Bull. Kitakyushu Mas. Nat. Hist., 16: 113-135. March 28, 1997

Vegetation Structure of a Secondary Grassland at a Line Corridor in Fukuchi Mountain System, Northern Kyushu

Tohru Manabe1, Kazuaki Naito2 and Nobukazu Nakagoshi3 'Kitakyushu Museum and Institute of Natural History, Nishihonmachi 3, Yahatahigashiku, Kitakyushu 805, Japan 2Department ofAnimal Production, Chugoku National Agricultural Experiment Station, Kawai-Cho, Oda, Shimane 694, Japan 3Faculty of Integrated Arts and Sciences, Hiroshima University, Higashi-Hiroshima 739, Japan (Received December 5, 1996)

Abstract Vegetation structure was studied at a line corridor which has been formed and maintained as fire prevention belt by periodic mowing at the ridge of a mountain system, northern Kyushu. Grassland herbs were dominant, although forest herbs which usually grow on forest floor occurred at the corridor. The grassland at the corridor was identified phytosociologically as Bupleuro-Miscanthetum sinensis belonging to Arundinello-Miscanthion sinensis alliance. The corridor was classified as trough corridor, because its vegetation height was lower than that of the adjacent matrix. The corridor was composed of various kind of tessera which were brought by many types of landscape element at the adjacent matrix, microtopography and surface geology at the corridor. Those tessera were distributed heterogeniously over the corridor. Distribu tion patterns of the plant species within the corridor were also affected by those of the tessera. Thus, this narrow line corridor had the function of habitat for many grassland herbs which are becoming rare. And variety of the tessera brought about variety of landscape structure generated high composition diversity of the plant species at the corridor.

Introduction Secondary grasslands, which were one of the main landscape element types (Forman and Godron, 1986), had been maintained by mowing and pasturage under the traditional sustainable land use systems inJapan (Naito and Nakagoshi, 1994). Popularization of petroleum fuels and chemical fertilizer in the 1960's, which is called the fuel revolution, led to abandonment of the traditional land use, and brought about rapid and drastic changes in distribution patterns of landscape elements (e.g. Kamada and Nakagoshi, 1996). Secondary grasslands are one of the typical landscape elements which have rapidly decreased in area since the fuel revolution (Someya et aL, 1989; Naito and Nakagoshi, 1994; Shoji et aL, 1995). The collapse of traditional land use systems also made some grassland species become rare, since 114 Tohru Manabe, Kazuaki Naito and Nobukazu Nakagoshi secondary grasslands were favorable habitats for them (The Investigation Commit tee for Important Plant Species and Communities for Conservation in Japan, 1989). In northern Kyushu, secondary grasslands are distributed on limestone plateaus showing patch shape in suburb landscape. These grasslands function as habitat for some grassland species (The Kitakyushu Municipal Education Board, 1973; Naito et aL, 1995). Secondary grasslands also distribute on the ridge of the moun tains located at the suburb. These grasslands, which have been formed and maintained as fire prevention belt by periodic mowing, take the form of line corridors. The line corridors with secondary grasslands are surrounded with various kinds of matrix such as coppices and conifer plantations, and are on the mountain ridges with various microtopography. These line corridors might contain many kind of tessera (Forman and Godron, 1986) which were brought by the various matrix, microto pography and so on. Thus, these secondary grasslands established at the line corridors might be also complex of microsites with various microenvironmental conditions for plant species. But, flora of these line corridors has not been reported because of its strip figure with nallow width. Further, there are few evaluation of these line corridors as habitat for grassland species. Corridors, which are defined as "narrow strips of land that differ from the matrix on either side" (Forman and Godron, 1986), perform well known functions in landscape such as habitat, conduit and filter (or barrier) as well as functions known empirically or conceptually such as source and sink (Forman and Godron, 1986; Saunders and Hobbs, 1991; Smith and Hellmund, 1993; Forman, 1995). Those five major functions of corridors in landscape are affected not only by biological attributes such as kind, number and /or biomass of living things growing within the corridors and at the matrix adjacent the corridors, but also by pattern and process of landscape structure such as width, length, connectivity and curvinearity of corridors, relationships to the matrix, origin of corridors and so on (Forman and Godron, 1986; Crome and Richards, 1988; Saunders and Hobbs, 1991; Smith and Hellmund, 1993; Forman 1995). Habitat and conduit functions of corridors are also important on the view pointof biological conservation, because these functions might havea roll to promote interchange among individuals in isolated sub-populations within fragmented landscape (Saunders and Hobbs, 1991; Forman, 1995). Although habitat and conduit functions of corridors have been analyzed for birds, mammals and insects growing within corridors (e.g. Adams, 1984;Opdam et aL, 1984, 1985; Kealv and Majer, 1991; Lynch and Saunders, 1991; Andreassen etaL, 1996), only a few works have done those for plants (McDowell et aL, 1991; Johansson et aL, 1996; Riffell and Gutzwiller, 1996). Structural and functional features of corridors have discussed mainly on wooded strip corridors at flat plain suchas hedgerows and on stream and river corridors such as riparian forests, but few on corridors with complex three-dimensional form such as those located at the ridge Secondary Grassland at a Line Corridor 115 of mountains in northern Kyushu. In this paper, we describe the vegetation structure of a secondary grassland at a line corridor which might contain many kinds of tessera and the matrix surrounding a corridor. And we discuss the feature of habitat and conduit function of the line corridor, especially those for grassland plant species. Nomenclatures are followed by Ohwi and Kitagtawa (1992) for Sperma- tophytes and by Iwatsuki (1991) for Pteridophytes.

Study Site The line corridor locates at the ridge of Fukuchi mountain system (33°43'N- 33°40'N, 130°48'E-130°49'E) ranging north to southeast across Kitakyushu City, Akaike-cho, Houjo-cho and Kawara-cho in Fukuoka Prefecture. This corridor has been formed and maintained as fire prevention belt by periodic mowing. Then, the corridor have secondary grassland owing to periodic human impacts, and it has been also using as the path along the ridges for mountain climbers. The corridor has about 10 m width, while it has more than 6,000 m length. That is, this corridor is classified as a line corridor (Forman and Godron, 1986). Many peaks within the corridor such as Mt. Fukuchi (900 m), Mt. Akamutanotsuji (791 m) and Mt. Yakitate (759 m) generate complex three-dimensional structure having ca. 250 m disparity in a.s.l. (Fig. 1). Various types of vegetation such as deciduous broad- leaved forests and evergreen broad-leaved forests occur over the mountains, although Japanese seder {Cryptomeria japonica (Linn, fil.) D. Don) plantations are most dominant. The surface geology of the corridor is classified as three types: northern part and southern part of the corridor are dominated by metagabbro and quartz- porphyry, respectively, the middle site formed by shale, sandstone or conglomerate is a little (Fukuoka Prefecture, 1970).

Methods

An actual vegetation map of the rectangle area, which contains line corridor, with 1 km width and 6 km length was drawn at a scale of 1: 25,000. The boundary of each landscape elements, which are recognized by homogeneous physiognomy, was determined by using air photographs taken in 1994 at a scale of 1 :20,000 by the Geographical Survey Institute, the Ministry of Construction. Phytosociological survey of vegetation (Braun-Blanquet, 1964) was conducted at thirty two of 2 mX2 m quadrats established at a central part of the corridor in the middle of May, 1996 (Fig. 1). Vegetation and microtopographical structure of the corridor and adjacent matrix were analyzed at 10sites in the middle of September, 1996. The inclination of each 1m was measured on a 10m line which was parallel to the corridor at each site. Another line was established at each site so as to cross Mt. Fukuchi 1,0001— Mt. Akamutanotsuji (901 m) (791 m) Mt. Yakitate (759 m)13^3*4 *1 21 90 B1 A H >

S 500 31 29 27

> O 3

oL 500 m

Fig. 1. Maps showing the study area. Dots; quadrats investigated composition of vegetation, asterisks; sites investigated vertical structure of vegetation.

Secondary Grassland at a Line Corridor 117 the corridor and both side of the matrix, that is, this line and the 10 m line were at right angles. Vegetation structure such as names and diameter at breast height (d.b.h.) of dominant species and vegetation height were recorded within both 1 m sides of the line. Microtopographic locations of the quadrats and the sites were also recorded according to Tamura (1980).

Results

Landscape structure The curvilinearity of the corridor on the horizontal projection was low, although the corridor branched complexity at the southern end of it (Fig. 2). The main part of the corridor ranging north to southeast connected five branch corridors from west side and one branch corridor from east, indicating that the corriodor formed relatively simple network. The twelve types of landscape element were identified such as evergreen broad- leaved forests dominated by Quercus acuta Thunb. and Neolitsea sericea (Blume) Koidz., evergreen broad-leaved and deciduous broad-leaved mixed forests (hereafter mixed forest) composed of above evergreen trees and Q. serrata Thunb. and Carpinus tschonoskii Maxim., pine forests dominated by Pinus densifiora Sieb. et Zucc or P. thunbergii Parlet., plantations of Cryptomeria japonica or Chamaecyparis obtusa (Sieb. et Zucc.) Sieb. et Zucc, apud Endl., dwarf bamboo communities dominated by Sasa senanensis (Franch. et Savat.) Rehder, grasslands dominated by Miscanthus sinensis Anderss., and so on. Most dominant landscape element was plantations, followed by mixed forests and evergreen broad-leaved forests. Grasslands were also classified two types due to the intensity of mowing: grass communities with low mowing impact, which mainly located at the matrix, where Miscanthus sinensis grew vigorously and saplings of deciduous broad-leaved trees (ca. 2-3 m in height) such as Quercus serrata and Carpinus tschonoskii grew sparsely, and those with strong mowing impact located within the corridor. The seven types of landscape element abutted on the corridor: evergreen broad- leaved forests, mixed forests, pine forests, plantations, grass communities with low mowing impact, scrub community and denuded land. Landscape element types other than scrub community and denuded land have taller vegetation height than that within the corridor (Table 1 and Fig. 3). The corridor was constituted twelve kinds of tessera by the combination of landscape elements at both sides: tessera abutted on mixed forests and pine forests, that abutted on mixed forests and plantations, that abutted on pine forests and plantations, and so on. No matter which cut the connectivity was within the corridor except for the rocks having tree species such as Ilex crenata Thunb., Eurya japonica Thunb., Cornus kousa Buerger ex Hance, and so on at a few sites (Fig. 3). 118 Tohru Manabe, Kazuaki Naito and Nobukazu Nakagoshi

Table 1. Vegetation and landscape structure of the line corridor and the adjacent matrix.

Plot Site Dominant Height of Width of Mean Type of Micro- Surface species" vegetation corridor slope topo- geology4' (m) (m) degree (°) graphy31

corridor Ms, Pa 2.1 6.6 4.5 MM HH MG matrix (cast side' Pj (29.8) 6.5

matrix (west side) Ck(13.8) 7.0 corridor Ms, Pa 1.0 10.7 18.5 MJ SS MG

matrix (cast side] CK(21.1) 6.5

matrix (west side ) Co (17.8) 6.1

corridor Zi, Ms 0.2 11.3 6.3 JJ CS MG

matrix (cast side] Co (15.3) 6.1

matrix (west side ) Co (16.1) 4.7

corridor Ms, Ic 1.4 12.5 3.3 MJ CS MG

matrix (cast side] Ck(ll.5) 4.2

matrix (west side ) Co (13.2) 5.9

corridor Ms, Ps 1.2 8.8 2.0 JG CF MG

matrix (cast side] Ms, Ck (8.8) 2.1

matrix (west side ) Co (16.7) 5.5

corridor Ms, Pa 1.2 9.0 11.5 JG CS MG

matrix (cast side] Ms, Ck (11.3) 2.4

matrix (west side) Co (15.9) 4.9

corridor Ms, Ss 0.4 9.5 27.8 PJ CS MG matrix (cast side] Pd (21.8) 5.5

matrix (west side) Co (12.6) 4.6

corridor Ms, Pa 1.3 9.5 3.5 EP HH SS

matrix (cast side] Qj(16.4) 5.1 matrix (west side ) Pt (21.2) 7.4

corridor Ms, Sa 1.9 7.5 3.3 MJ HH QP

matrix (cast side] Le(15.4) 7.9

matrix (west side ) Co (17.5) 6.4 corridor Mc.Ct 0.8 6.8 6.8 JJ CS QP matrix (cast side] Co (9.3) 2.7 matrix (west side ) Co (7.8) 2.5 1) Ms; Miscanthus sinsnsis, Pa; Pteridium aquilinum var. latiusculum, Pj; Pmusjamaszkura, Ck; Cornus kousa, Co; Chamaeciparis obtusa, Zj; Zoysia japonica, Ic; Ilexcrenate, Ps; Patrinia scabiosaefolia, Ss; Salix sieboldiana, Pd; Pinus densijlora, Pt; Pinus thunbergii, Cj; Cinnamomum japonicum, Sa; Solidago altissima, Lc; Lindera erythrocarpa, Ct; Cymbopogon tortilis. D.b.h. of dominant tree arc in parenthesis. 2) Two letters indicate vegetation types of matrix at the both side of quadrat: M; evergreen and decidous broad-leaved mixed forest, J; Japanese cedar and cyperes plantation, P; pine forest, G; grassland, E; evergreen broad-leaved forest 3) CS; crest slop, CF; crest flat, SS; sidcslope, HH; head hollow 4) MG; metagabbro, SS; shale sand stone, QP; quartz-porphyry Site D

-3»K- -SH«S- -X- -5t<£- -5 O

Site F

O

5*e Fig. 3. Vertical structure of vegetation at the corridor and the adjacent matrix at typical sites in Fukuchi mountain system. Abbreviations: CD; corridor interior, ME; matrix (cast side), MW; matrix (west side), Ap; Acer palmatum, Cj; Cinnamomum japonicum, Co; Chamaecyparis obtusa, Ck; Cornus kousa, Cs; Celtis sinensis var.japonica, Dc; Deutzia crenata, Ej; Eurya japonica, Ic; Ilex crenata; Ip; Ilex peduncculosa, Lo; Lyonia ovalifolia var. elliptica, Ms; Miscanthus sinensis, Ns; Neolitsea sericea, Pa; Pteridium aquilinum var. latiusculum, Pd; Pinus densiftora, Pj; Prunus jamasakura, Qa; Quercus acuta, Qs; Quercus serrata, Rj; Rhus javanica, Ss; 5a/ix sieboldii. Each bar indicates 2 m Table 2. Species composition of 2 mX2 m quadrats at the line corridor at the Fukuchi mountain system. Values are dominance and sociability in the ZM school of vegetation science.

Quadrat No. | 10 17 2 3 4 5 6 8 11 9 14 15 12 13 18 16 7 27 28 29 30 31 20 25 22 24 23 21 19 26 32 Above tea level (m) 650 740 650 670 700 730 760 780 780 750 770 690 680 730 710 660 660 780 540 540 540 530 530 610 530 620 560 600 610 660 530 550 Typa ormatrix1' MM JG MJ MM MJ MJ MM JJ MP JG JG MJ MP MJ MJ PC PJ JG JG MJ MJ MJ MJ MJ MM JJ JJ JJ JJ MJ JO MJ Microlopography2' HH CF CF SS SS SS SS SS CS CS CS CS CS CS CS CF CF CS CF CF CF CF CF CS CF CS SS SS CF CF CF CF Subsurface geology3' MG MG SS MG MG MG MG MC MG MG MG MG MG MG MC SS SS MC QP QP QP QP QP QP QP QP QP QP QP SS QP QP Htsta aUo-Karpntl* : II Aitmm atanida Hrlcmcpril arunlalii Dbponmjtssilt Bltckam mfmnan ttaaaudiajepauau Atttia aubata TritfrSs afloat Euphorbia sitboUiana Thtmtdajaponica Asterettratoidt$var. ovatas Calm Irachjsptmmm * Ahibiatrifoliota Rubaspartifoliut Knbiupalmatai : 1-2 Potima tcenins var. maitri • ! + lUx aenota •...'... Zoyiia japonica Ptritbtatba thine var. ciritSi Lanalacapitcta '"is:" ••• Uibmltta ananJjia Safin atholaHana Utptitta pitasa InJitofira puudo-tindtria Character speciesof Bupleuro-Miscanthetum sinensis association Sonoma ptkhdla f. nbinltpo + • • + tmpnatacjtinirica var. koenipi Characterspeciesof Arundinello-Mucanthionsinensisalliance 2-3 1-3 1-2 2-3 2-2 3-3 1-3 1-2 2-2 2-3 2-3 1-3 2-3 + 3-3 2-3 2-3 1-2 23 + + + 2 AnndaiUa hirt* > •3 + 1-2 1-2 + 1-2 + 1-2 Attn train + + • 1-2 1-2 1-2 1-2 1-2 II 1-2 + •2 1-2 + + + 1-2 Artemisiajaponic* o SoBaap roga-oitrra var. asiatica + + + + + o CattxJbribunao Ptmimm aasittnwn var. lahumhm 12 1-2 12 Ljsmochiaclithnuitt AinophoratriphjUe EupalatUm tkintnnvar. senpUdfetiain Usptdtta cmcala Lctpcdeiacyrtobotrya Diantkns nptrbusvar. longkaljcbus Characterspeciesof Miscantheluni sinensis class Miscanlhai rinmiit 22 23 2-3 2-2 2-2 22 3-3 2-3 3-3 3-3 2-3 1-2 3-3 3-3 3-3 2-3 3-3 3-3 33 1-3 3-3 1-3 23 33 2-3 1-2 3-3 + 33 3-3 1-2 1-2 1-3 1-2 1-3 + + + 1-2 Ixtrit dmtata 1-2 PotmtUla fnyniana Gtntisne tcabta var. bncrgeri Patygata japonica Cirsium japonicum Haloragis micranlha Accompanying species Viola grypoatas Ptcctraniha ujbxut + + + + + + + + Artemisia primps Angelica pubescent Smitax china Lonicera japonica Pueraria Malt Lmata maltS/bra PotenlUla fragamOes var. major Hypericum trectm Gnaphalhm japonicum Dktscotea japonica Hjdroaljtrsibtiorpuides Rosa wichuraiana SjnuntsextdtiB Piciris hicracioides var. glabttscent Viola dolacea Cousins orbiculata Ctlastntt orblcttlaha Brach/podiwn sjkaaam Peiinlaristettpineta Cirsium lincare var. discolor Ljsmackia japonica Rubas hirsutas Agrimonia japonica Etaeagrais umbcllata Chinopodhm micraatham Pulsatilla cemua Hjperieimlaxtan Lactuca indite var. laciniatt O Rhjnchosia acuminatifolia Chtorantbts japoninu IJIitm ardatta Rubus minusatlttt Poljgtmtumfekahm Valeriana femes Stemtenia hexaphjlta Ampdopsis brttipedunculate Deutzia crenela Chamaecyparis obtusa SaUx abopposita o Dwnaaa truncate Solidago attittima Botejchtam tctaatum Thesism chinense Opllsmtnus undalatifoHus var.japonicus Viola mandthurica Compositae sp. Smithie eitiata Chrysanthemum indiaan Erigeron amuut Carex brccieutmis Senecio tntegrijolixm var. spathttatus Epipactis tmmbergil 1) Two letters indicate vegetation types ofmatrix at the both side ofquadrat M; evergreen anddecidous broad-leaved mixed forest, J; Japanese cedar and cypres plantation, P; pineforest, G; grassland 2) CS: crest plope, CF: crest flat, SS: sidcslope, HH: head hollow 3) MG: mctagabbro, SS: shale sand stone, QP; quartz-porphyry 122 Tohru Manabe, Kazuaki Naito and Nobukazu Nakagoshi

Vegetation structure Miscanthus sinensis occurred at all quadrats and had the highest coverage at most of quadrats (Table 2). Character species of Miscanthetea sinensis class such as Ixeris dentata (Thunb.) Nakai, Potentilla freyniana Bornm. as well as M. sinensis and those of Arundinello-Miscanthion sinensis alliance such as Arundinella hirta (Thunb.) C. Tanaka, Aster scaber Thunb. and Artemisia japonica Thunb. occurred frequently. Grassland species such as P. fragarioides Linn. var. major Maxim., Synurus excelsus (Makino) Kitam. and Cirsium lineare (Thunb.) Schultz Bip. var. discolor Nakai also occurred at many quadrats, although they had low coverage. Further, Pulsatilla cernua (Thunb.) Spreng. which was designated as a threatened plant (The Inves tigation Committee for Important Plant Species and Communities for Con servation in Japan, 1989) was found at some quadrats, and sub-populations with more than one hundred individuals existed some sites other than quadrats. The quadrats with high coverage of Zoysia japonica Steud., Pleioblasius chino (Franch. et Savat.) Makino var. viridis (Makino) S. Suzuki and A. hirta had relatively low coverage of M. sinensis. These quadrats often located on crest slopes with steep inclination and side slopes with steep inclination. Some quadrats had forest herbs mainly grown on the forest floor and at the forest edge such as Hosta albo-marginata (Hook.) Ohwi, Asarum asaroides (Morr. et Decne.) Makino, Heloniopsis orientalis (Thunb.) C. Tanaka, Ranunculus japonica Thunb. and Tricyrtis afjinis Makino. These quadrats usually located near the mixed forests with large area which were distributed northern part of vegetation map drawn. Many species flowered and/or fruited at twostudy periods, suggesting that these species might maintain its population within the corridor.

Discussion Relationships between landscape structure and vegetation structure Origins, patterns of formation, structure of corridors as well as relationships to the matrix are important factors for the structure and dynamics of populations and communities within the corridors (Soul£ and Gilpin, 1991; Nicholls and Mar- gules, 1991). Corridors are classified into five types by their originssuch as disturb ance corridors, remnant corridors, environmental corridors, planted or introduced corridors and regenerated corridors (Forman and Godron, 1986). The corridor studied was classified as disturbance corridor, because it was formed and maintained by periodic mowing which was one of typical human disturbances. This periodic human impacts are one of important factors for maintaining secondary grasslands and species diversity of the sites (Itow, 1981). The corridor was typical trough corridor (Forman, 1995), because it had vegetation lower in height than the adjacent matrix. In corridors having trought Secondary Grassland at a Line Corridor 123 form, animals passing throughout the corridor may have much possibility to be caught by their predators. Then, animals such as frugivore do not often use the corridors for their habitat, suggesting that the corridor studied has weak effects on conduit and source functions for plants producing endozoochores and epizoochores seeds. But, the corridor plays important role of conduit for plants producing anemochores seeds. At trough corridors with narrow width, physical and biological features within corridors might be affected by those features at the surrounding matrix through strong edge effects (Murcia, 1995). Seedlings and saplings of trees growing at the matrix and forest herbs occurred within the corridor. Further, the sites having those species often located near the mixed forests with large area. The vegetation composition within the corridor, therefore, is also affected by the vegetation structure of the matrix through the edge effects. Thus, those sites might have functions of habitat and sink for forest herbs. Off-road motorcycles had often rushed at the corridor, although intrusion of the motorcycles within the corridor was prohibited recently. The effect of motorcycles on the surface ground is stronger than that of hikers and horses, and the effect of motorcycles are especially strong at the slopes (Weaver and Dale, 1978). Ruts with soil erosion and no vegetation were observed at the steep slopes within the corridor. Zqysia japonica, Arundinella hirta and Pleioblastus chino var. viridis often grew at those slopes with high coverage, indicating that vegetation of the slopes are retrogressive due to the effect of motorcycles. The corridor studied was composed of various kind of tessera which are brought by landscape elements of the matrix abutted on the corridor, microtopography and surface geology at the corridor. And those tessera were distributed heterogeniously across the corridor. Distribution patterns of the plant species within the corridor were also affected by the distribution patterns of those tessera. Thus, variety of the tessera brought about variety of landscape structure generates high compositional diversity of the plant species at the corridor.

Feature of the secondary grassland Secondary grassland established at the corridor had many grassland species. This grassland belongs to Arundinello-Miscanthion sinensis phytosociologically, since not only many character species of Miscanthetea sinensis class but also those of Arundinello-Miscanthion sinensis alliance often grew at the grassland. Typical associations belonging to Arundinello-Miscanthion sinensis in Kyushu are following four associations such as Themedo-Miscanthetum sinensis, Arundinario pygmaeae- Miscanthetum sinensis, Plectrantho-Miscanthetum sinensis and Bupleuro-Miscanth- etum sinensis (Itow, 1981). In the grassland at the corridor, character species of Arundinario pygmaeae-Miscanthetum sinensis such as Sanguisorba officinalis Linn, and Galium verum Linn. var. asiaticum Nakai forma nikkoense (Nakai) Ohwi and those of 124 Tohru Manabe, Kazuaki Naito and Nobukazu Nakagoshi

Plectrantho-Miscanthetum sinensis such as Plectranthus japonicus (Brum, fil.) Koidz. and Cirsium spicatum (Maxim.) Matsum. did not occur or occurred at low frequency, although Pleioblastus chino var. viridis which does not occur at Themedo-Miscanthetum sinensis grew at the grassland. On the other hand, character species of Bupleuro- Miscanthetum sinensis such as Saussurea pulckella Fisch. ex DC. forma subintegra (Regel) Kitag. and Imperata cylindrica (Linn.) Beauv. var. koenigii (Retz.) Durand et Schinz were occurred at high frequency. Many individuals of Bupleurum scor- zoneraefolium Willd. var. stenophyllum Nakai were observed in September, although those were not identified in May (c.f. appendix). Thus, the grassland at the corridor was identified as Bupleuro-Miscanthetum sinensis. Frequency of Bupleurum scorzoneraefolium var. stenophyllum and Bothriochloa parviflora (R. Br.) Ohwi, which are character species of Bupleuro-Miscanthetum sinensis at the grassland at the corridor was lower than that at Hiraodai plateau in Fukuoka prefecture (The Kitakyushu Municipal Education Board, 1973) and Akiyoshidai plateau in Yamaguchi Prefecture (Shiomi et aL, 1992). Further, Buglossoides zollingeri (DC) Johnston, which grows commonly at both Hiraodai and Akiyoshidai plateau, did not occur at the grassland. These trends indicate that the community of the grassland at the corridor is slightly different from typical Bupleuro-Miscanthetum sinensis. Those differences might be attribute to the following reasons: Bupleuro- Miscanthetum sinensis are thought to distribute at limestone plateau, retrogressive communities due to the effects of motorcycles are contained within the corridor. This secondary grassland growing many grassland species might be maintained, if periodic mowing are continue. And many species might be maintain its popula tion within the corridor, for those species flowered and/or fruited within the corridor. The line corridor studied has function of habitat for many grassland species which are becoming rare, although it have narrow width. Further, the corridor plays important role ofconduit for plants, especially those producing anemochores seeds.

Acknowledgments Financial support was provided by a Grant-in-Aid for Scientific Research (07308066) from the Ministry of Education, Science, Sports and Culture. Field assistance was provided by Yoshiaki Kameyama, Teruhiko Nakaishi and Akira Kikuchi of Hiroshima University whose help is greatly acknowledged. We thanks the Nougata District Forest Office and Shozo Ohishi for providing the valuable information. We also wish to express our gratitude to Dr. Masamichi Ota, the Director of the Kitakyushu Museumand Institute of Natural History for his guidance and encouragement through the work. Secondary Grasslandat a Line Corridor 125

References

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Appendix. The List ofvascular plants observed within the line corridor at Fukuchi mountain system in May and September, 1996.

Legends 1. The order of family was followed by Tagawa & Iwatsuki (1972) for Pteridophytes and Itow (1989) for Spermatophytes. 2. The names of species were refered to Iwatsuki (1991) for Pteridophytes and Ohwi and Kitagawa (1992) for Spermatophytes other than some naturalized species which were done to Osada (1976).

PTERIDOPHYTA

Lycopodiaceae Hikagenokadzura-ka Lycopodium serratum Thunb. Tougeshiba Ophioglossaceae Hanayasuri-ka Botrychium ternatum (Thunb.) Sw. Fuyunohanawarabi Ophioglossum petiolatum Hook. Kohirohahanayasuri Osmundaceae Zenmai-ka Osmunda japonica Thunb. Zenmai Dennstaedtiaceae Kobanoishikaguma-ka Pteridium aquilinum (Linn.) Kuhn var. latiusculum (Desv.) Underw. ex Hell. Warabi Blechnaceae Shishigashira-ka Blechnum niponicum (Kunze) Makino Shishigashira Thelypteridaceae Himeshida-ka Stegnogramma pozoi (Lagasca) K. Iwats. subsp. mollissima (Fischer ex Kunze) K. Iwats. Mizoshida

SPERMATOPHYTA GYMNOSPERMAE

Pinaceae Matsu-ka Pinus densiflora Sieb. et Zucc. Akamatsu Pinus thunbergii Parlat. Kuromatsu Taxodiaceae Sugi-ka Cryptomeria japonica (Linn, fil.) D. Don Sugi Cupressaceae Hinoki-ka Chamaecyparis obtusa (Sieb. et Zucc.) Sieb. et Zucc, apud Endl. Hinoki 128 Tohru Manabe, Kazuaki Naito and Nobukazu Nakagoshi

ANGIOSPERMAE DICOTYLEDONEAE RCHICHLAMYDEAE Salicaceae Yanagi-ka Salix sieboldiana Blume Yamayanagi Salix subopposita Miquel Noyanagi Betulaceae Kabanoki-ka Carpinus laxiflora (Sieb. et Zucc.) Blume Akashide Carpinus schonoskii Maxim. Inushide Fagaceae Buna-ka Castanea crenata Sieb. et Zucc. Kuri Quercus serrata Thunb. Konara Ulmaceae Nire-ka Celtis sinensis Pers. var. japonica (Planch.) Nakai Enoki Moraceae Kuwa-ka Ficus erecta Thunb. Inubiwa Santalaceae Byakudan-ka Thesium chinense Turcz. Kanabikisou Polygonaceae Tade-ka Polygonum lapathifolium Linn. Ooinutade Polygonum longisetum De Bruyn Inutade Rumex acetosa Linn. Suiba Caryophyllaceae Nadeshiko-ka Cerastium holosteoides Fries var. angustifolium (Franch.) Mizushima Miminagu-

sa Dianthus superbus Linn. var. longicalycinus (Maxim.) Williams Kawaranadeshiko Stellaria aquatica (Linn.) Scop. Ushihakobe Kauraceae Kusunoki-ka Lindera erythrocarpa Makino Kanakuginoki Neolitsea sericea (Blume) Koidz. Shirodamo Ranunculaceae Kinpouge-ka Clematis apiifolia DC. Botanzuru Pulsatilla cernua (Thunb.) Spreng. Okinagusa Ranunculus japonicusThunb. Umanoashigata Thalictrum minus Linn. var. hypoleucum (Sieb. et Zucc.) Miq. Akikaramatsu Lardizabalaceae Akebi-ka Akebia quinata (Thunb.) Decaisne Akebi Akebia trifoliata (Thunb.) Koidz. Mitsubaakebi Stauntonia hexaphylla (Thunb.) Decaisne Mube Menispermaceae Tsudzurafuji-ka Secondary Grassland at a Line Corridor 129

Cocculus orbiculatus (Linn.) Forman Aotsuzurafuji Saururaceae Dokudami-ka Houttuynia cordata Thunb. Dokudami Chloranthaceae Senryou-ka Chloranthus japonicus Sieb. Hitorishidzuka Chloranthus serratus (Thunb.) Roem. et Schult. Futarishidzuka Aristolochiaceae Umanosudzukusa-ka Asarum asaroides (Morr. et Decne.) Makino Tairin-aoi Actinidiaceae Matatabi-ka Actinidia arguta (Sieb. et Zucc.) Planch, ex Miq. Sarunashi Theaceae Tsubaki-ka Eurya japonica Thunb. Hisakaki Guttiferae Otogirisou-ka Hypericum erectum Thunb. Otogirisou Hypericum laxum (Blume) Koidz. Kokeotogiri Cruciferae Aburana-lca Arabis kirsuta (Linn.) Scop. Yamahatazao Saxifragaceae Yukinoshita-ka Deutzia crenata Sieb. et Zucc. Utsugi Hydrangea luteo-venosa Koidz. Kogakuutsugi Schizophragma hydrangeoides Sieb. et Zucc. Iwagarami Rosaceae Bara-ka Agrimonia japonica (Miq.) Koidz. Kinmidzuhiki Potentilla discolor Bunge Tsuchiguri Potentilla fragarioides Linn. var. major Maxim. Kijimusiro Potentilla freyniana Bornm. Mitsubatsuchiguri Pourthiaea villosa (Thunb.) Decne. var. laevis (Thunb.) Stapf Kamatsuka Prunus buergeriana Miq. Inuzakura Prunus jamasakura Sieb. ex Koidz. Yamazakura Rosa multiflora Thunb. Noibara Rosa wichuraiana Crep. Terihanoibara Rubus buergeri Miquel Fuyuichigo Rubus crataegifolius Bunge Kumaichigo Rubus hirsutus Thunb. Kusaichigo Rubus microphallus Linn. fil. Nigaichigo Rubus minusculus Leveille et Vaniot Himebaraichigo Rubus palmatus Thunb. Nagabamomijiichigo Rubus parvifolius Linn. Nawashiroichigo Rubus pectinellus Maxim. Kobanofuyuichigo Sorbus japonica (Decne.) Hedl. Urajironoki Leguminosae Mame-ka 130 Tohru Manabe, Kazuaki Naito and Nobukazu Nakagoshi

Albizia julibrissin Durazz. Nemunoki Dumasia truncata Sieb. et Zucc. Nosasage Indigofera pseudo-tinctoria Matsum. Komatsunagi Kummerowia striata (Thunb.) Schindler Yahadzusou Lespedeza cuneata (Du Mont. d. Cours.) G. Don Medohagi Lespedeza cyrtobotrya Miq. Marubahagi Lespedeza pilosa (Thunb.) Sieb. et Zucc. Nekohagi Lespedeza virgata (Thunb.) DC. Makiehagi Rhynchosia volubilis Lour. Tankirimame Rhynchosia acuminatifolia Makino Oobatankirimame Pueraria lobata (Willd.) Ohwi Kudzu Smithia ciliata Royle Shibanemu Sophora Jlavescens Solander ex Aiton var. angustifolia (Sieb. et Zucc.) Kitagawa Kurara Wisteria brachybotrys Sieb. et Zucc. Yamafuji Wisteria floribunda (Willd.) DC. Fuji Oxalidaceae Katabami-ka Oxalis corniculata Linn. Katabami Euphorbiaceae Toudaigusa-ka Euphorbia sieboldiana Morr. et Decne. Natsutoudai Mallotus japonicus (Thunb.) Muell. Arg. Akamegasiwa Rutaceae Mikan-ka Zanthoxylum ailanthoides Sieb. et Zucc. Karasuzansyou Zanthoxylum schinifolium Sieb. et Zucc. Inuzansyou Polygalaceae Himehagi-ka Polygala japonica Houtt. Himehagi Anacardiaceae Urushi-ka Rhus javanica Linn., pro p. Nurude Rhus succedanea Linn. Haze Rhus trichocarpa Miq. Yamaurushi Aceraceae Kaede-ka Acer palmatum Thunb. Irohamomiji Acer ruftnerve Sieb. et Zucc. Urihadakaede Aquifoliaceae Mochinoki-ka Ilex crenata Thunb. Inutsuge Celastraceae Nishikigi-ka Celastrus orbiculatus Thunb. Tsuruumemodoki Euonymus alatus (Thunb.) Sieb. forma ciliato-dentatus (Franch. et Savat.) Hiyama Komayumi Euonymus japonicus Thunb. Masaki Euonymus oxyphyllus Miq. Tsuribana Secondary Grassland at a LineCorridor 131

Vitidaceae Budou-ka Ampelopsis brevipedunculata (Maxim.) Trautv. Nobudou Ampelopsis brevipedunculata (Maxim.) Trautv. forma citrulloides (Lebas) Rehd. Kirehanobudou Parthenocissus tricuspidata (Sieb. et Zucc.) Planch. Tsuta Vitis Jkxuosa Thunb. Sankakuzuru Elaeagnaceae Gumi-ka Elaeagnus pungens Thunb. Nawashirogumi Elaeagnus umbellata Thunb. Akigumi Violaceae Sumire-ka Viola grypoceras A. Gray Tachitsubosumire Viola hondoensis W. Becke et H. Boiss. Aoisumire Viola mandshurica W. Becker Sumire Viola pumilio W. Becker Fumotosumire Viola yedoensis Makino Nojisumire Viola verecunda A. Gray Tsubosumire Viola violacea Makino Shihaisumire Haloragidaceae Arinotougusa-ka Haloragis micrantha (Thunb.) R. Br. Arinotougusa Cornaceae Mizuki-ka Cornus controversa Hemsley Midzuki Cornus brachypoda C. A. Mey. Kumanomidzuki Cornus kousa Buerger ex Hance Yamaboushi Araliaceae Ukogi-ka Acanthopanax sciadophylloides Franch. et Savat. Koshiabura Aralia cordata Thunb. Udo Umbelliferae Seri-ka Angelica decursiva (Miq.) Franch. et Savat. Nodake Angelica pubescens Maxim. Shishiudo Bupleurum scorzoneraefolium Willd. var. stenophyllum Nakai Mishimasaiko Hydrocotyle sibthorpioides Lam. Chidomegusa Sanicula chinensis Bunge Umanomitsuba

METACHLAMYDEAE Clethraceae Ryoubu-ka Clethra barbinervis Sieb. et Zucc. Ryoubu Ericaceae Tsutsuji-ka Lyonia ovalifolia (Wall.) Drude var. elliptica (Sieb. et Zucc.) Hand.-Mazz. Nejiki Primulaceae Sakurasou-ka Lysimachia clethroides Duby Okatoranoo 132 Tohru Manabe, Kazuaki Naito and Nobukazu Nakagoshi

Lysimachia japonica Thunb. Konasubi Ebenaceae Kakinoki-ka Diospyros kaki Thunb. Kakinoki Styracaceae Egonoki-ka Styrax japonica Sieb. et Zucc. Egonoki Symplocaceae Hainoki-ka Symplocos coreana (Leveille) Ohwi Tannasawafutagi Symplocos lucida Sieb. et Zucc. Kuroki Oleaceae Mokusei-ka Fraxinus lanuginosa Koidz. Aodamo Ligustrum japonicum Thunb. Nedzumimochi Gentianaceae Rindou-ka Gentiana scabra Bunge var. buergeri (Miq.) Maxim. Rindou Swertia japonica (Schult.) Makino Senburi Apocynaceae Kyouchikutou-ka Trachelospermum asiaticum (Sieb. et Zucc.) Nakai var. intermedium Nakai Teika- kadzura Asclepiadaceae Gagaimo-ka Cynanchum atratum Bunge Funabarasou Cynanchum paniculatum (Bunge) Kitag. Sudzusaiko Rubiaceae Akane-ka Galium trachyspermum A. Gray Yotsubamugura Mitchella undulata Sieb. et Zucc. Tsuruaridoushi Paederia scandens (Lour.) Merrill var. mairei (Leveille) Hara Hekusokad-

zura Verbenaceae Kumatsudzura-ka Clerodendron trichotomum Thunb. Kusagi Premna japonica Miq. Hamakusagi Labiatae Shiso-ka Clinopodium chinense (Benth.) O. Kuntze var. parvijhrum (Kudo) Hara Kuru- mabana Clinopodium gracile (Benth.) O. Kuntze Toubana Clinopodium gracile (Benth.) O. Kuntze var. multicaule (Maxim.) Ohwi Yama- toubana Clinopodium micranthum (Regel) Hara Inutoubana Plectranthus inflexus (Thunb.) Vahl ex Benth. Yamahakka Prunella vulgaris Linn. var. lilacina (Nakai) Nakai Utsubogusa Scutellaria indica Linn. Tatsunamisou Scrophulariaceae Gomanohagusa-ka Pedicularis resupinata Linn. Shiogamagiku Plantaginaceae Oobako-ka Secondary Grassland at a Line Corridor 133

Plantago asiatica Linn. Oobako Caprifoliaceae Suikadzura-ka Lonicera gracilipes Miq, Yamauguisukagura Lonicera japonica Thunb. Suikadzura Viburnum dilatatum Thunb. Gamadzumi Viburnum erosum Thunb. Kobanogamadzumi Valerianaceae Ominaeshi-ka Patrinia scabiosaefolia Fisch. Ominaeshi Patrinia villosa (Thunb.) Juss. Otokoeshi Valeriana fauriei Brip. Kanokosou Campanulaceae Kikyou-ka Adenophora triphylla (Thunb.) A. DC. Saiyoushajin Codonopsis lanceolata (Sieb. et Zucc.) Trautv. Tsuruninjin Peracarpa carnosa (Wall.) Hook, fil. et Thoms. var. circaeoides (Fr. Schm.) Makino Tanigikyou Platycodon grandiflorum (Jacq.) A. DC. Kikyou Compositae Kiku-ka Anaphalis margaritaceae (Linn.) Benth. et Hook. fil. var. angustifolia (Franch. et Savat.) Hayata Hosobanoyamahahako Artemisia feddei Lev. et Van. Himeyomogi Artemisia japonica Thunb. Otokoyomogi Artemisia princeps Pampan. Yomogi Aster ageratoides Turcz forma leucanthus Honda Yamasirogiku Aster ageratoides Turcz. var. ovatus (Franch. et Savat.) Nakai Nokongiku Aster scaber Thunb. Shirayamagiku Chrysanthemum indicum Linn. Shimakangiku Cirsium buergeri Miq. Himeazami Cirsium japonicum DC. Noazami Cirsium lineare (Thunb.) Schultz Bip. Yanagiazami Cirsium lineare (Thunb.) Schultz Bip. var. discolor Nakai Urajiroyamagiazami Erechtites hieracifolia (Linn.) Rafin. Dandoborogiku Erigeron annuus (Linn.) Pers. Himejyohn Eupatorium chinense Linn. var. simplicifolium (Makino) Kitam. Hiyodoribana Eupatorium lindleyanum DC. Sawahiyodori Gnaphalium japonicum Thunb. Chichikogusa Ixeris dentata (Thunb.) Nakai Nigana Kalimerisyomena Kitam. Yomena Lactuca indica Linn. var. laciniata (O. Kuntze) Hara Akinonogeshi Leibnitzia anandria (Linn.) Turcz. Senbonyari Ligularia japonica (Thunb.) Less. Hankaisou Petasites japonicus (Sieb. et Zucc.) Maxim. Fuki 134 Tohru Manabe, Kazuaki Naito and Nobukazu Nakagoshi

Picris hieracioides Linn. var. glabrescens (Regel) Ohwi Kouzorina Saussurea pulchella Fisch. ex DC. forma subintegra (Regel) Kitag. Himehigotai Senecio integrifolius (Linn.) Clairville var. spathulatus (Miq.) Hara Okaogur-

uma Solidago altissima Linn. Seitakaawadachisou Solidago virga-aurea Linn. var. asiatica Nakai Akinokirinnsou Synurus excelsus (Makino) Kitam. Habayamabokuchi

MONOCOTYLEDONEAE

Liliaceae Yuri-ka Aletris luteoviridis (Maxim.) Franch. Nogiran Aletris spicata (Thunb.) Bureau et Franch. Sokushinran Disporum sessile Don Houtyakusou Heloniopsis orientalis (Thunb.) C. Tanaka Syoujyoubakama Hosta albo-marginata (Hook.) Ohwi Kobagiboushi Lilium cordatum (Thunb.) Koidz. Ubayuri Lilium leichtlinii Hook. fil. var. tigrinum (Regel) Nichols. Kooniyuri Liriope platyphylla Wang et Tang Yaburan Polygonatum odoratum (Mill.) Druce var. pluriflorum (Miq.) Ohwi Amadokoro Polygonatum falcatum A. Gray Narukoyuri Smilax china Linn. Sarutoriibara Tricyrtis affmis Makino Yamajinohototogisu Dioscoreaceae Yamanoimo-ka Dioscorea japonica Thunb. Yamanoimo Dioscorea quinqueloba Thunb. Kaededokoro Iridaceae Ayame-ka Sisyrinchium atlanticum Bicknell Niwazekisyou Juncaceae Igusa-ka Luzula capitata (Miq.) Nakai Sudzumenoyari Luzula multifbra Lejeune Yamasudzumenohie Commelinaceae Tsuyukusa-ka Commelina communis Linn. Tsuyukusa Gramineae Ine-ka Agropyron tsukushiense (Honda) Ohwi var. transiens (Hack.) Ohwi Kamojigusa Agrostis clavata TRiN.var. nukabo Ohwi Nukabo Arundinella hirta (Thunb.) C. Tanaka Todasiba Brachypodium sylvaticum (Huds.) P. Beauv. Yamakamojigusa Calamagrostis arundinacea (Linn.) Roth var. brachytricha (Steud.) Hack. Nogar- iyasu Cymbopogon tortilis (Presl) A. Camus var. goeringii (Steud.) Hand-Mazz. Secondary Grassland at a Line Corridor 135

Ogarukaya Imperata cylindrica (Linn.) Beauv. var. koenigii (Retz.) Durand et Schinz Chigaya Miscanthus sinensis Anderss. Susuki Oplismenus undulatifolius (Ard.) Roemer et Schultes var. japonicus (Steud.) Koidz. Kochijimizasa Paspalum thunbergii Kunth Sudzumenohie Pleioblastus chino (Franch. et Savat.) Makino var. viridis (Makino) S. Suzuki Nezasa Poa annua Linn. Sudzumenokatabira Setaria glauca (Linn.) Beauv. Kin-enokoro Themeda japonica (Willd.) C. Tanaka Megarukaya Zqysia japonica Steud. Shiba Araceae Satoimo-ka Arisaema japonicum Blume Mamushigusa Cyperaceae Kayatsurigusa-ka Carex breviculmis R. Br. Aosuge Carex Jbribunda (Korsh.) Meinsh. Hikagsuge Carex nervata Franch. et Savat. Shibasuge Crchidaceae Ran-ka Cephalanthera falcata (Thunb.) Blume Kinran Epipactis thunbergii A. Gray Kakiran