450 m

00 m

200m

|||| Crestslope

Upper sidesiope

F=j Lower sidesiope

\/Vy\ Headmost wall

Head hollow

Footslope

bottomland

I 11.2 ha core plot

Fig. 5. Microtopography of the 4-ha plot in the AyaResearch Site (after Ohnuki etal, 1995). Contour interval is 10 m.

The research designs described below are similar to those used in the otherLTER sites managed by the Forestry and Forest Products Research Institute (FFPRI)in Japan, e.g., the Ogawa Forest Reserve (Masaki et aL, 1992; Nakashizuka et aL,1992) and the Kanumasawa Riparian Research Forest (Hoshizaki etaL, 1997).

a) Tree census

Research Years: 1989, 1991, 1993, 1995, 1997

Research Area: 4-ha plot

Species name, diameter at breast height (dbh), and location of all trees above 5cm in dbh in the 4-ha plot were recorded. Rectangularaluminum tagsmarked withserial numbers were used for identification. The recorded trees were divided into

the following four classes according to height and canopy condition:

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164 Tamotsu Sato et al.

Class 1, trees of more than 20 m in height and with the crown exposed to directsunlight;

Class 2, trees of more than 20 m in height that were suppressed by Class 1 trees;Class 3, trees of between from 10 m and 20 m in height; andClass 4, trees of less than 10 m in height.We regarded Class 1 and 2 trees as canopy layer trees, and Class 3 and 4 trees as

sub-canopy layer trees. The first tree census was conducted in 1989. The results ofthis census and the regeneration behaviour of the main tree species were reported byTanouchi and Yamamoto (1995). Since the first census, we have conducted tree

censuses every two years.

b) Seedling census and seed trapResearch Years: 1991-1998

4 m

4m

Fig.6. Research designof the 4-ha plot. Seedrainand Htterfall have been collectedbytraps (A) atpointsindicated byclosed and open circles (total 263 points). Aseedling census hasbeenalsoconducted monthly in subquadrats B adjoining the litter traps. Class S saplings were onlyrecorded in subquadrats Bat the closedcircles(143points). Class Msaplingswere recorded inBsubquadrats at the closed circles (143points) and in subquadrats Dat the closed triangles (257points) at the intersections of the 10-m grid lines throughout the 4-ha plot. Class L saplingswere recorded in quadrats C at the closed circles and quadrats E at the closed triangles.Classification of the saplings is described in the text.A: trap (each with a 0.5 m2 collection area)B:subquadrat for seedling census and sapling census (4 m )C: quadrat for sapling census in the 1.2-ha core plot (closed circles only; 16 m2)D: subquadrat forsapling census outside the 1.2-ha core plot (4m2)E: quadrat for sapling census outside the 1.2-hacore plot (16m)

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An LTER Site in an Evergreen Broad-Leaved Forest 165

Research Area: 1.2-ha core plotResearch Points: 263 points

In 1991, we set up 263 traps, eachwith a collection area of 0.5 m8 (Fig. 7), andmarkedout seedlingcensussubquadrats (2 mx 2 m) from each trap site (subquadratBin Fig. 6) in the 1.2-ha core plot. These traps with adjoining quadratswere set upat each intersection of the 10-m grid (143 points, shown as closed circles in Fig. 6)and at the center point of each 10 m x 10 m area within the grid (120 points, shownas open circles in Fig. 6) in order to observe seed dispersal patterns (Kominamiet al., 1998), seedling demography, Htterfall production. SinceJune 1991, we havecollected the contents of each trap monthly and sorted them into leaves, branches,flowers and seeds. We also have sun-eyed the emergence and survival of tree seedlings monthly within each of the seedling quadrat.

Fig. 7. Traps in the 1.2-ha core plot (photo byT. Sato).

c) Sapling census

Research Years: 1994, 1996, 1998

Research Area: 4-ha plotResearch Points: 257 points to measure Class L and Msaplings outside the 1.2-ha

core plot, and 143 points to measure all three Classes of saplings(L, Mand S) within the 1.2-ha core plot.

We began a sapling census in 1994 to obtain data for trees of less than 5 cm indbh. Saplings, defined as trees that are less than 5cm in dbh, were divided into thefollowing three classes according to tree height:

Class L, saplings of more than 200 cm in heightbut below 5 cm in dbh;Class M, saplings of between 30 cmand 200 cm in height; and

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166 Tamotsu Sato et al.

Class S, saplings of less than 30 cm in height.Number of Class L saplings were recorded in all 400 quadrats (4 m x 4 m)

marked out at the intersections of the grids in the 4-ha plot (quadrats C and E inFig. 6). Number of Class M saplings were recorded in all 400 subquadrats (2 m x 2m), which placed in the upper left corner of the quadrats (subquadrats B and D inFig. 6). The number of Class S saplings was recorded in the 1.2-ha core plot(subquadrat B in Fig. 6). Tree height for Class M and ClassS saplings and dbh forClass L saplings, were measured. All recorded ClassS saplings had emerged before1991, because all trees that had emerged after 1991 were recorded as 'seedlings' in

the seedling censuses.

Changes in stand structure caused by the 1993 typhoon

On 3 September 1993, a record-breakingstrong typhoon passedover a wideareaofJapan, from Kyushu to northern Honshu (Yamamoto elal., 1994). A maximuminstantaneous wind velocity of57.9 m s"1 was recorded at Miyazaki City, about 30 kmsoutheast of Aya (Yamamoto el al., 1994). The typhoon caused severe damage toparts of the ARS. Figure 8 shows damage to trees in the 4-ha plot. Most of thedamages, i.e., uprooted and stem broken trees, occurred near the border between

Fig. 8. Tree damage in the 4-haplot caused by the 1993 typhoon.(a), an uprooted Distylium racemosum; (b), stem breakage ofa Quercus acuta (photos by T.Sato).

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An LTER Site in an Evergreen Broad-Leaved Forest 167

a) 1992

Fig. 9. Canopy projection diagrams in the 1.2-ha core plot; a) before the 1993 typhoon (1992) andb) after the typhoon (1995). Grid interval is 10 m. Cray crowns indicate canopy layer trees(Class 1and 2), and hatched crowns indicate sub-canopy layer trees (Class 3 and 4).

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168 Tamotsu Sato et al.

the headmost wall and the upper sidesiope. Before the typhoon struck, canopy gapswere scattering around the 1.2-haplot (Fig.9a). The typhoon created new gaps andcaused expansion of the area of the old gaps (Fig. 9b).

The typhoon also caused changes in tree densities and basal area (BA). In1991, a total of 4,668 trees and 54.26 m2 ha"1 of BA were recorded in the 4-ha plot(Table 1). The results of the 1995 tree census, two years after the typhoon, showeda decrease in both stem densities and BA for most species. Densities of Pasaniaedulis, Daphniphyllum macropodum and Cornus controversa had decreased by more than25%. Bellingham et aL (1996) compared mortality and damage in dominantspeciesdue to the same typhoon in Yakushima, and they reported that Pasania edulissuffered a high mortality rate of stems and a drastic decrease in BA. On the otherhand, the number of stems of Castanopsis cuspidata var. sieboldii increased, but thisincrease was due to sprout recruitment. Eurya japonica and Cinnamomumjaponicumalsoshowed a slight increase in density between 1991 and 1995. Somesaplings ofthese species were recruited into the tree census class (> 5 cm in dbh) due to thechange in light conditions around the gaps. A large number of seedlings of somepioneer deciduous species, e.g., Mallotus japonica, Idesia polycarpa and Zanthoxylumalianthoides, emerged in the area beneath the canopy gaps after the typhoon.Emergence ofpioneerspecies beneathcanopy gaps were also reported in other EBLF(Naka and Yoda, 1984; Yamamoto, 1992).

Nakashizuka (1991) pointed out that long-term studies in forest dynamics areimportant toevaluate parameters oftreepopulation andcommunity dynamics aswellas the validity of some ideas (e.g., non-equilibrium theory). We have observedparameters of tree population and forest dynamics in the ARS since 1989. Duringthis observation period, a strong typhoon affected stand structure, causing thecreation of various-sized gaps (Fig. 9); from small gaps, which are thought tofavor shade-tolerant species, to large gaps, which favor shade-intolerant species(Whitmore, 1984; Brokaw, 1985). Naka (1982) pointed out that treefall gapscreated by typhoons were important for the regeneration of several species in theEBLF ecosystem. Environmental heterogeneity due to such disturbances maycreate micro-sites for the coexistence of some specieswithin the same stand. On alarger scale, this heterogeneity is what helps to produce the diversity of the forestlandscape (Foster, 1988b). Many studies have indicated that such episodic eventscan play an important role in forest dynamics over a long period (Canham andLoucks, 1984; Foster, 1988a). Since we have obtained data on forest structurebeforeand after the 1993 typhoon struckthe ARS, we can undertake research to testthese and other ideas about forest dynamics over the next decade.

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Species

Evergreen Broad-leaved Species* Distyliumracemosum* Persea thunbergii* Quercus acuta* Quercussalicina

Cleyera japonica* Persea japonica* Castanopsis cuspidatavar. sieboldii* Quercus gjlva* Camellia japonica* Pasania edulis

* Actinodaphne longifolia* Neolitsea aciculata

* Daphniphyllum macropodum* Cinnamomum japonicum

EuryajaponicaOther species (21 species)Subtotal

Table 1. Composition of dominant species in the 4-ha plot.

1991 1995 1991 1995

No. of

StemBA

No. of

StemBA

SpeciesNo. of

StemBA

No. of

StemBA

3JODeciduous Broad-leaved SpecieJS

1,508 46.31 1,507 47.14 * Cornus controversa 14 1.94 9 1.33 Cfl

142 36.34 122 31.76 * Carpinus tschonoskii 6 1.52 6 1.58ft

5'101 34.65 86 27.82 * Idesiapolycarpa 3 0.87 3 0.86

3

152 26.22 137 21.45 * Diospyros japonica 6 0.69 6 0.67<

1,056 12.88 1,043 12.58 * Cornusbrachypoda 3 0.25 3 0.25n

era

367 11.24 337 10.07 Other species (8 species) 25 0.82 24 0.78 nn

73 10.83 80 9.99 Subtotal 57 6.09 51 5.48 3

CO

338 7.43 38 7.36

357 5.96 354 5.97 Evergreen Coniferous Species a.

144 4.31 103 3.16 * Podocarpus macrophyUus 13 1.26 14 1.27r1ft

2125 3.23 114 3.02 Subtotal 13 1.26 14 1.27

<n

97 2.01 84 1.71 o124 1.96 17 1.26 Grand total 4,668 217.04 4,522 196.02

•nftVI

139 1.90 155 1.75

129 0.42 140 0.48 • All number and BA are shown <on the basis of the 4-ha plot.146 4.01 140 3.75 •UnitofBA:m2/4ha

4,598 209.69 4,457 189.27 • Asterisk shows canopy species

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170 Tamotsu Sato el al.

Acknowledgment

This study was supported by a grant-in-aid for the Bio Cosmos Project (BCP-98-III-A08), administered by the Ministryof Agriculture, Forestry and Fisheries. We aresincerely grateful for the field surveyassistance provided by members ofKyushu TokaiUniversity, Okayama University, Nagoya University, Kyoto University, Osaka CityUniversity, Iwate University, Hiroshima University, Chiba University, Shinshu University, Kagoshima University, Miyazaki University and Kumamoto University. We alsoacknowledge our debt to Mmes. M. Hatomura, Y.Tominaga and R. Fukushima fortheir assistance in data management in the laboratory.

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Appendix 1 List of vascular plants in the Aya Research Site.

Legends1. The ordering of family follows Tagawa and Iwatsuki (1972) for Pteridophytes

and Itow (1989) for Spermatophytes.2. For the nomenclature, we referred to Nakaike (1982) for Pteridophytes

and Ohwi and Kitagawa (1983) for Spermatophytes.3. Japanese names are written in square brackets after the scientific names.4. The abbreviations after the Japanese name indicate the research project

by which the species were recorded during the survey. Species not followedby an abbreviation were recorded during vegetation surveys in the period1989-1998.

5. Abbreviations: TC, Tree Census; SaC, Sapling Census; SeC, Seedling Census;S, Seed traps.

PTERIDOPHYTA

Plagiogyriaceae [Kijinooshida-ka]Plagiogyria euphlebia (Kunze) Mett. [Ookijinoo]Plagiogyriajaponica Nakai [Kijinooshida]Plagiogyria rankanensis Hayata [Takasagokijinoo]

Gleicheniaceae [Uraziro-ka]

Gleichenia japonicaSpr. [Uraziro]Hymenophyllaceae [Kokeshinobu-ka]

Lacosteopsis auriculata (Bl.) Nakaike [Tsuruhoragoke]Pteridaceae [Inomotosou-ka]

Pteris cretica L. [Oobanoinomotosou]

Pteris disper Kunze [Amakusashida]Aspleniaceae [Chasenshida-ka]

Asplenium wrightii Eat. ex Hook. [Kurumashida]Aspidiaceae [Oshida-ka]

Arachniodes aristata (Forst.) Tindale [Hosobakanawarabi]

Arachniodes sporadosora (Kunze) Nakaike [Kobanokanawarabi]Ctenitis maximowicziana (Miq.) Ching [Kiyosumihimewarabi]Diplaziumfauriei Christ [Miyama-nokogirishida]Dryopteris decipiens (Hook.) O. Kuntze [Nachikujaku]Dryopteris erythrosora (Eat.) O. Kuntze [Benishida]DryopterisfuscipesC. Chr. [Marubabenishida]

Leptogramma pozoisubsp. mollissima (Kunze) Nakaike [Mizoshida]Polystichum sp. [Inode sp.]

Polypodiaceae [Uraboshi-ka]

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Colysis elliptica (Thunb.) Ching [Iwahitode]Lepisorus thunbergianus (Kaulf.) Ching [Nokishinobu]Pyrrosia lingua (Thunb.) Farwell [Hitotsuba]

SPERMATOPHYTA

GYMNOSPERMAE

Pinaceae [Matsu-ka]

Abiesfirma Sieb. et Zucc. [Momi] SeC S

Pinus densijlora Sieb. et Zucc. [Akamatsu] SeC STsuga sieboldii Carr. [Tsuga] SeC

Podocarpaceae [Maki-ka]

Podocarpus macrophyllus (Thunb.) Lamb. [Inumaki] TC SaC SeC SCephalotaxaceae [Inugaya-ka]

Cephalotaxus harringtonia (Knight) K. Koch [Inugaya]

ANGIOSPERMAE

DICOTYLEDONEAE

ARCfflCHLAMYDEAE

Myricaceae [Yamamomo-ka]

Myrica rubra Sieb. et Zucc. [Yamamomo] SeC S

Betulaceae [Kabanoki-ka]

BetulagrossaSieb. et Zucc. [Mizume] S

Carpinustschonoskii Maxim. [Inushide] TC SeC S

Fagaceae [Buna-ka]

Castanopsis cvspidata (Thunb.) Shottky var. sieboldii (Makino) Nakai [Sudazii]TC SaC SeC S

Pasania edulis Makino [Matebashii] TC SaC SeC S

Quercus acutaThunb. [Akagashi] TC SaC SeC SQuercus gilva Blume [Ichiigashi] TC SaC SeC SQuercus glauca Thunb. [Arakashi]Quercus myrsinaefolia Blume [Shirakashi]

Quercus salicinaBlume [Urazirogashi] TC SaC SeC SQuercus sessilifolia Blume [Tsukubanegashi] TC SaC

Ulmaceae [Nire-ka]

Aphananthe aspera (Thunb.) Planch. [Mukunoki] SeCZelkova serrata (Thunb.) Makino [Keyaki] S

Moraceae [Kuwa-ka]

Broussonetia kaempferi Sieb. [Tsurukouzo]Ficus erecta Thunberg [Inubiwa] TC SaC SeC SFicuserecta Thunberg var. sieboldii (Miq.) King [Hosobainubiwa] TC SaC

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An LTER Site in an Evergreen Broad-Leaved Forest 175

SeC

Ficus sarmenlosa Roxb. var. nipponica (Franch.et Savat.) Corner [Itabikadzura]TC S

Madura cochinchinensis (Lour.) Corner var. gerontogea (Sieb. et Zucc.) Ohashi[Kakatsugayu]

Morus bombycis Koidz. [Yamaguwa] SaC SeCUrticaceae [Irakusa-ka]

Boehmeria sieboldiana Blume [Nagabayabumao]Pellionia minima Makino [Sansyousou]ViUebruneafrutescens (Thunb.) Blume [Iwagane]

Loranthaceae [Yadorigi-ka]Korthalsella japonica (Thunb.) Engler [Hinokibayadorigi]Taxillusyadoriki (Sieb.) Danser [Oobayadorigi] S

Magnoliaceae [Mokuren-ka]Michelia compressa (Maxim.) Sarg. [Ogatamanoki] TC

Schisandeaceae [Matsubusa-ka]

Kadsura japonica (Thunb.) Dunal [Sanekadzura] SeC SIlliciaceae [Shikimi-ka]

Illicium religiosum Sieb. et Zucc. [Shikimi] TC SaC SeCLauraceae [Kusunoki-ka]

Actinodaphne landfolia (Sieb. et Zucc.) Meisn. [Kagonoki] TC SaC SeCActinodaphne Longifolia (Blume) Nakai [Baribarinoki] TC SaC SeC SCinnamomum camphora (L.) Siebold [Kusunoki] TC SeCCinnamomumjaponicum Siebold ex Nakai [Yabu-nikkei] TC SaC SeC SLindera erythrocarpa Makino [Kanakuginoki] SNeolilsea aciculata (Blume) Koidz. [Inugashi] TC SaC SeC SNeolitsea sericea (Blume) Koidz. [Shirodamo] TC SaC SeC S

Perseajaponica (Sieb. et Zucc) Kosterm. [Hosobatabu] TC SaC SeCPersea thunbergii (Sieb. et Zucc.) Kosterm. [Tabunoki] TC SaC SeC S

Trochodendraceae [Yamaguruma-ka]Trochodendron aralioides Sieb. et Zucc. [Yamaguruma]

Lardizabalaceae [Akebi-ka]

Akebia trifoliata (Thunb.) Koidz. [Mitsuba-akabi] S

Stauntonia hexaphylla (Thunb.) Decaisne [Mube] SeC S

Menispermaceae [Tsudzurafuji-ka]Menispermum dauricumDC. [Koumorikadzura] SeC S

Stephania japonica (Thunb.) Miers [Hasunohakadzura] SPiperaceae [Kosyou-ka]

Piperkadzura (Chois.) Ohwi [Fuutoukazura]Chloranthaceae [Senryou-ka]

Chloranthus glaber (Thunb.) Makino [Senryou] SeC

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Aristolochiaceae [Umanosuzukusa-ka]

Asarum sp. [Kan-aoi sp.]Actinidiaceae [Matatabi-ka]

Actinidia arguta (Sieb. et Zucc.) Planch, ex Miq. [Sarunashi] TC SeC SActinidiapolygama (Sieb. et Zucc) Planch, et Maxim. [Matatabi] TC S

Theaceae [Tsubaki-ka]

CamelliajaponicaLinn. [Yabutsubaki] TC SaC SeC SCleyera japonica Thunb. pro p., emend. Sieb. et Zucc. [Sakaki] TC SaC

SeC S

EuryajaponicaThunb. [Hisakaki] TC SaC SeC SStewartia monadelpha Sieb. et Zucc. [Himesyara] TCTemstroemia gymnanthera (Wight et Arn.) Beddome [Mokkoku] TC SaC

SeC S

Hamamelidaceae [Mansaku-ka]

Distylium racemosum Sieb. et Zucc. [Isunoki] TC SaC SeC SSaxifragaceae [Yukinoshita-ka]

Schizophragma hydrangeoides Sieb. et Zucc. [Iwagarami] TCRosaceae [Bara-ka]

Prunusjamasakura Sieb. ex Koidz. [Yamazakura] TC SeC SPrunus spinulosaSieb. et Zucc. [Rinboku] SeC SRubusbuergeri Miquel [Fuyuichigo]Rubus crataegifolius Bunge. [Kumaichigo] S

Rubus hirsutus Thunb. [Kusaichigo]Rubus minusculus Leveille et Vaniot [Himebaraichigo]Rubus palmatus Thunb. [Nagabanomomiji-ichigo]Rubus sieboldii Blume [Hourokuichigo]

Leguminosae [Mame-ka]Albizia julibrissin Durazz. [Nemunoki] SeCCaesalpiniajaponicaSieb. et Zucc. Qyaketsuibara]EuchrestajaponicaHook. fil. [Miyamatobera]

Euphorbiaceae [Toudaigusa-ka]Antidesmajaponicum Sieb. et Zucc. [Yamahihatsu] TCMallotusjaponicus (Thunb.) Muell. Arg. [Akamegashiwa] SaC SeC

Daphniphyllaceae [Yuzuriha-ka]Daphniphyllum macropodum Miq. [Yuzuriha] TC SaC SeC SDaphniphyllum teijsmanni Zoll. [Himeyuzuriha] TC SaC SeC

Rutaceae [Mikan-ka]

BoenninghauseniajaponicaNakai [Matsukazesou]Evodiameliaefolia (Hance) Benth. [Hamasendan] SeC SSkimmiajaponicaThunb. [Miyamashikimi] SeCZanthoxylum ailanthoides Sieb. etZucc. [Karasuzansyou] TC SaC SeC S

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An LTER Site in an Evergreen Broad-Leaved Forest 177

Simaroubaceae [Nigaki-ka]Picrasma quassioides (D. Don) Benn. [Nigaki] SaC SeC S

Meliaceae [Sendan-ka]

Melia azedarach Linn. [Sendan] SeC

Anacardiaceae [Urushi-ka]

Rhusambigua Lavallee ex Dippel [Tsutaurushi] TCRhusjavanica Linn., pro p. [Nurude] SeCRhussylvestris Sieb. et Zucc. [Yamahaze] SRhus trichocarpa Miq. [Yamaurushi]

Aceraceae [Kaede-ka]

Acer mono Maxim. [Itayakaede] SeC SAcer nipponicum Hara [Tetsukaede] TC SaC SeC SAcerpalmatumThunb. [Irohamomiji] S

Sapindaceae [Mukuroji-ka]Sapindus mukorossi Gaertn. [Mukuroji] TC SaC SeC S

Sabiaceae [Awabuki-ka]

Meliosma rigidaSieb. et Zucc. [Yamabiwa] TC SaC SeC S

Aquifoliaceae [Mochinoki-ka]Ilexbuergeri Miq. [Shiimochi] TC SaC

hex goshiensis Hayata [Tsugemochi] TC SaC SIlex integra Thunb. [Mochinoki] TC SaC SeC SIlex latifoliaThunb. [Tarayou] SaC

Ilexmacropoda Miq. [Aohada] SIlexrotunda Thunb. [Kuroganemochi] TC SaC S

Celastraceae [Nishikigi-ka]Celastrus orbiculatus Thunb. [Tsuru-umemodoki] TC SeC S

Rhamnaceae [Kuroumemodoki-ka]

Berchemia racemosa Sieb. et Zucc. [Kumayanagi] TCBerchemia racemosa Sieb. et Zucc. var. magna Makino [Ookumayanagi] TC

Vitaceae [Budou-ka]

Ampelopsis brevipedunculata (Maxim.) Trautv. [Nobudou]

Ampelopsis cantoniensis (Hook, et Arn.) Planch [Udokadzura] S

Parthenocissus tricuspidata (Sieb. et Zucc.) Planch. [Tsuta] TC SeC S

VitisJlexuosa Thunb. [Sankakudzuru] TC

Elaeocarpaceae [Horutonoki-ka]Elaeocarpusjaponicus Sieb. et Zucc. [Kobanmochi] SaC SeC S

Thymelaeaceae [Jintyouge-ka]DaphneKiusiana Miquel [Kosyounoki] SaCWikstroemia trichotoma (Thunb.) Makino [Kiganpi]

Elaeagnaceae [Gumi-ka]

Elaeagnus glabra Thunb. [Tsurugumi] SeC S

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178 Tamotsu Sato el al.

Elaeagnus pungens Thunb. [Nawashirogumi]Flacourtiaceae [Iigiri-ka]

Idesia polycarpa Maxim. [Iigiri] TC SaC SeC SXylosma congestum (Lour.) Merr. [Kusudoige] SeC S

Myrtaceae [Futomomo-ka]

Syzygium buxifolium Hooker et Arnott [Adeku] TC SaC SeC

Cornaceae [Mizuki-ka]

AucubajaponicaThunb. [Aoki]

Cornus brachypoda C. A. Mey. [Kumanomizuki] TC SeC SCornus controversa Hemsl. [Mizuki] TC SaC SeC S

Araliaceae [Ukogi-ka]Aralia elata (Miq.) Seemann [Taranoki] SDendropanax trifidus (Thunb.) Makino [Kakuremino] TC SaC SeC S

Umbelliferae [Seri-ka]

Hydrocotylejavanica Thunb. [Oobachidome]

METACHLAMYDEAE

Pyrolaceae [Ichiyakusou-ka]

Monotropastrum globosum H. Andr. ex Hara [Ginryousou]Ericaceae [Tsutsuji-ka]

Vaccinium bracteatum Thunb. [Syasyanpo] SaCMyrsinaceae [Yabukouji-ka]

Ardisia crenata Sims [Manryou] SeC SArdisia japonica (Thunb.) Blume [Yabukouji]Maesajaponica (Thunb.) Moritzi [Izusenryou] SeC SMyrsine seguiniiL6\. [Taimintachibana] SaC SeC

Primulaceae [Sakurasou-ka]

Lysimachia sikokiana Miq. [Morokoshisou]Ebenaceae [Kakinoki-ka]

Diospyros japonica Sieb. et Zucc. [Ryukyumamegaki] TC SaC SeC SDiospyros morrisiana Hance [Tokiwagaki] TC SaC SeC S

Styraceae [Egonoki-ka]Styraxjaponica Sieb. et Zucc. [Egonoki] S

Symplocaceae [Hainoki-ka]Symplocos glauca (Thunb.) Koidz. [Mimizubai] TC SaCSymplocos landfolia Sieb. et Zucc. [Shirobai] TC SaC SeC SSymplocos lucida Sieb. et Zucc. [Kuroki] SaC SeCSymplocos myrtacea Sieb. et Zucc. [Hainoki] TC SaC SeCSymplocos prunifolia Sieb. et Zucc. [Kurobai] TC SaC SeC SSymplocos theophrastaefolia Sieb. et Zucc. [Kanzaburounoki] TC SaC S

Oleaceae [Mokusei-ka]

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An LTER Site in an Evergreen Broad-Leaved Forest 179

Ligustrumjaponicum Thunb. [Nezumimochi] SeCOsmanthus insularis Koidz. [Nataorenoki] TC SaC SeC S

Gentianaceae [Rindou-ka]

Gentianazollingeri Fawcett [Fuderindou]

Apocynaceae [Kyouchikutou-ka]

Anodendron affine (Hook, et Arn.) Druce [Sakakikadzura] TC SeCTrachelospermum asiaticum (Sieb. et Zucc.) Nakai var. intermedium Nakai

[Teikakadzura] TC SeC S

Asclepiadaceae [Gagaimo-ka]Marsdenia tomentosa Morren et Decaisne [Kijyoran] SeCTylophorajaponicaMiq. [Tokiwakamomedzuru] TC SeC

Rubiaceae [Akane-ka]

Damnacanthus indicus Gzertn. fil. [Aridooshi] S

LasianthusjaponicusMiq. [Ruriminoki] SeC SLasianthusjaponicusMiq. var. satsumensis (Matsum.) Makino

[Satsumaruriminoki]

OphiorrhizajaponicaBlume [Satsumainamori] SeCPaederia scandens(Lour.) Merrill var. maird (Leveille) Hara

[Hekusokadzura] S

Pseudopyxis depressa Miq. [Inamorisou]Randia cochinchinensis (Lour.) Merrill [Misaonoki] SaCUncaria rhynchophylla (Miq.) Miq. [Kagikadzura] TC

Verbenaceae [Kumatsudzura-ka]

CallicarpajaponicaThumb. [Murasakishikibu] SeCCallicarpa mollis Sieb. et Zucc. [Yabumurasaki] SeC SClerodendron trichotomum Thunb. [Kusagi] SeCPremnajaponica Miq. [Hamakusagi] TC SeC S

Loganiaceae [Fujiutsugi-ka]Gardneria nutansSieb. et Zucc. [Houraikadzura]

Caprifoliaceae [Suikadzura-ka]

Lonicera hypoglauca Miq. [Kidachinindou] TC SeC SLonicerajaponicaThunb. [Suikadzura] TC S

Sambucus sieboldiana (Miq) Blume ex Graebn. [Niwatoko] SeCViburnum awabuki K. Koch [Sangojyu] TC SaCViburnum dilatatum Thunb. [Gamazumi] S

Valerianaceae [Ominaeshi-ka]

Patrinia villosa (Thunb.) Juss. [Otokoeshi]Compositae [Kiku-ka]

Adenostemma lavenia (Linn.) O. Kuntze [Numadaikon]Carpesium dibaricatum Sieb. et Zucc. [Gankubisou]Carpesium koidzumii Makino [Hosobagankubisou]

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180 Tamotsu Sato et al.

Crasscephalum crepidioides (Benth.) S. Moore [Benibanaborogiku]Lapsana humilis (Thunb.) Makino [Yabutabirako]

MONOCOTYLEDONEAE

Liliaceae [Yuri-ka]

Lilium cordatum (Thunb.) Koidz. [Ubayuri]

Smilax china Linn. [Sarutoriibara] SeC S

Dioscoreaceae [Yamanoimo-ka]

Dioscorea bulbifera Linn. [Nigakasyu]Dioscorea tokoro Makino [Onidokoro]

Burmanniaceae [Hinanosyakujyou-ka]Burmannia cryptopetala Makino [Shirosyakujyou]Burmannia liukiuensis Hayata [Kirishimasyakujyou]

Gramineae [Ine-ka]

Miscanthus sinensis Anderss. [Susuki]

Oplismenus undulatifolius (Ard.) Roemer et Schultes var. japonicus (Stend.)Koidz. [Chijimizasa]

Araceae [Satoimo-ka]

Arisaema sp. [Tennansyou sp.]Zingiberaceae [Syouga-ka]

Alpiniajaponica (Thunb.) Miq. [Hanamyouga]Orchidaceae [Ran-ka]

Bulbophyllum drymoglossum Maxim. [Mamedzutaran]Bulbophyllum inconspicuum Maxim. [Mugiran]Calanthe aristulifera Reichb. fil. [Kirishimaebine]Calanthe reflexa Maxim. [Natsuebine]Cymbidium goeringii (Reichb. fil.) Reichb. fil. [Syunran]Dendrobium moniliforme (Linn.) Sw. [Sekkoku]Eria reptans (Franch. et Savat.) Makino [Osaran]Galeola septentrionalis Reichb. fil. [Tsuchiakebi]Lecanorchis sp. [Muyouran sp.]Liparis nervosa (Thunb.) Lindl. [Kokuran]Neofinetiafalcata (Thumb.) Hu [Fuuran]Phaius minor Blume [Gansekiran]

Saccolabiumjaponicum Makino [Kashinokiran]Sedireajaponica (Lindenet Reichb. fil.) Caray et Sweet [Nagoran]