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ISSN1346-7565 ActaPhytotax.Geobot.56(1):97-104(2005)
Cytogeographyof theAsterageratoidesComplex (AsteraceianeKo)rea
AKIKO SOEJIMAi, JAE-HONG PAK2, TATSUYOSHI MORITA3
and MOTOMI ITo"
t CoUege ofintegrate danAdr Stcsiences O,saka Prelfectur eU}iiversi S4aMkai, Osaka 599-8531, .lapan ;2
DepartmentofBiotogCoMtlegeofNaturSactience,K),ungpook Ndtional VhivensicM7Zieg"702-701,Korea;
34Bietogica ilnstitu thea,c"lt yqfEd"cation and H"man Science N,iigat cUihiversit Nyiigata, 950-2I8I ,Japan;
Department of Generat Syste mStsudie sG,rad"ate School qfArts and Science 7sb,kyo U}zivei:s 7ibck}ylo 153-
8902, Japan
This is the firs rteport on chromosome number of the Aster ageratoidks complex in Korea, which i sone
of the representatiyes of Sino-Japane seleements. Our cytogeographical research reveals tha tboth diploids
and tetraploids occur in Korea. Diploids are dominant and widety distribut tehdrough Korea including
Cheju Island w,hile tetraploids are limite dto the southeastern part of the Korehn Peninsula I.he leaf
shape of diploids shows a clinal variation from lanceola ttoe ovate toward south, At the south end of this
peninsula ,lanceoia tleeave ddiploid asnd ovate leave ddiploid asre sometimes sympatric in several pop-
ulations, Ibtraploi dalsso have a lea fshape variation from lanceolat eto ovate even within a population.
The plant swith elliptic-lanceolate or lanceola tleeaves are identifie das A. ageratoides var. agerateides,
Although a taxonomical treatment sti1 1waits for funhe rconsideration, the resemblance between the ovate
leaved plant sin Korea and A. ageratoides var, ovalijioli"s, known as endemic to Japan ,is noted,
Key words: Aster ageratoides, Asteraceae c,ytogeography, Korea, polyploidy
The Aste' rag'eratoides complex, comprised of poly- counted the chromosome numbers fbr the plants
plQid series, is one of the Sino-Japane seelements collected from about 60 populatio nisn Korea ,The
distribu twieddely from southwestern China ,Korea, results are describe idn relation to the geographical
Taiwan to Japan ,and has many intra-spec itfaixac distribut ianodn their morphologicul features.
and close]y related species (Kitamu r1a993, Seejima
& Peng 1998a) ,In the A. ageratoides complex, Materials and methods
one species and ten varieties in China (Lin g&
Chen 1985), one species and one variety in [ItLiwan More than 250 plant swere collected from 62 pop-
(Soeji m&a Peng 1998b), and fiv especies and three ulations in Korea ('Ri b1l, Feig .1) .Among them,
varieties in Japan (It &o Soejim a1995) were record- chromosome numbers were counted fbr a total of
ed, But in Korea, A. ageratoides var. ageratoides 189 plant scollected from 51 population Fso.r cyto-
with elliptic-lanceolate leave shas only been record- logica olbservation, root tip sobtained from the pot-
ed, and there is nQ record of it schromosome num- ted plant swere used. Root tips were stored in iced
ben water for about 24 hours, and fixed with ethanol and
In thi sstudy, we made fiel dobservation and acetic acid (3:1 )Ma.ceratio nand staining were
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98 APG su)15.6
TABLE1.Samplinglocalitiesan dchromosome numbers determined
Lveulity 2n' Pop.No.b
Seoul-siBukhan-san.Debong-gu 18 (4)18(3)lg(1)IS3456789le1112131415i6]8192123242S26?7293058S9602t720222831383940414243444546474S491333S36so5t3256573452S354S5376162
ayoeng-gi-do Gwangneung.Sohol-myeon,Pochcon-gun
Heungivong-sa,Idong-myeun.Gapyeang-gun
Gwangdeog,Sanae-myeon,Hwacheon-gun (2)18
Gwangsi-rycong,Yanggu-eup,Yanggu-gun
Saetee,Ildeng-myeon,Pocheon-gun (2)L8
Pung-san,Hwacheon-eup.Hwacheon-gun C3)18
Seoween,Sinbuk-eup,Chuncheon-si (6)lg
Cheoljung,Hwachen-mycon,Hengcheo]egun <2)l8
Ooga,Yangyang-eup,Yangyang-gun (3)18{11)IS
Yongpyeellg,Wangsan-myeon,Gangncung-si
Wondeng,Buk-myeon,Yeongwul-gun (4)IS(3)IS
Daehwa,Daehwa-myeon,Pyongchang-gun
Ro-dong,\ongpyeung-myeon,Pyongchang-gun (4)lg{3)18
Jangseong-dong,Taebaek-si
Sabuk,Sabuk-eup,Jeongscon-gun (5)18(])lg
Ssarijae,Gohan-eap,Jeongseon-gun
Chungcheongbuk-do Danyang,Danyang-eup,Danyang-gun (3)18
Shindang,Hansu-myeon,Jecheon-si
HHllam,Neeun-myeon.Chungiu-si (3)l8
GaTmok,Boeun-up,Boeun-gun (5)18
Piban,Huebuk-myeon,Boeun-gun (2)
Hagae,Tanhu-myeon,Boeun-gull
Ycon-gok,jineheon-eup,Jincheon-gun IS (2)18+1-2B
Chungcheo]gnam-do Taejo-an,Oesaii-myeon,Buyeo-gun (5)
Magek-su,Sagok-myeon,Gongiu-gun
Gueshim-sa,Unsan-myeon, Seosan-gun t8+O-IB{5}
Gyeengsangbuk-de Gasan-sanscong,Gasan-myeon,Chilgok-gun 18{4)tS
Hyeon-dong,Socheon-myeon, Bonghwu-gun
Bu-Seok,Buseok-myeon,Ycongtn-si (4)18
Jungryeeng.Punggi-eup,Ycungiu-si (S}36
Seowen, Naeseo-myeon. Sangiu-si
Seokgul-am,Yangbuk-niyeon,Gyeongiu-si (4)
Dosan,Dosun-myeon, Andong-si
Sacheon.Byeonggok-myeon,Yeongdeok-gun
Bengieong-sa, Seohu-rnyeon, A"deng-si 18 (2)
Jiri,Budong-rnyeon,Cheongsong-gun
Dcog-s,ye ,Hyeensee-myeon, Cheongsong-gun
Naeryong,Budong-m},eon,Cheongsong-gun 18(1)lg(1)
Ok-san,Dalsan-tnycon,Yeongdeok-gun
Seongnyu-gul,Geunnam-myeon,UIjin-glln
Bogyeong-su, So]igna-myeo Pno,hang-si 18 (2)
Scongu,Oajeong-myeon,Uljin-gun 18(3)
Naechi].Sannac-myeon,Gyeongiu-si 36 (3)
Hwayang,Chev]gdo-eup,Cheongdo-gun 36(t)
Daegu-si Palgong-san,Dohak-dong.Hak-dong IS (9)
Gycongsangnam-do Haein-sa, Gaya-mycon ,Gcochang-gun 18 (7)
Gaji-san,Sangbuk-myeen.MiJyang-gun 36 (5)
Nacwen-sa,Habuk-myeon,Yangsan-gun 36 C4)
Gosa,Jinjeon-rnyeon,Masan-si l8 (2)
Pyeonggu,Ssangbaek-myeen,Hapcheon-gun l8 (5}
Jeollabuk-do Jiri-san,Sannae,Namwon-gun 18 {2)
Gcumsan-sa,Geumgu-myeon,Gimje-gun IS+IB([)
Anshim-sa,Daedung-san.Bcolgok-myeon,Waaju-guii tg (4)
Jeollanam-do Muncheok,Muneheok-myeon,Gwangyang-gun ISU)
Ipam-san,Sipjcong-dong,Jeongeup-si IS(6)
Yuma, Mohu-san, Nam-tllyeon ,Hwasun-gun 18 (3)
Wolchul-san,Ycongam-eup,Yeongam-gun IS (6)T8+O-IB
BoTim-sa,Yuchi-myeon.Ga]gheung-gun (5)
Busan-siChGeejuui-pdjoeong-sanseong,Geunijeong-gu 36 (5)
NearthchotcLPukcheju-gun IS(3)
'"' South part ofMt. Hallasan, Sogvvipo-si 18(3)
/ NumbeTs in parenthEs s-heewss the numbeT of plan{ esxarnined. b /Pepulationnumbers are corresponding to Fig. 2.
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April 2UOS SOEJIMA et at.: Cytogeography ofAster ageratoides 99
P
65o
s4
ci?
elIS124.{
FiG. t, Samplin glocaliti aensd !ea fshape variation, Large open circle = diploid c.losed triangle = tetraploid ,sma]1 c]osed circle =
unknown. Scale bar = 5 crn.
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1OO APG V)1.56
tetrap]oi dhsad secondary constriction on thei rshort
arms. And the shortest chromosomes of both
diploid asnd tetraploids were longe rthan the half-
length of the longes otnes. One or two additional B-
chromosomes were observed in some plant osf four
([[hb1l)e.B-chromosomes
populations were small-
er than any other A-chromosomes. These kary-
ological feature sare coincident with the previous
studies of the most members ofAster ageratoides
complex in Japan (Huziwa r1a957, Matsuda &
Suyama 1980,Matsuda& Shinohara1985,Irifune
1990,Soejimal992,1993),China(Soejima
et al,
FiG. 2. Measurernent sef leave sand peduncle sa., lcaf blade 1999) and Thiwan (Soeji m&a Peng 1998).
lefe cn1g,t c 2hb,.., .l,.e, acf nb lias dues ewdi)d.th. c. peduncle lengt h(th aeverage Diploids and tetrapleids were not sympatric in
any population sexamined. The distributio nof
diploid and tetraploid populatio niss shown in Fig.
conducted simultaneously in liqui dof 1% aceto- I, It is obvious that diploi dpopulatio narse distrib-
orcein (45% acetic acid) and IN HCI (9: 1f)or about uted widely and very common in Korea .On the
24 hours befbre squash. The aerial parts of the other hand ,tetraploid populatio narse restricted to a
plant sgained in the fiel dwere prepare das voucher small region of southeastern part,
specimens for morphological analyses and are
deposite idn Kyo. Morphotogica lfleatt tirn erselation te the ploidyJ
For morpho]ogical analyses, the length and ZevelThe
the width of a leaf and the lengt hof peduncles importan ctharacters to distingui tsahxa within
were measured (Fi g2.), A leaf without serious the Aster ageratoides complex are such as leaf
damages was chosen from the larges lteaves of a shape, hair densit yon stems and leaves h,ead size,
specimen.-As the peduncle length, the average and color of Iigule sI,n Korea, leaf shape varies
length of two to seven peduncle sin a cluster was rather widely from elliptic-lanceolate to ovate. But
used. A cluster was determined as a group of heads it seems that there are not marked variation in hair
arranged above the uppermost bracteo lwehich had density ,head size, and color of ligule sT.he stems
distin cmitdvein. For measurement, a cluster was and the lower surface of leave sare nearly glabrous
chosen from the clusters closest to the main axis. or sparsely pubescent with short hairs ,and the
upper surface of the leaves is usually strigullose
Results and Discussion with sparse rigid short hairs T.he diamete rof the
heads including the ligule sis about 1,5-2 cm at
Ch romosome .feat uies and distribution anthesis. The ligule asre usually white but some-
Chromosome numbers were detertnine dfor the 189 times tinged with pale purple on the outside espe-
plant fsrom 51 populatio nAsm.ong them, 167 plants cially befor eanthesis. Among other characters, the
from 45 populatio nwesre diploid s2,n = 18, and 22 lengt hof peduncle sof the heads shows a relatively
plant sfrom 6 populatio nwsere tetraploids, 2n = wide variation range.
36, based on x = 9. The longes ttwo chromosomes The lea fshape varies continuously from ellip-
of diploid asnd the longes tfour chromosomes of tic-lanceola tteo ovate in both diploid sand tetra-
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25
20
vee.
15qg.dse
lo8
5
o
1.5e-1.7 51.75-2.0 e2.00-2.2 52.25-2.5 e2.50-2.7 52.75-3.0 03.00-3.253.25-3.5e3.50-3.753.75-4.0e4,OO-425
lengthfwidth
FtG. 3. Frequency distributi oofn lea fiengthtwidt lhength,
20
15veE-ngO
10iggZs
o
O.O-2. 5 2.5-5. 0 5.0-7. 5 7.5-10. e10.0-12 .512.5-15. 015.0-17 .517.5-20.020,O-22 .225.5-25. 025,O-27,5
length (mm)
of peduncle
FiG .4, Frequcncy distributi oofn length ofpeduncle
ploids .There is not statistically significant difference diploids and tetraploids, tetraploids tend to have
between diploid sand tetraploids, but tetraploids shorter peduncle tshan diploi d(sFi g4). .It seems that
tend to have oyate leaves in comparison with there is a weak correlation between leaf shape and
diploids (Fig 3,). Long peduncle smake the inflo- peduncle length p;lants with ovate leaves tend to
rescence loose ,while short ones make them dense, have short peduncles (Fi g5.). However, the corre-
Although there is no significant differenc ebetween latio nbetween the leaf shape and the peduncle
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4.5
4,O
3.5
ep-))-uagT--
3,O
2,5
2.0
1.5
1.0O,O
5,O 10.0 15,O 20.0 25.0 30.0
length (mm)
of peduncle
FIG. 5 Scatte rdiagram showing relationship between leaf length/le awifdth and peduncre length.
length is not statisticall ysignificant, In a total, the ed int otwo morpholegically distinguisha gbrloeups,
ranges of merphological variatien of tetraploids Considering the parapatri coccurrence of
are include din those Qf diploids. tetraploids with diploid s(Fi g1.) and their ranges of
morphological variation, which do net exceed these
Geographic cline of diploid s(Fi g3,-5) ,the origin of tetraploids could
Diploids are distributed w{de]y and abundant haye been related to diploi disn the southern part of
throughout Korea, but their variation of the leaf Korea. The rather wide morphological variation
shape shows a geographic cline as oyate leaves shown among tetraploids could be attributed mul-
occur in the southern and seutheastem part of Korea tiple polyploidizati oinn situ, or some genetic
(Fi g1,) .As is shown in Fig .1, only lanceolate-exchange through backcross such as unreduced
leaved diploids appear in the northern part ,but gametophyte of diploi adnd reduced gametophyt eof
both lanceolat eleave dand ovate leaved diploid satre tetraploid.
sympatric in some southern population ss,uch as
Populatio n4s6, 53, and 54 (Fig 1.) .The leaf shape 7inxonomic considerations
varies from elliptic-lanceolate or lanceola tleeaves In Korea, only Aster ageratoides var. ageratoides
to ovate leaves continuously within a population.has been reported in the A. ageratoides complex.
Ibtraploid stend to have ovate leaves (Fi g3.) ,but Followin gIto & Soejima (1995 )t,he plant swith
they also have rather wide variation range in the Ieaf lanceola ttoe elliptic lanceola tleeave sare identified
shape from el]iptic-lanceolate to ovate even within as var. ageratoides. But the plant wsith ovate leaves
a population (37 4,8 in Fig, 1) .The leaf shape vari- found in this research qujt eresemble to A, agera-
ation is continuous and they cannot be clearly divid- toides var. ovalijblius, which is considered to be
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endemic to Japan. The variety ovaldelius occurs tion 61 have lanceolat eleaves and are identified
on the Japan Sea side, mainly in the Hokuriku dis- as var. ageratoides, Although the size difference
tric tincludin gYamagata and Niigat aPrefectures is noticeal )le ,here we consider the plant sof both the
as it scenter, however ,it interrnitte onctculr$y west- two popuiatio nosf Cheju Islan das Aster agera-
ward along the Japan Sea side to Chugoku districttoide svar. ageratoides.
and Tsushima Islands ,The population sare separat-
ed by certain geographic distanc eisn west part of its VVe are gratef utlo Dr, H, Nagarnasu fbr his taxonomical
distribut iaorena, which could suggest that the var. suggestion and help in drawing picture sO.ur special
oval(fblius of Japan has differe onrtigin frem that of tMhsa. nKk isamre dYubeu tnog D-r.o kCfhoor i tKhyeuirn cgo,r dMisa,l Saosns iOstka-nnceeo imn, aonudr
the ovate leaved ones of southern Korea, fiel wdorks, This study was supperted by a Grant-in-Aid
The morphological variation ranges of va: of Scientif Riecsearch from the Japanes eSociet yfor the
ageiutoides and var. ovatijiolit{s are continuous in promotion of Scienc e(No ,07041126 to M. Ito and No.
Japan as is the same to the situation of both dip]oids 15310161 Lo H. Yamaguchi),
and tetraploids in the southern part of Korea ,That is
References
the reason why var, ovalijbtius is recognized at
variety rank. This study revealed the cytogeo-
graphica dlistributi oannd the morphological varia- Huziwara ,YL 1957. Karyotype analysis in some gener aof
tion in Korea, and showed the possibil iotfy exis- Ctooimdepso sgirtoauep , .IAImI e,Trh,e J k ,Baorty,o 4t4y:p 7eo8f3 t-h7e 9A0s.ter agera-
tence of dip!ei dand tetraploid var, ovalijblius in Irifun eK,, 1990, Karyornorphologi cstauldy on speciation
Korean Peninsula .On the other hand, only tetra- of the Aster ageratoides subsp. amplexijbtius com-
ploid swere known in var, ovalijblius of Japan to plex in Japan ,J ,Sci .Hireshima Univ., Ser .B, Div. 2,
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Ito,M, & A.Soejima.1995,AstenIniIwatsuki,K.,T.
tionshi pbetween var, ovatefolius in Japan and the
ovate-leaved diploid sand tetraploids in Korea, tax- Yofa mJaazpaakn i3, bD: .5 9E-. 7B3e .uKfefdbanrsd h&a ,H ,[ [Obhkbyao.. (eds. F)l,ora
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Ling,Yl& Y.-L.Chen.1985.Astenin:Ling,Y.,Y.-L.
latio n62, fbund in Che,iu )sland ,are very small in
the height T,hey have lanceola tleeaves and quite C1h3e3n- 2&5 4C.(. SihnCihhi .n(eesdes ).Fl) ,Repub. Popul .Sinica e7.4i
similar to the var. ageJutoictes, The size of the heads Matsuda, T, & K, Suyama. 1980. Pelyploids and distrib-
is not so much smaller than others, but the plant ution ofAster agerutoides subsp. Ieiqphyil" sgroup in
height is between 10-30cm at anthesis. This popu- the Kanto distric atnd surfounding area. I .Sci .Rept.
latio nappeared along a small stream of the moun- Ybkohama Natl. Univ., Sec. 2, 27: 7-18,
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the
environmental condition could cause miniatur- Ybkohama Natl.Univ.,Sec.2,32i11-26,
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of the plant sat anthesis is 80-100cm, which is sim- wirh special reference to ploidy level ,Bot. Mag.
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104 APG Nbl.56
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Received September 18,2004;accepted lanuaryI3,2005
'
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