Comparison - J-STAGE

The Society of Vegetation Science

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The Society ofVegetation Science

Pegetation Science 2Z: 1-14, 2005

e e wrsme

Comparison

natural, and

Japan

of species

secondary

Hiroaki ISHIDAi, Tamotsu

and Yoshiaki TAKEDA2

composition and richness

lucidophyllous forests on

HATTORIi

i

es'eeiasimgig#i･ i. s-wi-ew,,l

among primeval,

Tsushima Island,

ilnstitute ot'Natural und Environmenta1 Sciences, Unii,ersity of Hyogq Sanda

669-1546, JapanZI:acutty

ot' Hurnan Deyelopmcnt, Kebe Universil},, Kobe 6S7'OOj l, Japan

Other than artificiul forests, three major types oflucidophyllous (evergreen broad-leaved) forests are tbund in

Japan, accoTding to the degree of human impact: (1) primexral; (2) natuTal-those that have been subjectedto occasional low-impact hurnan disturbance, such as tree or understory cuuing, and to a high degree offragmentation by human development ; and (3) secondary-coppices that was used as a source offire",ood and

charcoal. In this stud),, species composition and richness w'ere compared among primes,al, natural, tLnd

secondary lucidophyllous t'orests on Tsushima Island, Japan, to claTify the effects of human disturbance and

forest fragmentation. The vegetation of 76 plots of 1oo m2 (30 prjmeval, 20 natural, and 26 secondary plots)was investigated. The results ofdetrended correspondence ttnalysis showed that : (1) the spec{es compositionclearly difiered among the forest types ; and (2) the species composit{on ofthe natural forests was in betweenthe compositions ofthe primeval and secondary forests, showing similarities to both, The dilkrences inspecies composition among the three fbrest t)pes seemed to greatly refiect the degrees of both past hutnan

disturbanee and forest fragrnentution. Distyfium racemosum, which hus a low coppicing abitity, dominatedin the primeval forests, but this ",as not or rarely the case in the natural and secondary tbrests, Speciesrichness (number of species per 1oo m2) differed signifieantSy among the forest t}pes, fi'om highest (primeval)to lowest (secondary). The number of al1 species per 100 m2 did not haye a clear relationship with forestfragrnentat{on. However, the nuinber of species whose disnibutions were biased toward the prirneval and

natural forests was highly and positively correlated with patch urea of the primeval and natural forests.

suggesting that fbrest fragmentation is partly responsible fbr the ditierences in species richness among the tbrest

types.

Key words : forest fragmentation,human disturbance, lucjdophy[lous forest, prjmeva]'lorest, species richness

ew INTRODUCTION

Lucidophyllous (evergreen bread-leaved) forest,dominated by evergreen Fagaceae or Lauraceae species, is

a representative forest type in Japan. These fbrests once

covered most of the lowlands west of the Tohoku region,but the few primeval forests thut remain can be found only

in Miyazaki Prefbcture, Kagoshima Prefecture, and the

isrands of Tsushima, Iriernotejima, \akushima, and Mi-kurajima (Hattori & Asami 1998), While old-growth

]ucidophyllous forests remain in such places as shrines and

temples, most are small and isolated. Because these frag-mented fbrests are subjected to some degree of humandisturbance, such as tree cutting understory cutting, and

foot traMc, they are not strictly pristine. On the Qther

hand, extensive managed lucidophyllous coppices can befbund along the Pacific coast west of the Boso Peninsula

(ltow & Kawasato 197g; Suzuki 1982; Miyawalci &

Okuda 1990), Thus, there are several types of lucidophyl-lous forests in Japan, according to their degree of human

lmpact.

Hattori & Asami (1998) roughly classified luc{dophyi-lous forests in Japan (excluding anificial forests) into three

types according to their naturalness: (1) primeval, with

few detectable human disturbances and with original

conditions persisting; (2) natural, in which no periodic

clear cutting or understory cutting has occurred, but wherethere has been occasional, low-impact human disturbanceas well as a high degree of fragrnentation by humandeve]opment (the shrine!temple remnant forests represent

this type); and (3) secondary, including eoppices thatwas used as a source offirewood and charcoal. As far as

primeval and natural forests are concerned, many studies

have been conducted to clarify species composition (Suzu-ki & Suzuki ]973;Miyawaki et aL 1978;Yamanaka]979; Hattori 1985), species diversity (ltow 1984; It6

The Socieb, of Vegeration Science asReccived February 20, 2004/Acceptcd Septembei' 28, 2004

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2 Vbgemtion ScienceVel.22, No, 1. 2005

1997;Ishida et aL 1998), dynamics (Naka 1982;

Yamamoto 1992), and habitat conditions (Hattori 1992;

Hattori et al, 2003). However, no study has clearly

defined primeval and natural forests and then thoroughly

compared the two, Consequent]y. the efiects of human

disturbance or forest fragmentation on the species composi-

tion and diversity of natural Iucidophyllous forests huvenot been f'u[ly elucidated (lshida et al, 1998; Hattori &

lshida 2ooO). These efibcts must be clarified if we ai'e to

correctly understand the actual state of natural lucidephyl-lous forests and establish forest protection nieasures.

While the distribution and species composition ofsecon-

dary lucidophyllous tbrests have been analyzed (Yamana-ka 1969; MiyawakH980; Itow T9g3; Isogai 1994), there

have heen few quantitative studies on species diversity

(Ishida & Hattori 1998 ; Aiba et al. 200I). In addition,

although it would appetLr reasonable to assume that the

species composi{ion and diversity ofsecondary lucidophyl-lous fbrests difi'er fi'om those of primeval or natural lueido-

phyllous fbrests due to human disturbances such as clear

cutting and understory cutting (Itow 1981a), there has been

insurncient Tesearch to confirrn this.

We sought to clarify the eil'ects of human disturbance

and forest fi'agmentation on the species composition and

riehness of lucidophyllous forests on Tsushima Js]and inJapan. The specific objectives ofthis study were: (1) to

identify the di[ferences in species composition and richness

(number of species per unit area) among primevul, natural,

and secendary lucidophyllous forests; and (2) to exam-

ine the relationships between these diiferences and both

human disturbance and forest fragrnentation,

three types were anaiyzed and compared (Fig, 1). A largeforest patch (ca. 70 ha) on the northern slope ofMt. TateTa

in the town of lzuhara served as a sampling site for

primeval lucidophyllous fbrest. This primeval 'fbrest

bas

been designaled as a national natural monument, as a

special protection area inside the lki-Tsushima Quasi-National Park, and as a national forest for the protectionof its biological and genetic resources, To examine the

eflctts of forest fragmentation on species composition and

richness, fifteen forest patches that preserved in the pre-

cincts of shrines, ranging in area from O, 17 to 4.4 ha, servedas the sampling sites fbr natural Iucidophyllous fbrests.

We chose natural foresls whose climatic ¢onditions were

similar to those of the primeval and secondary 1ucidophyl-

lous forests. Also, to eliminate the efTbcts ofstrong coastal

winds, only natura] tbiests that were at least 50 m C'rom the

36" -iliV'

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#g MATERIALS AND METHODS

Study s;tes

Tsushima, an lsland that was once parL oi' mainland

Asia, is ]ocated in the Sea ofJapan between the Kyushu

Island and the Korean Peninsula, The island's total area

is 682 km2, and its highest point is 649 m. The island is

composed mainly ot' Tertiary sedimentary rocks of the

Tsushima group (mudstone, sandstone, etc.), but in some

areas, granite, quartz porphyry, and other igneous rocks are

found. The island is in a warm-temperate climatic zone,

The island's lucidophyllous forests are dii,'ided into two

altitudina[ zones (Itow & Nakanishi 1987) : a ([lastanopsis

zone at lower altitude (< ca, 350 m) and a 2uercus zone

at higher altitude (> ca. 350 m),

ln this study, the lucidophyllous forests in the Castanopsts zone on Tsushima Is. were classified into three types-

primeval, natural, and secondary-according to the

classification system of Hattori & Asarni (1998). These

O 5 10km-

1-30

e

!gll32ix2`ic

36

51-76

,48

37

>lf

40, 45

Fig. L Map sho",ing the study sites. Numera[s (1'76) indicate stand numbells in Appendix 1, (A), Primeval forest;(e). Natural forest;(-). Secondar}r forest,

Name of locations : numerals 1 30, Uchiyama ; 31-32,

Kisaka; 33 34, Nii; 35'36, Tsutsu, 37-38, Shilne: 39,

Tsutsunaiin;40, Kechit 41-42, Ogata;43, So;44,

Kokubu; 45, Kofunegoshi; 46, Nii: 47, Kusuho; 48,

Ashiura; 49, Kaiguchi; 50, Tsutsu; 5I-76, Uchiyama.

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Species eomposition and richness of lucidephylleus forests en Tsushima lsland 3

coast were examined. A coppice (ca. 40 years old) on Mt.

Kayaba, near Mt.Tatera, served as a sampling site fbr

secondary Iucidophyllous fbrest, This tbrest was not frag-

mented. CIear cutting had been eonducted in this forest

every 15 to 20 years for production of firewood and char-

coal, but the forest has been abandoned for about the last

40 years because its utility value had been degraded,

Methods

The fietd sunrey was conducted from May 1998 to June

1999. In each primeval, natural, and secondary forest,vegetation was investigated by establishing plots on slopes

of less than 35 degrees where tall evergreen broad-leaved

trees were the dominant species. Ridges and valleys were

excluded. Each plot was 1oom2 (10m × 10m). Whiletwo plots were set in each of five relatively large natural

fbrest patches, only one plot was set in each of the other,

smaller natural forest patches, In total, 76 plots (30 prime-val plots, 20natural plots, and 26secondary plots) were

studied.

Each plet was divided i"to different fbrest layers, and

the height and coverage (%) of each layer were measured.

All species of vascurar plants were Iisted for each layer, and

the coverage (%) efeach spocies was estimated. For some

species there was only one seedling, so to simplify data

processing the minimal unit of coverage was set at O,Ol%.

Epiphytes that grew in the tree and sub-tree layers wereexamined using a pair of 12X binoculars, The habitat

conditions "'ere recorded fbr each plot, topography, slope

degree, slope aspect, and soil condition,

To clarify the natufal environmental conditions of the

stands, we determined six natural environunentat variables i

altitude (ALT), mean temperature of the coldest rnonth

(MTCM), annual precipitation (AP), distance from the

nearest coastline (DFNC), slope degree (SLD), and slope

aspect (SLA), MTCM and AP were estimated from therecords of the closest meteoTological station (Meteorologi-cal Agency I958, 1959), The temperature decrease with

elevation was set at O.6DCflOOm, The runges in ALT,MTCM, AP, DFNC, and SLD for all stands were 15-200m, 3.4-4.5"C, 1489-2]09mm, O.05-3.9km, and 5-35degrees, respectively (Appendix 1).

Data analysis

To clarify the ditferences in species composition among

the forest types, the ordination of stands was conducted bydetrended correspondence analysis (DCA; Hill 1979)based on the presence or absence of species with overal1

fi-equencies of at least 5%.

The species identified in the plots were divided into two

groups aocording to the classification system of Hattori &

Minarniyama (2oo1): the component spocies of' lucido-

phyllous forests (lucidophyllous elements) and others

(non-lucidophyllous elements). The lucidophyllous ele-

ments aTe mostly shade-toierant species that can growunder the closed canopy of lucidophyllous forests. The

non-]ucidophyllous elements are relatively shade-into]er-

ant, mostly deciduous, and comprised of pioneer species

(e,g., AraiZa elata, Madotus juponicus, Athiaia juthtissin,Rhtas javanica, Rhus sybesttis, Euscaphis J'qponica,Zanthexylum piperitum) and spocies showing a strong

habitat preference fbr grassLand, forest edge, canopy gap, or

summergreen forest (e.g., Midettia japonica, Dioscorea

japonica, ettercus serrata, Carpinus tschonoskii, Rhus

tribhocarpa, Styrax japonicus, daiZiearpa moi?Zy, dni2lita-

pa .faponica, Rrunus jumasakura, }Visteria .fombundo).The lucidophyllous e]ements were subdivided intQ the

foIIowing seven lit'e fbrms: trees, shrubs, ground herbs,

grouncl ferns, saprophytes, epiphytes, and c]imbers, The

Iife forms were classified according to the life-forrn system

of Miyawaki et al. (l978), Species richness was evaluated by the number of species

per plot (per unit area). To clarify the differences in

species richness among the forest types, the mean number

of species per pLot fbr each of Che above species groups was

calculated fbr each forest type.

Using simple correlation analysis, we examined the

relationships between the stand seore obtained by DCA or

the number of species per plot and both the natural

environmental variables and the area of a fbrest patch,For this analysis, the SLA was categorized using a three-

grttde index (NW-NE, 1; NE-SE, 2; NW-SW, 2; and

SE-SW, 3), according to the amount of light,

The Kruskal Wallis H-test was used to test for statisti-

cally significant diflbrences in the mean height and mean

eoverage of each forest layer and the mean number of

species per plot among the three tbrest types. When there

were significant diferences among the three types, multiple

comparisons were conducted using the Mann-Whitney U-

test with Bonferroni correction,

es RESULTS

Stand structure The structure of the natural stands was more similar to

that of the primeval stands than to that of the secondar}J

stands (Table 1). There were great differences in the mean

height of the tree layer and the mean coverage of the herblayer between the pyimeval or natural stands on the one

hand and the secondary stands on the other.

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The Society ofVegetationScience

4

Table 1.

VegetationSeieneeVol, 22. No. 1, 2005

Comparison ot' height and coverage of eachfbrest layer among the three forest types,See Appendix t for abbreviations.

}'orcst layerPR NA SE (1) (2)Mean SD.Mean S,D.MeanS.D. All typesPR and NA PR and SE NA and SE

Height {m) Tree layer

Subtree laver"

First shrub layer

Second shrub laver J Herb Laver "

19.813.S7.42.4O,52.52.ll.OO.5O.117.810.56.32.le.63.52.3L5G.2O,113.310.07,222O,41,6 p<O.OOI].2 p<O.oo1O,9 p<O.05

O.4 p<O.05

O,1 p<O.OOI

p<O.05 p<O.Ol

p<O,Ol p<O.Ol

p<O,05 n.s,

p<O,OS n,s,

n,s, p<O,OI

p<o.otn.s.n.s.n.s.p<O,Ol

Coverage (%) Tree layer

Subtree Iayer"

First shrub layer

Second shrub layer

Herb Layer

81.9 14,O

33.6 16,9

22,l ll,6

20.9 10.8

48,O IS.l

91.2 7.1

24.1 16.E

33,O 19.9

25.7 12.6

62.8 16,6

94.419,263.S13.S15,352 p<O.oo]

8,l p<O,0518.4 p<O.OOI6.5 p<O.Ol7,5 p<O.OOI

p<O.05n.s,n.s.n.s.P<O.05p<O.Olp<O.05p<O.Olp<O.05p<O.Oln.s.ns,p<O.Olp<o.oip<O.Ol

S.D. / Standard deviution.S/

Values for stands with subtree Iayer (29 primeval s[ands, l7 natural stands, and 10 secondary stands).

(1 ) : Significant level ot' the difference annong the thrce forest types by the Kruskal Wallis H-test ; (2) idifference between the two tbrest types by the Mann-Whitney U-tesL with Bonferroni eorrectioi, n.s, :Significant

level of the

not significant.

Species composition

The dominant species were Distyfium racemosum and

Castcrnqpsis cuspidota var, sieboldti for the primeval stands,

CL cuspidtita var. sieboidil t'or the natural stands, and C

eztspiddta var. sieboidti and {2uercus salitina for the secon-dary stands (Table 2).

Eigenvalues fbr the first to fourth axes obtained by DCA

were O.320, O.110, O.080, and O.061, respectively. Appendix1 shows the stand scores of axes1 and2 for ea ¢h stand, and

Fig.2 shows the stand ai]angement based on the stand

scores of axesl and 2 for each forest type, Along axis 1,

the three forest types were arranged in the order from leftto right ofprimeval. natural, and secondury. In aU stands,

the stand score of axis1 had no high correlation with

ALT, MTCM, AP, DFNC, or SLD, but it did have a highco"'elation with SLA (Table3). Howeyer, when we

examined the relationship between the stand seore of axis

1 and SLA fbr each t'orest type, we fbund no sigriificant

coiTelations between thern in any fbrest type (p>O.05).On the other hand, the stand score of axis2 in all standshad no high correlations with any of the six variables.

To clarify the eff'ect of forest fragmentation on the

species composition ofthe primes,al and natural forests, sN'e

examined the relationships between the scores of axes 1 and

2 and the fbrest patch area for each of the fo11owing two

stand groups : a natural stand group (all natural stands)

Table2, Specics composition of

frequency (%), and numericultrcc

laycr for each fb]'est type. Numerical i,alue shows

value in paTentheses shows the mcan value of coverage peroccurrencetoo m2 (%).

SpeciesSpecies

typF..

PR NA SENo. ofp[ots

Ca,stanopsiv cuspidata s,ar, sieboldii

euereus salicina

Disiylium racemosum

euercus acr{taPersea thunbergii

euercus serrataPt'unus jamas'akuraDendropanax tttlidus

da,era J'oponicaRhus succedanea

SJ'nlploeos iu(･ido

Athizia .iulibrissin(Rest is omitted)

L{e)

L(e)

L(e)

L(e)

L(c)NL(d)NL(d)

L<e)

L(e)NL(d)

L(c}NL(d)

36,7

3.380.0

.6.7

.

.16J16,7

.

.

.

(21,7)C O.5)(45.9)

C ].5)

( 2.5>( 3.2)

90.0 {62,7)30.0 ( 9.5)

5.0 ( l.O) .2o.o

{ g.3)

'

.

.15.0

C 2,2) . .

100 (48,9)100 {25.9) .38,5

(10.0)15.4 ( l,4)34.6 { 4.2)30.8 ( 2.4)3.8 ( O.2) --7.7

( O,3)15.4 ( 1,9)15.4 ( 1.5)

5533251010986s544

L: Lucidophyllous elements:NL ' Non-lucidephyllous elements ;e/ Evergreen; d/ Deciduous.

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Specics composition and richness of lucidopbyllous forests on Tsushimu Island 5

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180

160

140

120

1OO

80

60

40

20

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Table4. Simple correlution coeMcients between the

stand scores obtained by DCA or the number of

species per 100 m2 and the fbrest patch area for the

primeval and natural forests.

ArcaLog area

N'atural stund group Stand score Axis 1

Axis2

rvumber of species per 100 m2

All species

Lucidophy]lous eLement

Non-lucidophyllous element

". .. Species groups A und B

-O,543･k-O.O13

-O.040

O.067-O.174

O,537*

-O,418

O.029

-O.089-O.060--O.079

O,383

O 50 100 150 200 250 DCA axi$ 1

Fig. 2. 0rdination diagram ofstands using scores for the

first and second axes ef detrended correspondence

analysis (DCA). (A), Primeval forest; (e), Natura]

forest ; (I), Secondary forest,

Primeval-natural stand group

Stand score

Axis 1

Axis 2

Numbcr of species per IOO m2

All species

Lucidophyllous clement


Species groups A and B

-O,931*t-O.021

O.709*s O.85.1,ks

-O,504**

O.937**

･-O,928*"-O,O]3

O,6T3*e e.816x,*-O,5oo*.

O.929*i:

*,p<O.05;** p<O.OOI.

and a primeval-naturul stand group (combination of al1

primeval stands and a]1 natural stands), The results

showed that the score of axis 1 had a significant correlation

with patch area in both tbe natural stand group and the

primeval-natural stand group (Tttble4), The correlation

was particularly high with the primeval-natural stand

group (F'ig. 3).

To examine the changes in species distribution accord-

ing to the arrangement pattern of forest types along DCA

axis 1, the occurrence frequency (%) und the mean value of

total coverage per plot (%) of each species were calculated

for each tbrest type and then compared among the types,

This analysis distinguished five species groups : (A) thosebiased toward primeval stands ; (B) those biased toward

prjmeval and natural stands; (C) those biased toward

natural and secondary stands ; (D) those biased toward

secondary stands; (E) Lhose commonly occurring in the

three forest types (Table5), The basis of this species

ciassification was the phytosociological criterion for distin-

guishing an exclusive species CSasaki 1973),

Species richness

In all stands, the number of spec.ies per plot for al1

species and that fbr lucidophyllous elements had high

correlations with SLA (Table3). However, when we

examined the relationship between the number of species

per plot fbr each of the two species groups and SLA fbr

each forest type, we fbund no significant cen'elations

between them in any forest type (p>O.05), The number of species per plot for all species and that

fbr Lucidophy[lous elements had significant positive corre-

Table 3. Simple correlation coefiicients bet",een the stand scores obtained by DCA or the number ofspocies per 1oomZ and natura] environmental variab]es, See Appcndix 1 for abbrcviations,

ALT MTC:M AP DFNC SLD SLA

Stand score

Axis1

Axis 2

O,170-O,030-O.l74

O,320*s

O.059--O.101-O.IS9--O.064 O,267e-O,283.

Number of specjes per 100 m2

At[ species

Lucidophyllous element


O,022

O,061-O.138

O,130O.0370277: O,130

O.I82-O.2]7 0299**

O.3ss*e-O.295*

-O,372*#-O.387**

O.164

O.596.**

-O,089

-O.624*.*-O.668***

O.343**

*:p<o.os,s* p<o.oi,#**:p<e.(nl.

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6 Vegetation ScienceVol. 22,No, 1, 2005

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120

1OO

80

60

40

20

o

.te

e

...

.--

..fe

.

.

..

..sll

O.1 1 10

Area (ha)

Fig.3. Relatjenship between the stand

axis 1 and the forest patch area (log primeval and naturul forests.

s--ll100

score of DCA

scale) fbr the

lations with the forest patch area in the primeval-natural

stand group, but not in the natural stand group (Table 4),

For all spocies, there were significant diffbrences in the

mean number of species per plot arnong the three forest

types (Table 6). This was true also for the mean number

of species per plot fbr lucidophyllous elements. Thenumber of species per plot for non-[ucidophyllous ele-

ments differed significantly between the primeval and

natural stands and between the primeval and secondary

stands, but not between the nutural and secondary stands

(p>O,05). Of the lucidophytlous elements, trees were the

most prevalent life form in all fbrest t}pes. There was a

significant difference in the number of lucidophyllous-

element tree spocies per piot between the primeval and

natural stands, as weli as between the primeval and secon-

dary stands, but not between the natural and secondary

stands (p>O.05).

Table 5. Species groups classified by sirnilarities in distributional patterll among the three foresr types. Numerical

value shovvs oceurrence frequency (%), and mimerical value in parentheses sho",s the mean vaLue of total coverage

per IOO m2 (%).

SpeciesSpecies

typePR NA SE

No. ofplots

Species group A

Distylium racemosttm

Michelia conu)i'essa

Anodendeen q17fneActinodaphne longij2,fia

Daphniphythim teiismannii

Podocarpus macrophyaus

Euehresta j'aponiea

S)implocos prunijbtiaMarsclenia tomentosa

Daphne kiusianaiWonotropastrum globosum

Desmodium laxutn

Species group B

Damnacanthus indicus

Arachniodes sporadbsora

Elaeagnus glabra Ikzx integra

72?rnstroemia gymnanthera

Cle.yera ,iaponica Neontsea sericea Arachniode,s artytata

LemmaphyUitm tnicrophyUum

Ardisia cretzata

Attcuba japonica Gardneria nutansSpecies group C

Liriope pkr4phytla

L(e)L(e)L(e)L(e)L{e)L(e)L(e)L(e)L(e)L(e)L(d)L(e)96.7 (76,1)83.3 ( O,6)80.0 ( 1.1)86J ( O.6)SO.O ( O.7)70.0 ( 1.6)73.3 ( 02)50,O ( ].1)SO.O (O.05)43.3 (O.05)33,3 (O,03)30.0 (+)

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5.0

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........'...30292827252222161614119

L(e)L(e)L(e)L(e)L(e)L(e)L(c)L(e)L(e)L(e)L(e)L(e)tl93.3 (10.1)90,O ( 72)

73.3 (0.06)83.3 ( 2,7)go.o < 1083.3

(10.0)53.3 ( O.5)36J ( 3.9)60,O ( O.2)40,O (O.03)26,7 ( O.1)200 {O.03)

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40,O (l2.8)50.0 ( 02)40.0 ( 1.l)30,O < 1.3)25,O C O.6)35.0 ( 2.9)65.0 (21.8)25.0 (O.02)20.0 (± )25.0

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SpeciescomposiLionandrichness of lucidophytlous forests onTsushimalslalld 7

Species group D

euercus acuta

Smilax china

9uercus serrata

Pi'unus juniasakura T)hea sinensis Dumasia truncata

E7beagnus pungens

Dioscorea euinqueloba

SO,rrax ,]'oponica Dioscorea tokoro

Athisia julthrissin Rohdoa .imponicaSpecies group E

71rachelospenmum asiaticum var, intermedium

demetha J'aponica

Ckestanopsis cuspidata s,ar. sieboldii

Ligustrlim jcrponicum Plersea thunbergii

Stauntonia hexaphydo

euercus saficina

Kadsura ]'aponica

Eurp,a J'aponica

Cinnamomum japonicum Symplocos lucidn (lphiopogon ohvvii

Dendropanax b'ijZius

Dnyopteris eu,throsora

Neolitsea aciculata

Ai'disia jmponica QFmbidium goet'ingii

Actinodaphne lanc(fofia Iiedera rhombea

Otherti

Persea J'aponica Itbx rotttnda

Parthenoctyst{s "it'uspiclata Flit'us erecta

Calanthe sieboldii

Rhus succedonea

Damnacanthus ma.ior

Clephalotaxus harrin.otonia

Litsea j'aponica

P.vrola J'aponica Calanthe artstutijlera

GootO'era schlechtendafiana

(]pltiopogon jaburan flyrrosia lingua Paeden'a scanclens var. mairei

Jlllbium religiosum

Lepisorus thunhergiantts

Ardisia pustila

Euonymus ,iqponicus filtus sarmentosa var, nipponica

Ftrtsia joponica DnyIJopteris vaFia vEtr, kikonensis

Rlex chinensts(Rest is omitted)

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8 Vegetation Science Vol. 22, No. 1,2oes

Table 6. Comparison of the number of species perIOO nn2 among the three forest types.

PR NA SE (1) {2)

Mean S,D, Mean S.D.Mean S.D. A]ltypesPR and NA PR and SE NA and SE

All species

Lucidophyllous element

Non iucidopliyllous element

33.S 2,6 26.1 5.0

32.9 22 23.3 4.1

O.9 O.9 2.8 2.6

21.4 3,4 p<e,eOl

17.9 2.6 p<O.oo13.5 1.5 p<O.oo1

p<O.Olp<o.elP<O.Olp<O.Olp<O.Olp<o.elp<o.osp<O.Oln,s.

Life form of lucidophyllous element

Tree Shrub

Ground herb

Groulld ferll

SapTophyte

Epiphyle Climber

15.6 1.85.6

l.322 1,32.3

O.7O,4 O,6O,8

O,9S.8 12

8.4 2,94.9

1,73,4 O.72.3 O.SO.t

02O,4

O.74.0 1,5

7.5 l.34,3

O.921 O,8O,3

O.5 oo oo3.2

O.9

p<O,OOIp<O,OIp<O,Olp<o.oelp<e.oolp<O,OOIp<o.oo!p<o.o]n.s,p<O.Oln.s.p<O.05n.s.p<O,Olp<O.OTp<O.Oln.s.p<O.Olp<O.Olp<O.Olp<O.Oln.s.n,s,p<O,05p<O.Oln.s,p<O.05n.s.

(1) / Significant level ofthe difi'erence among the three forest types by the Kruskal Wallis H test ; (2) /rhe t"'o focest t)'pes by the Mann-Whitney U-tcst with BonferToni correction.

Significant level of the difi'erence betwecn

g･ge, DISCUSSION

Differences in speeies composition

As shown in Fig. 2, the species composition differed

among the primeval, natural, and secondary forests, and

the arrangement pattern of these forest t,ypes along DCA

axis l corresponded to the degree of human disturbance of

the forests, which increases in the fo11owing order : prime-val, natural, and secondary. Furthermore, the stand score of

axis 1 ofthe primeval and natural fbrests was significantly

and highly correlated with the forest patch area (Fig. 3).These results suggest that the differences in speeies compo-

sition annong the three fbregt types greatly refiect the degree

of forest fragmentation as well as the degree of past humandisturbarice.

In all stands, SLA correlated highly with the stand score

of axis 1 and with both the number of species per plot tbrall species and that fbr lucidophyllous elements (Table 3).

However, no eflbcts of SLA on species composition and

richness were found in any ofthe three forest types. Thus,

we consider that SLA is not an important factor in the

difibrences in species composition und richness among the

fbrest types,

No previous study has shown the differences in species

composition between primeval and natural lucidophyllous

forests. However, the present study revealed that the

species composition of natural lucidophyllous forests on

Tsushima Is. clearly differ$ from that of the island]s prime-i,al lucidophyllous fbrests, This may be true in other

regions of Japan.

There were marked differences in species composition

between the primeval or natural tbrests and the secondary

forestg (Table 5). Similar findings have been reported by

Itow (1983>, who documented that the character species of'

natural Castanopsis forests were missing or rare in secon-

dary Castanqpsis forests in the Kyushu region, However,according to a study by Kamijo et al, (2001) on natura]

and secondary Castanpmis-Persea forests on Hachijo-

jima ls. in Japan, there was not much ditference in speciescomposition between the two fbrest types. Kamijo et al.

(2oo1) attributed the lack of dfl'erences between the twoforest types to the history of frequent natural disturbanceby typhoons. However, on Hacliijo-jima Is,, trees have

been cut for firewood und charcoal all over the island

(Ohba 1994). Thus, we consider that human disturbancehas changed the species composition of natural forests on

Hachijo-jima Is. and that, as a result, the species composi-

tion ofthese natural forests was similar to that of secondary

fbrests,

Aiba et al, (2oo1), who investigated old-growth and

secondary dnstanqrsis forests on Yakushima Is, in Japan,

reported that species composition wus similar between thetwo fbrest types, Their findings disagree with the presentresults. The reason fbr this disagreement may be that

Aiba et al. (2001) examinecl only trees whose tiunk diame-ter at breast height "ias at least 5cm, Ifall vascular plantshad been analyzed, there ",ould have been a significant

dilference in species composition between the t",o foiest

types.

Differences in species distribution

The clistributions ofspocies groups A and B were biased

toward the primeval and natural fbrests (Table 5). This

suggests that the populations of these species may be

isolated, According to Shaffer (1981), the isolated popu-lations are prone to localized extinction because of demo-

graphic, genetic, and environmental stochasticity, and their

probability of extinction depends on the size ofthe habitat.

Thus, the fbrest patch area is expected to have some eiibct

on the distribution of species groups A and B, To exam-

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Species composition and richness of lucidophyUous forests on Tsushima lsland 9

ine this point, we analyzed the relationship between thenumber of species per plot for species groups A and B and

the fbrest patch area fbr the natural stand group and the

primeval-･natural stand group by using simple correlation

analysis. The number of spec{es shewed significant posi-

tive correlations with the patch area in both the nttturalstand group and the primeval-natural stand group (Table4). The primeval-natural stand group showed a panicu-larly high correlation between the number of species and

the patch area (Fig, 4), Therefore, the fbrest patch areaappears to be related to the distribution of species groupsA and B in the primeval and natural forests.

Many species found in the primeval or natural forests

(spocies groups A and B) were missing or sparse in the

secondary fbrests (Table 5). One of the main reasons fbr

this is thought to be that these species have a low capacityfbr vegetative reproduction fo11owing clear cutting. This

is supported by the ftict that species with high ceppicing

abilities (e.g,, dastanopsis cuspiclata var, siehotdti, 2uereussafitrina, Symplocas incidtx, Persea thunbergii, CkemeiXin

.ioponica, Cinnamomum joponicum, Eurya J'aponica, and

Ligustrum J'oponicum) were included in species group E,Furthermore, species groups A and B included species that

prefer shade and rnesic soil, such as Micheha conu)ressa,

Aetinodaphne longij?)ha, Podocarpus macmp1ij]iins, Eu-chresta japonica, Desmodium laxum, Arachniodes spo-adhsora, Eldeagnus glabra, IVI?olitsea sericea, Arachniodlers

aristata, LemmapIrydeni micropiu,iZtim, Aucuba J'aponica,and Gatxineria nutans (Ishida et al. 1998;Hattori et al,2oo3), suggesting that their paucity in the secondary forests

can be attributed to rapid environmental changes caused

by clear cutting, such as increased exposure to sunlight and

drier soil.

Even when species with low reproductive abilities and

low tolerances for rapid environrnental changes are tempo-

rarily eliminated from one area of a sDcondaty forest byclear cutting, their seeds (including spores) can enter the

deforosted area from surrounding areas. Hence, with time,

the species composition of secondary forests should come

to resemble that of primeval and natural forests. How-

ever, the results of the present study showed that even 40

years after clear cutting, many component species ofprime-

val and natural forests were absent in the secondary forests,suggesting that either seeds cannot enter the secondary

fbrests or seeds cmmot germinate there. Regarding the

fifst hypothesis, we suspect that there are no plants c]oseenough to supply seedsto the secondary forests. As to the

second, the low arnount of light inside the forests may beinvoived. Itow (1983) reported that the canopy of socon-

dary lucidophyllous forests is completely closed, so littlelight reaches the fbrest floor and floor vegetation is thus

20

paEoo

15r

Lg8'6

108

cots

ts 5£2

o

.eoe

re

.

..

--..oee

.

.

........

O.1 1Area

(ha)10100

Fig. 4 Reiationship between the number of species per 1oo mU tbr species groups A and B and the fbrest patch area (log scale) for the primeval and natural forests.

poorly developed. Simi1ar findings were obtained for theseconda]y fbrests in the present study (Table 1). Thislack

of ]jght could explain the absence of some species in the

secondary fbrests.

In the socondary forests, shade-intolerant deciduous tree

spocies were found (euercus serrata, Rrunus jamasakura,Styrax jmponieus, and Athizia juhopissin), as ",ere decidu-ous climber species (DiascoJea quinqueloba, Smikix china,and Dioscorea tokoro), although they were missing or

sparse in theother two fbrest types (Table 5), Ofthese, thedeciduous tree specie$ occurred mostly in the tree layer(Table2). Similar findings have been obtained jn other

parts ofthe Kyushu region (Kohyamtt & Aiba, 1997 ; ltow1981a; Ishida & Hattori 1998). The reason why the

deciduous tree species in species group D can survive in thetree layer for a long time after clear cutting would be thatthey grow at a rate faster than or at least comparable to

evergreen broad'･leaved trees after clear cutting,

Relationship between dominant type and humandisturbance

Dtstylinm raeemosum is one of the dominant species inJapan's lucidophyllous fbrests. D. racemosum-dominantforests occur widely in western Japan, and are distributedin both lowland and mountainous areas of the Kyushu

and southern Shikoku regions (Nozaki et al. 1996), OnTsushima ls., we found that D. racemosum dominated,together with Castanqpsts cuspidata var. siebotdti, in the

primeval forests, but not or rarely in the natural and

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10 Vlegetation Science Vol. 22,

secondary fbrests (Table2). The reduction ef D.racemasum in the natural and secondary forests would beattributable to its extremely low coppicing ability (Aiba etal. 2001), whjch prevents it from remaining in the cornmu-

nity after tree cutting. Hence, based on the results of the

present and past studies (Itow et al. 1976, Itew I98]b), the

dominant type of fbrest communities (excluding coastal

fbrests) in Tsushima's Ckestanqpxsis zone would be alteredin the fo11o",ing sequence when subjocted to human distur-

bance: C, cuspidata var. sieboidii-D. racemosum forest;

C cuspidata var. sieboldli-2uercus satlitrina forest ; and 9serrata"PlalycaoTa strobhacea forest,

Differences in species richness

The total species richness and the lucidophyllous-ele-

ment species richness differed significantly among tbg three

forest types, and was highest fbr the primeval forests,foIIowed by the natural and then the socondary tbrests

(Table 6). Because the number of species per plot fbr al1species and that for lucidophyllous elements did not haveclear relationships with forest fragmentation (Table 4). thedilTerences in species richness among the forest types

appear attributablc mainly to humun disturbance. How-ever, the number of species per plot for species groups A

and B was highly and positively corretated with the forest

patch area (Fig,4). Therefore, fbrest fragmentationwould be partly responsible fbr the differences in speciesrichness among the forest types.

There was a relatively large difi'erence in the number of

species per plot for lucidopltyllous-element trees betweenthe primeval forests and the natural or secondary ['orests

(Table6), The specjes richness of lucjdophyllous-e]e-ment trees may be more susceptible to human disturbancethan the species richness of other life forms.

Hattori & Asami (1998) have pointed out that the

number of species per unit area for lucidophyllous ele-

ments is a usefu1 index of the naturalness of a lucidophy]-

lous forest. The present results would support their asser-

tion.

The -umber of al1 vascular plant species per 100 mZ fbr

secondary summergreen ({2uereus serrata-C}uerctts varL

iahids) lbrests in the Castanopsis zone of the Seto lnland

Sea region in Japan tended to be higher than that fbr

natural lucidophyllous forests with similar climatic, geo-]ogic, and topographic conditions (Ishida & Hattori 1997),

This is because human disturbances, such as c[ear cutting

and understory cutting, increased the llumber of non-

lucidophyllous elements more than it decreased the num-

ber of lucidophyllous elements. However, in the presentstudy, the number of non-lucidophyllous elements in-

creased only slightly as human disturbance increased.

No. 1, 200S

This is because non-lucidephy]lous elements would

receive insuMcient light in the seeondary lucidophyllousforests and therefore could not grow there. Thus, speciesrichness would increase when the intensity and frequency

of human disturbance on the secondary lucidophyllousforests increase, and as a result the canopy becomesdominated by deciduous trees rather than evergreen broad-

leaved trees. [n fact, the mean number ofspecies per leOm2 for al1 yascular plant spocies was over 44 for secendarysurnmergreen (Cuereus senrata-Platycat:ya strobildcea>

forests in the CZistanqpsty zone on Tsushima Is. (Takeda et

al., unpublished data), which is greateT than that for the

primeval fbrests.

ig', CONCLUSION

The present stucly revealed that there are clear differencesin species composition and riehness among the primeval,natural, and secondary lucidophyllous forests on Tsushimalsland, and these diFferences are reluted to the degree of

past human disturbance and of forest fragmentation.Thus, we conclude that even in the warm and wet reglons

where secondary lucidophyllous fbrests can exist, lauman

disturban¢ e and fbrest fragmentation have significimt

eflects on the species composition and richness of lucido-

phyl]ous forests.

kg ACKNOWLEDGEMENTS

We thank H, Akamatsu, K, Asami, S. Yamamoto, M,Yamato and Y, Yagura, and the members of the Labora-

tory of Vegetation Science, Kobe University fbr their helpin field sunreys. We also thank Nagasaki District Forestry

Otiice fbr permission for fieldwork in Mt.Tatera. The

comments of two anonymous reviewers and an editor

significantlyimprox,edthequalityofthernanusciipt. This

study was partly supported by a Grant in-Aid forScientific Research from the Ministry ofEducation, Science

and Culture, Japan (No. 16510I78),

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environment ofnature park in Hachijo-jimu Island. Vol,3

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Natienal Parks Associaiion of Japan, Tokyo {in Japanese),Sasaki,Y,(ed), 1973. Phytosociology. Lectures onecolo.uy4.

Kyoritsu S'huppan Publishers, Tokyo (in Japanese).Shaffer, M.L. 198I. Minimum population sizes for species

conservatlon. BioScience, 31: 131'134.Suzuki, K. 1982. Disiribution or evergreen and summergreen

broadTteaved forest in Japan. Bulletin, Institute of Environ-

mentul Science and Technology, Yokoharna National Univer-

sit)', 8: 151-I63,Suzuki, T. & Suzuki, K. 1973. Vegetation ofwarm'temperate

zone in Japan. In: Phytosociology. Lectures on ecelogy 4

(cd, Sasaki, Y,), 18-24, Kyoritsu Shuppan Publishers, Tokyo

(in japanese).Yaunamoto, S, ]992, Gap characteristics und gap regeneration

in primary evergreen broad-leaved forests of western Japan,



The 　Soolety 　of 　▽ egetatlon 　Solenoe

12 Vegetation　Science　Vo122 ，　No 　 L　2005

　Botanical　Maguzine，　Tokyo ，105 ： 29−45．Yamanaka ，　 T．1969　 Ph｝tosocio ［ogical 　 studies 　 of　 secondary

　fc〕rests　ln　southern 　Shlkoku，　Japan．　Research　ReporLs　of　the

　K6chi　University（Natural　 Sclence），18 ： 1−14 （in　Japanese

　with 　English　summary ）Yalτ1anaka ，　T 　1979　 Forest　vegetatK 〕n　ofJapan ．　了sukljl　 sho ・

　kan 　Publlshers．　Tokyo 〔hl　JaPanese ）

繋要約

　対馬における照葉樹の原生林，自然林，二次林の種

組成および種多様性の比較．石田弘明・服部　保（兵庫

県立大学自然・環境科学研究所）・武田義明（神戸大学発

達科学部）

　入工林以外の日本の照葉樹林は自然性の違いによっ

て大きく 3つのタイプに区分される，すなわち，（1）

ほとんど人の手が加わっておらず，原生状態を維持し

ている樹林（照葉原生林），（2）薪炭林のような定期的

伐採や下刈りは行われなかったが，社寺林のように，

不定期で部分的な人為攪乱や著しい断片化によって自

然性が低下し，原生状態とはいえない樹林（照葉自然

林），（3）かって薪炭林として利用されていた萌芽林

（照葉二次林）の 3タイプである．本研究では，これら

の森林タイプの種組成および種多様性の相違と人為攪

乱および断片化の関係を明らかにすることを目的とし

て，対馬の照葉原生林，照葉自然林，照葉二次林に 100

M2 の調査区を合計 76 区設置し，全ての維管束植物を対

象とした植生調査を行った．全層の種組成の類似性に

基づいて DCA によるスタンドの序列を行った結果，

（1）照葉原生林，照葉自然林，照葉二次林の種組成は

互いに明らかに異なっていること，（2）照葉自然林は

照葉原生林と照葉二次林の中間的な種組成を有してい

ることが明らかとなった．また，照葉原生林および照

葉自然林の DCA 第 1軸スコアと樹林面積の間には高い

有意な相関が認められた．これらのことから，森林タ

イプ問の種組成の相違は人為攪乱と断片化の程度を大

きく反映していると考えられた，照葉原生林ではイス

ノキとスダジイが優占していたが、照葉自然林や照葉

二次林ではイスノキはほとんど優占していなかった．

この現象は，イスノキの萌芽力の低さと自然林および

二次林で行われた樹木の伐採に起因していると考えら

れた．種多様1生（species　richness ）の尺度として調査区あ

たりの出現種数を算出し，この値を森林タイプ問で比

較した．全出現種と照葉樹林要素の出現種数は森林タ

イプによって有意に異なっており，照葉原生林，照葉

自然林、照葉二次林の順に減少する傾向にあった．全

出現種および照葉樹林要素の出現種数と断片化の問に

は明瞭な対応関係は認められなかった．しかし．照葉

原生林や照葉自然林に偏在する種の出現種数は樹林面

積と高い正の相関関係にあったことから，森林タイプ

間の種多様性の相違には人為攪乱だけでなく断片化の

影響も関係していると考えられた．

N 工工一Eleotronlo 　Llbrary 　




Speciescornposlttonandrichnessoflucidophyllousforests onTsushimaIsland 13

Appe]dix 1. score of

Characteristics of

DCA axis 1.the

standsinvestigated.Stands arearrangedaccording tolncreasmgrhestand

StandNo,Forest

typeALTMTCMAPDFNCSLDSLA(1)(2)(3)(4)(s)

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14 Yegetation Science Vol. 22, No. I, 2005

5455S6575SS9606{626364656667686970717273747576SESESESESESESESESESESESESESESESESESESESESESESE200l851801801902oo185195]80190185190185190l85190190l9S190[85195l95L853,43.53,S3,53,53.43.53.53.53.5353,53,53,53.53.53,53,53.53,53,S3.53,521092t09210921092I092109210921092!0921092109210921092109210921092I092109210921092109210921093.163.113,113.103,153,L63.ll3.173.123.183,L23.173,L73.183.153.133,IS3,16].193.193.173.153,16201525202022252020]52015202220l510510I95

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l)R: Primeval forst ; NA / Natural forest ; SE : Secondary tbrest.

ALT / Altitude <m) ; MTCM / Mean temperature of the coldest month ('C) ; AP : AnnuaL precipitation (rnm) ;

DFNC / Distance from the nearest coasttine (km) ; SLD : Slope degree (') ; SLA : Slope aspect.

(1): Stand seore ofDCA axis 1 ; (2) / Stand score ofDCA axis 2; (3) i Nurnber ofull species ; (4) : Number of

species for lucidophyllous eLements ; {5) : Number of species for non-lucidoph},11ous elemeiits,

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