Mosaic of various seral stages of vegetation in the Satoyama, the traditional rural landscape of Japan as an important habitat for butterflies

2007·1.9 MB·

TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan
axtwh 7>'ansI.epidS.oc.JqtJaf5r8(1):69-90,Junuary2007

Mosaic of various seral stages of vegetation in the SatQyaina ,the traditional
rural landscape of Japan as an important habitat for butterfiies
YasuakiNisHiNAKA* MinoruIsHii
and
Entomelogicul Laboratory, Graduate Schoo lof Agricultur eand Biologica lSciences,
Osaka PrefectureUniversitSya,kai,Osaka,599-8531Japan

Abstract Assemblage snmctures of butterfl iweerse studied by the transec tcount method in Farm
ancl Coppice landscap eisn a Satoyama, the traditional rural landscape of Japan, around Mt Mikusa
in northcni Osaka, central Japan fi'om April to Oetober ,2004. Transects in Farm and Coppice land-
scapes were classified into three and six Iandscape components, respectively, aceording to vegeta-
tion and landscape. To analyzc charactcristics of buttcrfi aysscmblage struetures in the Sato.vama,
the seral rank of each butterf wlays determine bdased on seral stages in NN･hich the main larva flood
plants occur, and compositions ef each seral rank species were cornpared among landscapes and
landscap eeomponents. A tota lof 1322 individua bleslongin gto 56 species from seven faniilies
were recoTded throughout the whole study area. Both species richness and dcnsity of butterfiies
were highe rin Coppice landscap e(4 6and 20.4 respectively) than in Fanm landscap e(3 9and 13.1
respectively), while species diversit }(,1-A )and evcnness (.J w)crc higher in Farm landscape (O,91
and O.75 respectively) than in Coppice ]andscape (O.8 5and O,66 respectively). Coppice landscape
and the whole study area were characteri7ed by the dominance of three dwarf bamboo feeders,
Lelhe diana, L, si(/elis and Neope goschkevitschi i,ln addition, both species richness aiid densities
of univoltine butterfli eisncludin ggrass ,vielct and deciduou soak tree feeder swere abundant in
Coppice landscap eA.nalysis based on the SR index revealed that species richness and densitie sof
butterfli edsependent on plants occuring in young forest sand deciduous broad-leaved forest sas lar-
val foods were high in Coppice landscape ,whereas those dependen ton plant osecuring in short to
tal lgrassland swere high in Farm landscape .On the other hand, some species of intermediat erank
showed a wide range of distributio inn Farm and Coppice landscapcs in thc study arca. The results
demonstrat tehat buttcrfi idccspenden ton plants in wide rangcs of seral stages of vegetation inhabit
,Sato.vam aT.he mosaic structure of vegetation in terms ot' the seral stage is important for conserva-
tion of species diversit iofe sbuttcr'fi iicn sSato.vama.
Key words Butteril yassemblage, seral stage, species diversity J,apan, Sato.vama ,coppice, farm,
transectcount,
Introduction
The "Sato},amaT' in a broad sense is the tradition arlural landscap eof Japan consisting of
(Sato},aimna farmlands fields
coppices a narrow sense) and such as paddy and semi-natural
(Ishi20i0,1a), havebccn for byhuman intervention
grasslands which maintained centuries
farming(Takeuch20i0,3;Ishii2,001a,b).Sato>)amais
such as coppicing, mowing and an
importan thabita ftor an abundant variety of wjldlife includin gspecies endemic to Japan and
eastern Asia, although it is secondary in an ecological sense (Ish i2i0,01a, h).
However, the fue land fertili zreevrolutions in the 1960s resulted in the destructi oannd
decline inhabitinSgatoyama(e.
abandonment of coppices, and a consequent of FwIilldalbifaeta , g,
Moriyama, 1988; Ishii et al., l993; Hattori et at,, 1995; 1997), Habitats for wildlife
in Satoyama farmland shave also been degraded by paddy fiel dconsolidation fbr the mech-
anization of agriculture, the use of agricultural chemicals such as insecticide sand herbi-
*Corresponding author. E-mail: [email protected]
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70 Yasuaki NisHiNAK Aand Minoru IsHii
cides, and abandonment or destructio nof thatch field ssince the 1960s (Moriyam a19,97;
Ishii ,20e5), Thus it is recognized that the recent change in land use is aprimary cause for
the declin eof biodiversi itny Japan ,and indced many species ef wild animals and plant sin-
cluding butterfl iiens Satoyama now appear on Red Data List sof the Environmcntal
Ministr yof Japan and loca lauthorities in Japan ,Although no native butterfl yspecies has
become extinct in Japan, there has becn a considerable net decline of local butterfl ypopula-
tions inSato},ama(Ishi1i9,96;Nakamura, 2003; Inoue,2005),

In this study we investigat tehde butterf lasysemblage in differe nltandscap ecomponents of
a Sato),am ain northern Ogaka, central Japan, and analyzed the relationship between vegeta-
tion type and species divcrsi tofy butterfli etshere by using a newly developed index based
on seral stages in which the major food plant sof each butterf lspyecies occur. From the re-
sultg, we discug sthe characteristics of butterf lasysemblage in Satoyama, and the conserva-
tion of gpecies diversit yof butterfli etshere,
Study site and methods
This study was carried out in a Sato.yam ain and around Mt Mikusa (34057" N1,.35022'E,
564 m a.s.].), northern Osaka, central Japan, from April to October, 2004. A nature reserve,
`LMt, Mikusa Zeph)rrus Coppice" is situated on the southeastern slope of Mt Mikusa, and a
]ocal conservation body has been conducting the management of the coppice, There arc
remnants ofcoppice woodlands that are dominated by decjduou soaks, such as 2ueicu sser-
e. e. farmlandsincluding fields,
rata, acutissima, aliena, etc,, and rice paddies, vegetable
chcstnut orchards, etc., in this area, We diyide dthe study area into two landscape sF,arm
and Coppice ,and fixe da transec troute through major landscap ecomponents in the two
landscape s.The transect was divide dLnt o26 sections according to landscape components
(Fig 1.). Details ofeach section are shown in fable 1.
In the Farm landscape t,he transec twas divide dint othree sections, corresponding to three
landscap ecomponents, P]ai npaddy, Village and Yatsuda (padd fyield dsevelope don a nar-
row valley bottom) ,The transect of Plain-padd ysection was fixe don a leve ebetween rice
paddie son the plai nwhere frequen tmowing and grass-burni nwegre conductcd and short
herbs of Leguminosae and Gramineae dominated .The transect of Village section was fixed
along a narrow road through settlements with horticultura lplants ,plantatio nosf thc
Japanese cypress, Chamaec')li?ar iosbtuse ([faxodiace tahee )J,aptmes ecedar, Cr)ptonzeria
.iaponica (Cupressace aeth)e, chestnut Castanea crenata (Fagacea ea)nd the bamboo,
Ph.ytlostac hsyps., gardens ,and open space with short and tal lherbs ,climbing plants and
the dwarf bamboe, Pleiohlast ucshino (Gramineae )T.he transect of the Yatsuda section
was fixe dalong a road surrounded by rice paddies ,levees and slopes with short and tall
herbs such as bnperata cytindrica and Miscanthus sinensis (Gramineae a)b,andoned fields
with tall herbs and shrubs such as M, sinensis and Athizzi jaL{libr (iLsesgiunminos achees)t,-
nut orchards, Japanese cypress plantatio nansd coppiccs on the vaLley bottom.
In the Coppice landscape t,he transect was divide dint o23 ･section sbelonging to six land-
scape components, Glade, Cutover-lan dC,oppice edge, Medium coppice, Tall coppice and
Cypress plantatio n(Fi g1,) ,All components of the Coppice landscape were dominated by
dwarf bamboo with high densitie son the fores tfloo rexcept Cypress plantatio nG.lade sec-
tions (Gl and G2) were along traiI sthrough an open grassy slope, where short herbs such as
Clinopodi"m gvacil e(Labiat aean)d V}ola gr)poceras (Violace ateal)l ,hcrbs such as
Phytolacca americana (Phytolacca cancda cM). siHensis, and dwarf bamboo 1-3 m in
height were dominant ,The Cutover-lan dsection (CL) was established in a 4th-year cop-
pice after cutting. The dominant trees were e. acutissima and e. aliena 3-5 m high, and
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Satoyatn aas an Importan tHabitat for Butterfiies 71
Fig .1. Locatio nof the study area (a )and transec troutes established in Furm (b )and Coppice (c)
landscap eisn and around Mt Mikusa ,northern Osaka ,central Japan.
the ground was covered with bushes of dwarfbamboo from 1 to 3 m in height ,Coppice-
edge sections (CEI-4 w)ere fixe dalong a trai lon the boundary of a chestnut orchard (CE1)
or coppices of deciduou soaks (CE2-4 a)nd open space. Section sCE3 and 4 were adjacent
to 1st to 3rd-yea rCutover-lan ddsominated by Q. ac"tissima and C. aliena. Medium-cop-
pice sections (MCI-4) were along a trail through coppices consisting of e, acutissima and
e, atiena about 10 m in height .[fa11-coppic esections (TCI-8 )also occurred along a trail
through coppices dominated by deciduou soaks, although the mean heights (abou t15 m)
were taller than those of the Medium-Coppice sections. Cypress-plantat iseocntions (CPI-
4) occurred along a trai1 on boundaries between Japanese cypress ptantation s(mea nheight:
18 dominatedbydecjduous (15 inheighO.
m) and coppices oaks m
Data cotlection on butterf iasysemblages was conducted using the transec tmethod (Pollard,
1977, 1984; Thomas, 1983; Pollar dandYates, 1993) .Counts of butterfl iweerse conducted
in all sections of the transect throughout the study area a total of 14 times, twice a month
from April to October ,in 2004. Each count was conducted from 9:30 to 15:OO hrs local
time as a rule under fin eweather conditions with calm to ligh twinds, We recorded the
number of adult individual sof each butterfi yspecies sighted withjn a 1O m width (5 m each
side of the recorder) and up to 5 m heigh talong the transect. Butterfii neots readily iden-
tified were captured by net and released immediately after identification.
Analysis
Community indices

We calculated species diversi tbyy the Simpson's index of diversit y1,-A (Simpso 1n9,49),
and species evenness by the equitability index ,J' (Pielo u19,69) as follows:
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72 Yasuaki　Nls田 NAKA 　and　Minoru　IsHII
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Sa　to．vama 　as　an　Important　Habitat　for　Butterflies 73
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and
Table 2.Five seral stages determined in this study, vegetation type and characters of p]ants in each stage.
SeralstageVegetation Characters of plant isn each seral stage
type
Hcrb and grass Climbing plant Woody p]ant
123 Short grassland Short herbs in grassland
Ta]1 grass]and Tal1 herb sin grassLand
Young foresL Tall herb sin young Climbin gplant isn foi'estDcciduousshrubs
forestShort edgc
herb sin edge of Young deciduous trees
deciduousforest

4 Deciduou sforest Tall herb sin deciduous Climbing plant isn Dcciduous trccs
forestShort deciduousforest

herbs Deciduous in
with shrubs
relatively high shade deciduous forest
torelance Young evergreen trees
5 Ex,ergree nforest Herbs in cvergreen Climbing plant isn Evcrgrccn trces
forest forest
evergreen
Shrubs in evergreen
forest
1-A (n,-1)(LANP-1)
=1-En,
J'=-E ((n,/Nlo)gS(nliV)

'is
where n, the number of individua] sof the ith species, 7Vis the tota lnumber of individuals
of al 1component species, and S is the total number of species.
Tb group the butterfl yassemblages from nine landscape ¢omponents according te the simi-
larit yin species composition or assemblage structures, we conducted UPGMA cluster
analyses based on Pianka's a index (Piank 1a9,73) as fo11ows:
(n,,n2i)()n1,(,1(7)2V}()n],,ZIV})2)ieA(,AJ})
a =(E
where Ni and IV 2represent the total number of individua lors species in landscape compo-
nents 1 and 2, ni, and n2, represent the number of individual sor species of ith gpecies or
group in landscap ecemponensts 1 and 2, respectively,
Ecological butterflies
classification of and analyses of assemblage structures
We classified butterfii ersecorded in this study into 2-4 groups according to voltinism and
distributiondescribedinNishinaka Ishii(2006)W,e
geographical as and compared species
richness and abundance of each butterf lgyroup between landscape sand among landscape
In determinedthe (SR)for butterfly
components. addition, we seral rank each species ac-
cording to seral stages in which major 1arva lfood plant soccur, and analyzed butterfl yas-
semblages in the study area by the proportio onf each seral rank (SR) .Major larva lfood
plant sand seral stages in which the plant species occur were determine daccording to Unno
and Aoyama (1981 )F,ukuda et at. (198 21,983, 1984a, b), Makino (198 21,983) and Okuda
<1997 ).Seral stages of vegetation (St w)ere categorized into five stages (stl- 5ac)co,rding
to the successional proces sin temperate regions in central Japan ,short grassland (stl t)a,ll
grasslan d(st2 y)o,ung fores t(st3 d)e,ciduou sfores t(st 4an)d evergreen fores t(st 5()[[hble
2),
SR fbr each butterf ispyecies was calculated as follows:
SR=2 ･
£ stLltt-1
wherc sti is the score of ith seral stage (1-5 )an,d n is the total number of seral stages in
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Saio.vam aas an Important Habitat for Butterfli es 7S
which foodplant osf the species occur. St and SR for each species are shown in Tbble 3.
[Ib group butterf lasysemblages from nine landscap ecomponents with respect to the sera]
UPGMA based Pianka's
rank of component species, we conducted cluster analyses on
index a for the evaluation of overlap of species and/or abundance of species belonging to
each seral rank.
Results
Characteristicsbutterfiy inFarm Coppicelandscapes
of assemblage and
A tota lof 1322 individua lbselongin gto 56 species from seven familie wsere observed in
the study area ([rbb 4l)e ,Out of the 1322 individual 4s1,2 and 910 individua lbselongin gto
39 and 46 species were seen in Farm and Coppice landscape sr,espectively, Community in-
dices ,1-A and J' were higher in Farm landscap e(O. 91and O.75 respectively) than in
Coppicelandscape(O.85 O.66).
and
The dominant species in the study area were Lelhe diana (290 jndividual sI)V,eop ego,s'chke-
(148)L,. (101)},iPthima (94) CoXias (69)indecreasing
vitschii sicetlis atlgtts and erate
order, and the proportion of the fiv especies was 53% in the number of individua l(s[[Uble
4). The top fiv edominant species in Coppice landscap ewere al 1satyrids, the same species
as those in the whole study site, except that Minois dr.va swas one of the five deminant
species instea dof C. enate. In contrast, in Farm landscape five dominant species were all
grasslan dspecies and differe dconsiderably from those in the whole study site: they were C.
erate, L.vcaena phlaeas ,Pieris rapae, V. argus and Eveies argiades in decreasin ogrder.
Out of the 56 species recorded from the study area, 17 and 1O species were observed only in
Coppice and Farm landscap erespectively, while 29 were common between the two land-
scapes ([fab 4l)e, Major species unique te Coppice landscap ewere woodland species such
as Er:ynni smontanus, Ndratttft zjaponic aFl,avoniu scognatus, Arg.vnni spaphia and
M.vcalesi s.hancisca. On the other hand, openland species such as C, evate, P. rapae,
Zi.7.eer intaaha, C. vn.thia cantui and Ptirnara gittta tweare among the major species unique
to Farm landscape. The 29 species seen in both landscape sinclude dthe species of forest
edge such as }i .angus, Pieri smelete, IVepti sscrppho, T:Fzores svaaria and Eurema hecabe.
The top four dominant specics in the study arca, L. diana, N. goschkevitschi iL,. sicellis and
Y. ai:gus, were of the Sino-Japanes geroup ,while the fift dhominant species, C evate, was
of the Siberia ngroup (Tabl 4e) .Bcsides the dominant species, Sino-Japanes sepecies such
as E. montanus, F. cognatus and M, .francisc awere seen in Coppice landscape ,while
Siberian species such as L. phlaeas, P. r(u)ae and E. acgiades and Malayan species such as
Z, maha, C, cantvti and P, g"ttata were abundant in Farm landscape .On the other hand,
Sino-Japanes sepecies such as P, melete, T. varia and C, acuta, Siberia nspecies such as M.
dr.va sand N. sqt?pho, and Malayan species such as E. hecabe and K. canace were seen in
both landscape s.Thus species richness of the Sino-Japanes egroup (3 1spp.) was 1arger
than those of the Siberia n(1 1spp,) and Malayan (14 spp,) groups in the whole study site,
and the proportio nof the Sino-Japanes geroup in species richnes swas highe rin Coppice
landscap e(65% )than in Farm landscap e(51% )(Fi g2.) .In densit yt,he proportion of the
Sino-Japanes geroup was much highe rin Coppice landscap e(83% )than in Farrn landscape
(29%) w,here the Siberia nspecies was the most dominant .Furthermore t,hree out of the 31
Sino-Japanes sepecies were those endemic to Japan, and the proportio inn densit ywas larg-
er in Coppice landscape (27%) than in Farm landscape (19b )due to the abundance of N.
goschkevitsc hainid L. sicettis,
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76 YasuakiN MinorulsIIII
［sH艮NAKA 　and 　 　
Table　3，　The　seral　rank （SR）of　butte且ry　species 　recorded 　in　thi　sstudy．　Major 】arval　food　plants，　and　their
　　　　seエal　stages 　are　also　shown ．　See　text　fQr　calculation 　Qf　SR．
Species　namc M　a　jo　r　　］（aFrvaami］　lfyeo）　d　　pl　an　　t 　1　　2　Ser3　a　lst4a　ge5　SR
Hesperiidaeセ セ リチ ョ ウ科
　Ei》，nnis 　inOntanuh ’ミヤマ セセ リ Deciduou　soak （Fag） 　 十 十 65544533
　D α競の 皹 1り，sダイミ ョ ウセ セ リ Black　bryOIly（Dio） 　 十
　Thoressa　vanla コ チャ バ 不セセリ Dwarfbamboo （Gra） 十 十 十
　∬sote’inon 　tarrフtro1spilus ホソバ セ セ リ Grass（Gra） 　 十 十
　Potanthttsflavttm キマ ダラセセ リ Grass（Gra） 十 十 十 十
　Po　1．vtremis 　pettucidaオオチ ャバ ネセ セ リ Grass（Gra） 　 十 十 十
　Pelopidas　mathias チャバ ネセセ リ Grass（Gra） 十 十 十
　Pa　rnara 　gzattatイa チモ ンジセセ リ Grass（Gra） 十 十 十
Papilionidaeアゲハ チ ョ ウ科
　Bvasa　alcinottsジヤ コ ウアゲハ Aristoi’ohcia　SpP ．（Ari） 十　　十 485
　Graphiutn　sarpedon ア オス ジアゲハ Camphor 　tree〔Lau） 十　　十
　Papili　xouthus ナミアゲハ Zitnthoxylum　SPP ．（Rut） 十 ．
　P．helenusモ ンキアゲハ Zとirithox｝’lum　SPP ，（Rut．） 十 十 65566
　P．Pfotenot’クロ アゲハ Zanrhox），lum　SPP ．（Rut） 十
　P．ma 〔lilenttts オナガ アゲハ Orixa　sPP ．（Rut） ⊥T
　　pp．．Jbniaaαncokiriカミラヤスマアカゲラハス アゲハ ZZiとinntthho‘，．xt｝），’lluint　　TmSSPPPP ．．（（RRuしttt）） 十十 十十
Pieridueシロ チ ョ ウ科
　Colia、g　erate モ ンキチ ョ ウ Clover（Leg） 十 　 　 　 十 14313
　Eurema 　hecabeキチ ョ ウ Bush　cl〔，ver （Leg） 十 十 十 十 十
’
　Anthochari　sscol］　mus ツマ キチ ョ ゥ Bitte　rcress 〔Cru） 十 十 十 十
　Pieris　rapae モ ン シロ チ ョ ウ Cabbagじ（Cru） 十 　 　 十
　P．Jnelete ス ジグロ シロチョ ウ YcllOW　cress （Cru） 十 十 十 十
Lycaenidaeシジミチ ョ ウ科
　Narathttrα．iapofticムaラサキシジ ミ Evcrgrecn　oak （Fag） 　 　 十 十 十 766
　　JAanptoingii．cuas　αstatepielsit’arkiaミta．ス ＊イウロラオナナミガアシカジシミジ ミ DDeecciidduuoouしsLs　　ooaakk （（FFaagg）） 　 　 十十 十十 　 ．
　1”avonius 　cognatus ：i‘ヒ ロ オビ ミドリシジミ Deciduous　oak 〔Fag） 　 　 十 十 　 675611634
　　RCaαρ」如al卿a　aヴra3taノ猷ト擢ラa コフツシバジメミ JWaipsatnaef’isae　　SaPnPdro．mc（dLaeg）〔Eri） 　　 　　 十十 十　 　十
　Tarahα　ha．madafi ゴ イシ シ ジ ミ Aphids§ 　 十 十
　　LZi｝，zCCeieenraia　　pmahhtaaeaヤsベマ ニトシシジジミミ SOhxeaelpis　　sSePiPTe，l（O（PxQaユ｝） 十十 　 　 　 　　
　Ceiαsti’ina　argiolus ル リシジミ Kudzu （Leg〕 　 十 十 十 十
　Everes　argiades ツバ メ シジ ミ Tares（Leg） 十 十 十
　C齠 濡 o傭 α ウラギン シジミ Kudzu （Leg〕 十 十
Libytheidaeテ ングチ ョゥ科 ∫．
　ムめyr加α副 翻 テ ン グチ ョ ウ Hackberry （Ulm ） 十　　十 0
Nymphal’idae タテハ チ ョ ウ科
ADNae　rmpgyohfraarongon　ns，yeangnai　nsrac　tasnl＊aadtn’‘aメ）’＊omス＊enオeグ4 オロ’　’ウkヒクラliモギウガンモタスンヒジョヒウョモウモン ン FFFooorrreeessstl　　　tVVViiiooollleccttt（（〔VVViiio（o》））） 十十十 555535552552
Arg．1，nnis　paphi．a＊＊ ミドリヒ ョ ウモ ン Fores【Violet（Vio．） 十
Arg．vreus 　h）’perbius＊＊ ッマ グ ロ ヒ ョ ウモ ン Vlo】et （Vio） 十 十
Ladoga　cα｝nitiα イチモ ンジチ ョ ウ ムonicera 　SPP 、〔Cap） 十
L．glon）icaアサマ ．イチモ ン ジ ムo’1．icera　SPP ．〔Cap） 十
Nゆ諭 、照1η物o コ ミス ジ Kudzu 〔Lcg） 十
Pol．v，gonia　c−CtLtreuJn キタテハ 魚 硼 1乙‘∫’SPP ．（Mor ） 十　　十
Kaniska　canace ル リタテハ Snli‘乙lr　‘／hin‘1　（Lil） 　 　 十 　
Vane∬α　indicアa カタテハ Ralnie（Urt） 　 十 十 十
Cynthia　card “iヒメ アカ タテハ Arす8刀廊∫どα SPP．（Corn） 十 十
一
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TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan
Scrtoyam aas an Importan tHabitat for Butterflies 77
Dichorragia nesimachus - - ::'} 'fi )' Meiiosma spp. (Sab) +++7
Apatura metis' - A 7 A;' k Saii xbabytonie (aSal) ++ 6
Satyr'id Va' ct' 7 pt ff 'g VF['
}?)thimaargus t p t9 7 :} '- }V' it 7 pt Gras s(Gra) ++ +++++++++++ 34666344
Minois dn'as V' "x 7 1 f n V Grass (Gra) +
Lethe h'ice"tis Et )i jetf 'g V Dwarf bamboo (Gra)
L. diana b P e )t lb" Dwarfbamboo(Gra)
Ne(ij) egoschkevitschi -i)' I NS' R' 5r i'7 tt tJ lf Dwarf bamboo (Gra)
M.ycales giostama t pt V' ft t7 pl Grass (Gra) ++
M..francisc aU V "r 7 S '? Gras s(Gra) +
Melanit ipshedima b U U 7 f H V Grass (Gra) +
Family names of larva lhost plants .Ari, Aristolochiaceae ;Cap, Caprifbliace aCeo;m, Compositae;Cru,Leg,Sab,
CruciferaeD;io.DioscoreaeeaeE;ri,Ericaceae;Fag, Fagaceae; Gra, Gramineae; Lau. Lauraceae;

Leguminosae; Lil,Liliaceae;Mor, Moraccae; Oxa, Oxa]idaceae;Pol,Polygonaceae; Rut,Rutaceae;

Sabiaceae;Sal,Salicaccae;Ulm, Ulmaceae: Urt,UrticaceaVei:o,Violaceae.

eAlthough 72zrak haamada is carnivore, -,e rreated this species as "Dowarf bamboo feeder "becauselarval
food is the aphids inhabitin gon leaves of dwemf bamboos.
*: Hairstrea kspecics; **': Fritilla rsypecies.
Species richness of uniyoltine species was large rin Coppice landscape (12 spp,) than in
Farm landscap e(fiv sepp.), and all the univoltine species in Fann landscape were also
recorded from Coppice landscape (Tabl e4). A total of seven univoltine species were
recorded only from Coppice landscape include dthree deciduou soak feeders E,. montanus,
J, saqpestriata and F. cognattts, thre efbrest-vio lfeeteders ,A. ruslana, Nephargynnis
feederA,nthocharis
anatlyomene andA. pophia and a crucifer scoZymtts.
Characteristic osf butterfl yassemblage in each landscape component
Specie srichness of butterfi iweass differe namtong the nine lanfdllsecialpl ecomponents: i twas
highes tin Yatsuda (3 3spp,), followe dby Coppicc edge (30) , coppice (29) M,edium
(26)G,lade(24) Village(21), it lowestinCutover-la(n1d1)
coppice and while was and
Cypress plantati o(n11 )followe dby Plain paddy (13 )(fabl 4e). Density of butterfli ewsas
higher in Coppice landscape (20, 4individualsA(m )than in Farm landscap e(13,1 a)nd a]so
differen tamong landscap ecomponents. The densit ywas highes tin Medium coppice
(26,9 )fb,llowed by Coppice edge (24.2 )Gl,ade (2 1,1) ,while it was lowest in Village (8.0),
followed by Yatsuda (14. 1an)d Plain Paddy (15.4').
SpeciesdiveTsit(y1-A) 1argely landscape
ofbutterfly assemblage varied among components
(Tab l4e). It was highcst (>O.90 i)n three Iandscape cQmponents, Villag eY,atsuda and
Glade,followedby Cutover-land(O.90) Coppice (O.89), it lowestin
and edge while was
Cypress (O.55I)n.Cutoverland, but (J)
plantatio n not o 'nly species cliversity also evenness
was high (O.9 2i)rrespect iovfe poor species richness (1 1spp,), but the species diversi tiys
overestimated due to the shert transect and the lowest abundance (32 individual sI)n. con-
trast, species diversi twyas relatjvely low (O.76 i)rrespect iovfe high species richness (29
spp,) in [[hl lcoppicc. This is due to the dominance of L. diana, as shown by the low eve-
(O.63).
ness
The number of species unique to a particula rlandscap ecomponent ("uniq uspeecies") was
the larges itn Yatsuda ,Glade and Tal ]coppice (3 spp. each), and they were 7larak haamada,
Ladoga glorijic aand Apatura metis in Ytttgud aI,]blytremi spelhtcida ,Byasa alciizous and
Anthochari sscolymus in Glade, and Ptipili ohelenus, P. maackii and Melanitis phedima in
Tttl cloppice (rlab 4l)e .In addition, P. x"thus andArgyronome r"slana, Jlriponi scaaqpestri-
ata and Dichorragia nesimach"s, and Giephitt msaripedon were unique to Coppice edge,
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78 Yasuaki　NlsHtNAKA　and 　Min 〔｝ru　IsH【匸
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