A Study on the Morphological and Cytological Features

The Japanese Society for Plant Systematics

The JapaneseSociety forPlant Systematics

ISSNooOl-6799

Acta

^{PhytotaxGeobot.44}

(1):

^35-51

(1993)

A Study on the Morphological and Cytological Features

^of

Aster leiophyUus Complex (Compositae) ^{in Kyushu, Japan}

AKIKO

^SOEJIMA

U}iiversity

of

^{Osaka Ptqfecturq CoUege}

of

^{lhtegratedArtsand}

Sciences,Gakuen-cho, Sakai.Osaka⁵⁹³

Abstract The AsterleiQphylluscomplex istaxonomically complicated and includesmany related

taxarepresenting polyploid^series. Ithasbeenreported thattwotaxa^ofthis group^are distributedin Kyushu, However,there^{are many} herbarium specimens collected inKyushu^{which cannot} be identifiedtoeitherof them. Inthisstudy, morphological variations were examined inrelation to ploidylevel,and taxonomic consideration ^was given, As a result, threespecies and one variety are

recognized inKyushu: A. IeiophyUus^{var. teiophyUus}

(2x,

^6x),A. semiamptexicautis (2x),A.^ieio- plryllus^var. intermedius(4x)^{var.nov. and} A. satsumensis

(2x)

^{sp, nev, The} distributionrange of

diploidA.teioplp,llus^var. Ieiophyllusislimitedtoasmall areaof Fukuoka,Nagasaki^{and Miyazaki,}

while thehexaploids^{occur widely} inand around volcanic regions suchas Mts,^{Aso-Kujiu,Unzen and} Kirishima.A. ieic\u,lliLs^var. intermediusisdistributedinthenorth-eastern part,A.semiamptexi- caulis inOitaand the northern part^ofMiyazaki,^and A. satsumensis mostly inthesouthern part of

Kyushu.

(Received

^{April10,1993;Accepted May 26,1993)}

Key words: AsterieiqphyUusmorphological ^variation, polyploid^{complex, taxonomy.}

^{The Aster}ageratoides sensu

Kitamura (1937,

^{1981)comprises a well developedpolyploid}

complex

distributed

widely inEastAsia. Inthistaxon,two

distinctly different karyetypes

are known. One

karyotype is

^{called as}

L-type,

^{which haslong}chromosomes only, the other

is

LS-type which

has long

and short chromosomes halfand halfwithin a

karyotype.

^The

plants ^with L-type karyotype are morphologically variable, and regarded as ^a taxonomic complex. A polyploid^{series ranging}

from diploid

tononaploid except forheptaploid

in Japan (Huziwara, ^{1953, 1954,}

^{1955, 1956, 1957a, b; Irifune,1990; Irifuneet al.,}

^1985).

Previous

authors called this^{complex variously;}

A.

crgeratoides subsp. amplexijblius complex

(Irifune,

^1990),A. ageratoides subsp.

IeiophyUus

group

(Matsuda & Shinohara,

^1985;

Matsuda & Suyama, 1980), A. IeiQpdyllus

complex

(Soejima,

^{1992). Here, I}

^follow

^my

previouspaper

(Soejima,

^{1992)and use thename}

^A. Ieiophyitus

complex.

The Aster leiopityllus

complex ismorphologically variabie comprising many taxa.

Natural

introgression

^and!or

inter-

^and

intraspecific hybridization between

thetaxa^of the complex are

believed

to

happen

occasionally

(Kitamura,

^{1937,1981). Thus thetaxonomy}

within this complex

is diMcult,

and

interpretations

of the

boundaries

and the

interrelationships

among thetaxaare commonly conflicting.

InKyushu, two taxa of the Aster leiophyltuscomplex, 'Shiro-yomena'

(A.

ageratoides var. adustus,

A.

crgeratoides var.

harae f. teucanthus, A.

ageratoides ssp. IeiQphyUus,A.

teiopIryllus)and 'Inaka-giku'

(A.

ageratoides ssp. amplexijblius,

A.

semiamplexicaulis)

have been known

to

be distributed (Amakawa,

^{1975;Baba ed., 1964;Hara, 1952;Hatusima ed.,}

1986;Hatusima, 1989;Kitamura, 1937,1981;

Ohwi, 1953;

Toyama,

1980; Yamashiro

^et al.

ed.,

1969). But

^{among specimens}

in

the

herbaria,

thereare ^{many specimens collected} in

The Japanese Society for Plant Systematics

The JapaneseSociety for Plant Systematics

36

Acta Phytotax.Geobot. Vol.44

Kyushu

^{which cannot} be identifiedtoeitherof thetwo species nor teother taxa of the A.

teiqphyUus

^complex.

In

_general,the existence of

polyploid

^{series may}

be

one of the

factors

of taxonomical

diMculties.

Although

diploid,

tetraploid and hexaploid are reported in Kyushu

(Irifune,

^{l987,1990;Irifuneetal.,1985),the}relationships

between

taxaand ploidy

levels

were not discussed.The aim of thispaperisto^recognize themorphological variation ofthe

A. leiop1tyllus

compiex

in

connection

with

the ploidyleveland to

discuss

thetaxonomy

of the complex inKyushu.

Materials

^and

Methods

Plants^{were collected}

from

56

populations in

Kyushu

(Table

^{1,Fig.1). The aerial} parts

of the plants^were

kept

as

herbarium

voucher specimens

for

morphological analyses. The subterranean parts^{were cultivated} inpots^at Tokyo Metropolitan Universitytobe used for

cytological investigations.The _{voucher specimens are}

kept in MAK.

^{The three}

1argest

leaveswere chosen from each specirnen and examined for _eight morphological characters

(Fig.

^{2). The methods of} morphological and cytological analyses

followedare those of Soejima

(1992).

^For taxonomical consideration, the specimens inthe herbariaKYO, MAK, TIand TNS

were studied.

Results

I. Cytological

features

1. Ploidylevel

The basicchromosome number of the genusAsterisx=9

(Huziwara,

^{1953,1954,19S5,}

1956,

^1957a,

b). In

the

present

^study,

diploid,

tetraploid,

pentaploid

^and

hexaploid

were observed. Pentaploidswere rather few and occurred inpopulations^with diploidsand/or

hexaploids. The ploidy

level

^observed

in

^each _population

is

^shown

in

Table 1and Fig.1.

In

the

Aster leiopityllus

complex, the

length

of chromosomes

in

a

karyotype is

relatively constant, and ina karyotype,the lengthof the shortest chromosome islongerthan one half

of the lengthof thelongestone. Many observed plantshaveone tofiveB chromosomes

in

a

karyotype

which can

be

easily

distinguished

^{from normal ones bytheirextremely small size.}

The number of B chromosomes isstable withinan individualplantbutvaries even within a

population

^{and seems to}

have

no taxonomic significance. The numbers of B chromosomes were net taken intoconsideration

in

thisstudy.

Two diploids

areobtained

from

a

population

^ofFukuoka Prefecture.Arnong 14plants

from

six

populations

^of

Nagasaki Prefecture,

threeplantsfrom the populations^of NS1 & 2 are diploids,while other eleven plants^of NS3rv6 are all

hexaploids. Six diploids,

one

pentaploid

^{and six}

hexaploids

are

found in

the population ^of KM2, in Kumamoto Prefecture.Diploids,tetraploids and

hexaploids

are

found

ⁱⁿ65plantsfrom 14populations

in Oita Prefecture. Nine diploids

are

found

ⁱⁿ the populations ^of OI15 & 16. The populations

OIIN4,

6･v8

& 10

^consist of tetraploids.Hexaploids occur inOI9, 12-v14.

Three pentaploids

^are

also found in

the population ^of OI14. From 12 populations ^of Miyuzaki Prefecture,44plants^{were examined.}

MZ6"v8

are the

populations

^of hexaploids, and alltheother tenpopulations^{consist of} diploids.InKagoshima Prefecture,

al1

52plants

examined from 12

populations

^{are diploids.}

During ^my fieldtrip

in

SagaPref.,thenorthern part^of

Nagasaki Pref.

and thesouthern

part

^{of Kumamoto Pref., Icould not findthe plantsof thiscomplex except fora population}

The Japanese Society for Plant Systematics

The JapaneseSociety forPlantSystematics

August1993 SOEJIMA: Asterleiophyllus

in Kyushu 37

Table1. Colleetionsitesand ploidylevel Population

ne. Locality Voucher -speclmen Ploidy^level

(No.

^of plants) FOI

F02 NSI NS2 NS3 NS4 NS5 NS6 NS7KM1KM2

OII OI2 OI3 OI4 OI5 OI6 OI7 OI8

OI9OIIOOIIIOI12OI13OI14OI15OI16MZIMZ2MZ3MZ4MZ5MZ6MZ7MZ8MZ9MZIOMZIIMZ12

KSI KS2 KS3 KS4 KS5 KS6 KS7 KS8

KS9KSIOKSIIKS12KS13 ::::::::::

::::::::

::::

t:

::::;:::::::::::::::::::

Chikushi-gun,^{Nakagawa T.,Funyduo} FukuokaC.,Sawara-ku,Hinatapass HiradoC.,^Mt. Shijiki

Oomura C.,Kuroki

Minamitakaki-gun,^{Kunimi T,,Uematsu} Minamitakaki-gun,MizuhoT,,Miyanoji Minamitakaki-gun, Ariake^T.,Oona Minamitakaki-gun, NishiarieT.,Tounosaka Minamitakaki-gun, Obama ^T.,Kitano Aso-gun, Oguni T.,Nishizato Ase-gun, Oguni T.,Kamida

Higashikunisaki-gun,Aki T.,Karami Higashikunisaki-gun,Aki T.,Yurugi Higashikunisaki-gun,KunisakiT,,Inagawa Higashikunisaki-gun,Aki T.,Mt. Futago Bungotakada C,,Hashirimizu pass Bungotakada C,,Tazome

Hayarni-gun,Yamaga T.,Hisashi Usa-gun,Ajimu T.,Higashishiiya Oita-gun,Yufuin T.,Kawakita Kusu-gun, Kokonoe T.,Kabeyu Naoiri-gun,KujiuT,,Kujiu Takeda C.,Ogawa

Takeda C.,Takeda Takeda C.,Takeda OitaC.,Karnihetsugi Usuki C.,Nakausuki Miyazaki C.,Kagarnisu

Higashiusuki-gun,KitagoV,,Yamakariya Higashiusuki-gun,KitagoV,,Motokariya Minaminaka-gun, Nango T.,Yowara Kushima C.,Agebaru

Miyakonojyou C.,Miike

Nishimorokata-gun, TakaharuT.,Hirowara Kobayashi C,,Mizunote

Nishimorokata-gun, NojiriT.,Amagatani Nishimorokata-gun, NojiriT,,Imabyu Higashimorokata-gun, Takaoka T,,Uchiyama Higashimorokata-gun, Aya T.,Minamimata Soo-gun, ShibushiT.

Kanoya C.,Kamitakakuina Kimotsuki-gun,SataT.,Hetsuka Kimotsuki-gun,NejimeT.,Ootakeno Kimotsuki-gun,Nejime T.,Yokobeppu IbusukiC.,Hatakekubo

Kawanabe-gun, ChiTan ^T.,Higashibeppu

SendaiC.,Aoyama

Hioki-gun,HigashiichikiT.,Yuda Hioki-gun, Higashiichiki T.,Takatsuka Satsuma-gun, Hiwaki T.,Tounohara SendaiC.,Nishikata

Idzumi C.,Kamiookawauchi

Soojima90os61 Soojima 90os62 Soojima90a563 Soojima90a564 Soojima9005as Sony'ima9a0566 Soojima 900567 Soojima 900SciP Soojima90057e Soojima 89IO12 Soojima891013 Soojima 891001 Soojima 89IO{12 SoojimaS910CB Soeg'ima891004 Soojima891005 Soojima891006 Soojima891007 Soojima891008 Soojima891e09 Soojima 891010 Soojima 891016 Soojima S91017 Soojima 891018 Soojima8910i9 Soojima^891a20 Soojima891Cl21 Soojima⁸⁹¹¹²² Soojima891123 SeojimaS91124 Soojima 891125 Seojima891126 Soojima891146 Seojima89i147 Soojima89i149 Soojima891150 Soojima 891151 Soojima89J152 Soojima S91153 Soojima89JI3Q Soojima 891I31 Soojima89J132 Soojima891133 Soojima 8Pl134 Soey'ima89JI36 Soojima89J137 Soey'ima891139 Soojima891140 Soey'ima891I42 Soojima 89i143 Soojima 8Pl144 Soq'ima 891145

2x(2)2x(1)2x(2)6x(2)6x(2)6x(4)6x(3)

2x(6),5x(1),6x(6)

4x(2)

4x(11)

4x(7)

4x(7)

4x(4)

4x(4)

4x(5)

6x(1)

4x(5)

6x(3)

6x(3)

5x(3),6x(1)

2x(5)

2x(4)

2x(S)

2x(4)

2x(4)

2x(2)

2x(11)

6x(2)

6x(4)

6x(2)

2x(3)

2x(4)

2x(2)

2x(1)

2x(8)

2x(8)

2x(5)

2x(5)

2x(2)

2x(3)

2x(7)

2x(5)

2x(5)

2x(4)

2x(5)

2x(3) FO:KS:FukuokaPref.,

Kagoshima Pref.NS!

Nagasaki Pref.,KM: KurnarnotoPref., OI:Oita Pref.,MZ:Miyazaki Pref.,

The Japanese Society for Plant Systematics

The JapaneseSociety for PlantSystematics

38

Acta

Phytotax.Geobot. Vol.44

of NSI, inHirado

Isl., Nagasaki.

2. Distributionof diploids,tetraploids and hexaploidsinKyushu

(Fig. ¹⁾

The

occurrence of

diploids is

not uniform inKyushu. Although the

diploid

plants^are

scattered widely throughout the sampling

localities,

most of them are clustered

in

the

southern partof

Kyushu. A few diploids

appear

disjunctly in

some

populations

^ofFukuoka,

Nagasaki, Kumamoto

^and Oita.

AIthough therearemany tetraploid

populations in

thenorth-eastern part^of Kyushu, no

tetraploidsare

found

intheother localitiesof Kyushu.

Fig.

O A e

t

.

OI4

11

Z12

2

1 Collectionsites and distributionof cytotypes. Open eirclesrepresent diploids,solid triangles tetraploids,solid circles hexaploids,a star pentaploid. Smail closed circles represent populations, the ploidy levelsef which arenot known. Forabbreviations, see Table1,

The Japanese Society for Plant Systematics

The JapaneseSociety forPlant Systematics

August 1993 SOEJIMA: Asterleiophptll"sinKyushu 39

? ^l

1

,Z8m

43

g RQ g

^Fig.2 MeasurFments ofleaf. 1:Ll

aeaf

^length),2:L2

(length

^ofbroad part), 3:Wl

aeaf

^{width), 4:W2}

(width

^{at narrowing point),5:W3}

(width

^atthe

^midd!e point between thenarrowing point^{and the}leafbase),6:HD (hair densityoftheabaxial surface), 7:GD

(gland

^{densityof theabaxial surface),}

8:HDD (hairdensityof theadaxial surface).

The ^{occurrence of}

hexaploids

tendsto

be

concentrated

in

and around thethree^volcanic

regions, namely

Unzen (NS3--6),

^Aso-Kujiu

(KM2, ^{OI9, 12-14)}

^and

^{Kirishima (MZ6--8).}

II. Karyotype features

Diploids

^{have nine} pairs^{of normal} chromosomes often with a

few B-chromosomes.

^In

most

diploids,

^{each chromosome of} the

longest

pair

has

asatellite

located

on the short arm, and theothers

do

^not

have

a satellite

(Fig.

^{3A). The}

^diploids

^ofthe populations^{of southern} Kyushu

(MZI-5,

^{KSI, 2,4-13)}

^{have different karyotype. They have}

^{one ortwo additional}

satellites on thechromosomes ofthesecond pair. The additional satellites are sometimes on

the^{end of the} longarm

(Fig.

^{3B) and sometimes on theshort arm.}

Tetraploids have 18

pairs and

hexaploids

^{have 27} pairs^{of normal} chromosomes often with several B-chromosomes. The

karyotypes

of tetraploidsand

hexaploids

seem to be

multiple of the

diploid

with two satellitechromosomes. Among the 36 chromosomes of

tetraploids,each ^of the

four longest

^ones

has

^{a satellite on} theshort arm, and among the

54

chromosomes of

hexaploids,

there are six chromosomes with a satelliteon the^{short arm.}

III. Morphologicalvariations

1. Hair density

Diploids^{show a wide variation range}

for hair density. The hair density

^isrepresented

by HD (hair

^{number cQunted within 4mm2 on the abaxial surface of the}

^leaf).

^{The range of}

HD varies

from O

tomore thanthree-hundreds.

Figure 4

shows the^{variation range of} HD of

The Japanese Society for Plant Systematics

The JapaneseSociety forPlantSystematics

40 Acta Phytotax.Geobot. Vol.44

Fig.3 Somaticmetaphasechremosomes. A: A.semiatmplexicaulis(OIIO,B:A,

^rows indicatechromosomes with a satelliteon thedistalend of theshort arm.

^{chromosome with a satelliteon the}distalend of the longarm.

satsumensis(KS8), Ar- An arrow hcad indicates

fi

350 300 250 200 150 100

50

o

N-a ^{T- c.) so u')} co ^a N- ea N lr) rs NMNeo ocl u'} KD

^2ENEEEE:5g ge ^{wa wa} ee ^{65 6- E 6- g}

Fig.4 ThevariationefHD. Aboxshowstherangebetweenaverageplus-minusstandarddeviation. Each ^{end ofthe} barsmean minimum and maximurn yalue inapopulationrespectively. Allthe individualsof

^the populationsNS3 and NS6 have hairlessleaves;HD==O, Forabbreviations and themorphological types^{of each} population,^{$eeTables1& 2.}

Fig.SHaiTs on theabaxial surface of theleaf.A:OI15

(D-3),

^{B: OI8}(T-1).x150,^Bar==20Qum.

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The JapaneseSociety for Plant Systematics

August ^{1993 SOEJiMA,} AsterteiophytlusinKyushu 41

Tab]e 2. Themorpliological ty})esot'each population D-1D-2D-3T-lH"iF02, NSI, MZ9.12

NS2,KM2(2x),MZI-5, OJIS, l6

OIi.8,10

NS3-6,KM2(6x), OI9,

KSI.l3

12-14,MZ6-8

some populations.

Although

the^variation seems to

be

centinuous from lowerto higher,itis possible^todividethe

diploids into

three subgroups

based

^on

both

the

hair density

^and

hair length.

The

length

^of the

hair

^varies that the

longest hair is

^as threeto

four

timesas longas the shortest ones

(Fig,

^{5). The shoTtest ones}

(Fig.

^{5A) censist of one er two cells, while the}

Songest

enes

(Fig. ^SB)

^{of threeto}

^four

^{cells. 'Iihere isatendency that the planthavingdense}

hairs has long hairs

and

intermingled

^{short ones, and the densjtyof}

long hairs is higher

than thatof short hairs.The plant^of

low

hair

density has

^short

hairs

^only, The gland

density

^en

the abaxial surface of the

leaves is

alrikost

ifi

_proportion^to thehairdensity. But thevariation range of the gland

density is

narreweT than that of

hair deRsity, Therefore,

the

hair density is

used toclassifythiscomplex inte_subgroups. ^'E'he morphologScally recegnized subgroups

distinguishedrnainly

based

^on

hair density

^are tentativelynamcd as types

D-1, D-2

^and

D-3.

'Themorpholegieal featuresof each type are

deseribed

^helow

(Table

^2).

Type D-1: HDi=O--･30, hairsshort, The

dipaoids

of this type scarccly

have hairs

nor

glands

^{on the abaxial surface of the leaves. Stem isa2se glabreus. Thc diploidsof the}

popaxiationsF02, NSI, MZ9-l2

belong

t.othistype,

Type D-2:

^{HD =:SO-- 1SO,hairsshort and numerous witha fewlonghairsen the abaxial}

Fig.6 The abaxial surface ofthe leaves.A: OT12

(H-D,

^B:OIS

<T-l),

^C:MZ5 (D-2),^D:OII5 (D-3).^x5e,

Bar^=' lmm.

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42

Acta Phytotax.Geobot. Vol.44

surface of the leaves

(Fig. 6C).

There are

densely

suppressed short

hairs

on thestem.

All

the diploids_obtained from

Kagoshima

and

Miyazaki

prefecturesexcept

for

enes

from

the populationsMZ6-12belong _to thistype. The

diploids

^of thepopulationsNS2 _and

KM2

are also

included in

thistype.

Type D-3:

HD=i150N300, abaxial surface of the

leaves

and stem

densely

_pilose^with

long

hairs

and

intermingled

^short ones

(Fig. ^{6D). There}

^{are a}

^lot

^of glands ^on theabaxial surface of the

leaves. The diploids

^of thepopulations

OI15 & 16 belong

to thistype.

Compared

with thediploidplants,thetetraploidsand

hexaploids

^show a ^rather narrow variation range of

hair density,

and each of them cannot

be divided into

^any morphological subgroups.

The

tetraploidsand hexaploidsare tentativelynamed as T-1 and H-1 and the

morphological

features

^are

described below.

Type T-1:HD=::(5"-)10--50(-s-90),most

hairs

^{short, with} exceptional occurrence of

long

hairs (Fig. ^{6B). The hair density}

^ofthis typevaries widely

from

^ca.

10to

^near 100inthe

value of HD. Allthetetraploidsobtained

in

thisstudy belong tothistype.

Type H-1: HD=Otv5,

^{rarely more} than

20, hairs

short. The plant body isalmost glabrous,^and the

hairs, if

_present,^{are short}

(Fig. ^{6A). They have}

^scarcely glands. Allthe

hexaploids

obtained

in

thisstudy

belong

tothistype.

A few

pentaploids^{are obtained}

from

two populations,

OI14

^and KM2. The hair density_of them

is low

;

HD

^==O--20.

2.

Leafshape

N?E

6

5

4

3

2

11

2 Fig.7 Scatterdiagrams of leafshapes,

Fig. 2.

3

4

MfWl

Each plot^{represents an} individualplant.

5 For abbreviations, see

The Japanese Society for Plant Systematics

The JapaneseSociety for Plant Systematics

August 1993 SOEJIMA:

Aster leiophyltus

^inKyushu

43

^{Figure7shows the variation}

^{in broad}

^{partof a}

^leaf

^{of the}

^Aster

^{leiophyUuscomplex in}

Kyushu.

Almost

^alltheplants^examined

have

oblong-lanceolate or

lanceolate leaves;

^the

ratios L2fWl ^=2.0 --4.0,

WllW2=1.0

^{--4.0. The} graph

is

plotted forthe plants^{of which} the ploidy

levels

are

known.

^{Itseems thatthe}

diploid

plantstendto

have

more elongated, and wider

based

^{leavesthan those of tetraploidandlor}

hexaploid.

Discussion

1. Taxonomic treatmentsof theAster

leiQphyllus

complex

in

Kyushu

Five

^{subgroups are recognized within theAster}

leiophyllus

complex

in

^{Kyushu based on}

the

ploidy

^leveland morphological

features.

^{Three of the}

five

^{subgroups are}

diploid,

one ^of the others

is

tetraploid and the^another

hexaploid. Gene fiow between

^the

ploidy levels

must

be greatly

limited.

^{However, as} pointed^out

by

^Tateoka

(1975),

gene

flow

may ^not

be

^of

.

crucial consequence

for

aspeciesto rnaintain

its

unity, thus

different

ploidy

levels

^can coexist within a taxonomic ^unity.

In

conclusion, Irecognize threespecies and ^{one variety}

in

Kyushu. The taxonomical treatment^and

its

morphological

features

^are

described below.

Aster leiophyllus

^Fr.et

Sav.

^var.

leiophytlus;

^{D-1 and}

H-1

^{The two subgroups,}

D-1

and H-1 resemble each other

having

^a glabrous ^{or nearly} glabrous

leaf (Fig. ⁴⁾

^{and stem, and scarcely}

^{have glands}

^{on the abaxial surface of the}

^leaf.

The hairs,

if

extant, are short, one or two ^cells

long. It is

^noted thatthe

diploids

^{of the}

populations NSI and MZ9-12, and the

hexaploids

^of

OI14

are ^a

little

more

hairy (HD= ^15--30)

^{than typical plantsof var.}

IeiQpbyUus.

^Aster leiophyllusFr.et Sav.var.

intermedius

^{Soejima var. nov.;}

T-1

^{The subgroup}

T-1 has

an intermediatevariation range of

HD between

^Aster

leiophyltus

var.

Ieiopdyllus

and ^Le4.semiamplexicaulis.

Concerning

^the

hair density, it

^resembles

A.

satsumensis

(D-2),

^{but A.} satsumensis does not seem to

be

the

direct

progenitor^of T-1

because

of theirkaryotype features.Allthetetraploids^observed

in

this study

have four

satellite chromosomes. They should have

been derived from diploids

with two satellite chromosomes.

On

the^other hand, most plants^ofA. satsumensis have three or

four

^satellite

chromosomes inthe

diploid karyotype.

^{Therefore,T-1should}

be

treatedasan independent taxon fromA. satsumensis. The hairsofA. ieiqpdyllusvar.

leiQphyllus

^{and T-1are mostly}

short ^and those of A. semiamplexicautis are ^mostly long. These cytologieal and morphological

features

^{show closer} resemblance of T-1 to A.

Ieiqplryllus

var.

leiop]ryllus

than to

A.

satsumensis andlorA. semiatuplexicaulis.

I

regard thissubgroup ^{as a variety of}

A. leiophyllus

and propose the^{new name}

Aster leiophyllus

^var. intermedius.

Aster satsumensis Soejima

sp.

^{nov.; D-2}

^The plants^of thesubgroup

D-2 have

^{many long}

hairs

^and _glands

but

^{not so many as the}

plants ^of

D-3.

^{The variation range of HD of thissubgroup seems to be continuous}

from lower

to

higher,

which

is intermediate

^{between those of D-1 and}

D-3 (Fig.

^{4). D-2 is}

distributed

^{mainly inthe southern} part^of Kyushu and haswider distributionarea thanD-1

and/or D-3

in

Kyushu.

Most

plantsof thistaxon have three or

four

satellitechromesornes

(Fig. ^3B).

^{Here I}propose ^{a new species}

Aster

satsumensis Soejima sp.^nov. forthistaxon.

Allthe

diploids

^ofA. satsumensis

in Miyazaki

^and

Kagoshima

prefectures

have

threeor

four

satellite chromosomes. The diploidsof NS2 and

KM2,

^which are morphologically diMcult to

be distinguished from D-2

and

be included

^inthistaxon,

have

two satellite chromosomes.

Irifune (1990)

investigatedthe

karyotype

of theAster ieiophylluscomplex inKyushu. He observed seventy-nine

diploids

^{havingthree or foursatellite} chromosomes and eightdiploids

with two satellitechromosomes

in Miyazaki

and Kagoshima. According to him, all the

The Japanese Society for Plant Systematics

The 　Japanese 　Sooiety 　for 　Plant 　Systematios

44 Acta　Phytotax．　Geobot ．

Vol

．44

diploid

　plants^ofthe　other　

localities

^except

for

^one

dip 且

oid

in

thepopulation^ofTomari _， Notsu　T ．，

Oita

　

Pref

．，

have

　only 　two 　satellitechromosQmes ．

　　　．4 ．semiamplexicaulis 　

Makino _；

D −3

　　　The

^{subgroup 　D −}

3

　is　characterized　

by

　extremely 　dense_，　long　

hairs

^and_pienty^of_glands

on the　abaXial　surface of the

且

eaf．

　This

^{subgroup 　is　identified　to ／}

t

．se〃ziamplexicautis

Makino _（

Syn

，：・

4

．　ageratoides 　ssp ．　amplextfolius ）．

2

．

Notes

　on 　the　

distribution

　pattern^of　the ．4ster　leiophyllus　complex 　

in

Kyushu

　　　 Usually

，the　p

^亘

^ants　of／Aster　

leio _ρ h _ア〃

us　complex 　appear 　commonly 　ontheroad sideor

forest

　margin ．　

However

_，　

in

　Saga　Pref．_，　the　northem 　part^ofNagasakiPref．and the

As 　　

A． 　　 A． 　　 A

　　2x^：★，^{un   Own ：}

☆

Fig．8



Distribution　of　the／Aster　leioph_ンllus　complex 　in　Kyushu ，

　　

bariumisunknown ，

The　ploidylevel　of　thespecimens ofher一

The Japanese Society for Plant Systematics

The JapaneseSociety forPlant Systematics

August 1993

SOEJIMA;

^AsterteiQphyilus

in

^Kyushu

45

southern part^of Kumamoto

Pref.,

^{I could not}

find

them except

for

a population;

NSI.

Although 'Shiro-yomena'

(A.

ageratoides ssp.

IeiophyUus) is

^reported to

be

^{common inSaga}

Pref.

(Baba,

^{1964),theoccurrence of the complex inthisprefecturemay}

^be

^{rarer than}

ⁱⁿ

^the

other prefecturesinKyushu. My investigation

is insuMcient

^{inFukuoka, northern} part of

Miyazaki

and the ^northern part ^of

Kumamoto

prefectures.For these places,

herbarium

Fig.9 Type specimen ofAster leiophyUusFr,et Sav, var. intermedittsSoejima

The Japanese Society for Plant Systematics

The JapaneseSociety for Plant Systematics

46 Acta Phytotax.Geobot. Vol.44

.specimens

are surveyed

(Fig. ^8).

^{The occurrence of diploidAster} leiopltyUusvar.

teiophyUus is

^rare in

Kyushu. In

this

study, the

diploids

obtained

from

three disjunctlocalities;western

Fukuoka (F02),

^Hirado

Is.,

Nagasaki

(NSI)

^{and southern}

^Miyazaki (MZ9-l2).

^{The hexaploidsoccur commonly}

ⁱⁿ

thevolcanic regions, such as

Unzen, Aso-Kujiu

and

Kirishima.

According to theherbarium

specimens,

A. IeiopiryUus

^var.

leiopityllus

a]so occur

in Kumamoto (Fig. ^8).

Aster

leiophyltus

^var.

inter:medi"s

^{occur commonly}

in

thenorthern part ^of

Oita.

Inthe herbaria,thereare specimens of ^var.

ieiop1tyllus

^{and var.}

intermedius

^collected inFukuoka, Oita_and thenorthern

part

^of

Miyazaki (Fig.

^8).

Astersemiamplt:xicaulis

is

rare

in Kyushu. In

thisstudy, theoccurrence of thistaxon in Kyushu

is

restrictedtothesmal1 area

in

Oita. Only threeherbarium specimens are

identified

to A. semiamplexicaulis.

Two

of them were collected

in Oita,

^and the another

in

the northern

part

^{of Miyazaki}

(Fig.

^{8). About the}

distribution

^of thistaxon, see thenote of the

description

of

A.

satsumensis.

Astersatsumensis is_common

in

thesouthern

Kyushu;

southern part^of Miyazaki and

Kagoshima. A few

_plants^of thistaxon are alsocollected

from

thenorthern part^of

Kyushu,

Nagasaki and

Kumamoto.

3.

The taxonomical position^of thepentaploids^of

OI14 (Oita ^Pref.)

^and

^KM2 (Kumamoto

Pref.)

Three pentaploids and a

hexaploid

are obtained

in

the population OI14. The pentaploids ^resemble tothesympatric

hexaploid.

Both have oblong-lanceolate or

lanceolate leaves

with a few short

hairs

on theabaxial surface.

The hexaploids in

the other region of

Kyushu,

such as populations NS3-6, OI12, l3,KM2 and MZ6-8,

have

glabrous

leaves;

HD=O-5(-10).

The hexaploids

and pentaploids^of

OI14 have

more

hairs;

HD=15-20

(Fig.

4).

Here,

both

the pentaploidsand the

hexaploid

are

identified

to

A. leiqphyllus

var.

Ieiop1tyllus,

tentatively.

However, it is

_possiblethat tetraploid

plants

^ofA.

teiophyllusvar.

intermedius which

also

occur near the population

OI14,

^related to the origin of these pentaploids^andlor

hexaploid.

In

the population

KM2,

a pentaploid ^{appears sympatric with sorne} diploidsand

hexaploids. The diploids

of thispopulation ^are

identified

toA. satsumensis and the

hexaploids

to A.

leiop]ij,llus

^var.

leiopdyllus. The

morphological appearance of the

pentaploid

^{resembles thatof the hexaploidsand isidentifiedto var.} IeiQpJtyllus.

In ^{summary, three species and one variety of} the

Aster ieiop1tyllus

^{complex are}

distributed in

Kyushu :A.

leiophyUus

var. Ieiophytltdsconsisting of diploidand hexaploid, A. laiopdyllusvar.

inter:medius (tetraploid), ^A.

semiamplexicaulis

(diploid)

^{and A.}

satsumensis

(diploid).

Descriptions for

new

taxa

1. Aster

leiophyllusFr.et Sav.var. intermediusSoejima,var. nov. Fig.9.

Aster ageratoides subsp.

IeiQphyUus

x subsp, ovatus

Kitam. in J. Jpn. Bot. 12: 651 (1936),

pro parte.

Aster

crgeratoides

Turcz.

subsp. amplexijblius

(Sieb.

^et

Zucc.) Kitamura,

pro parte.^excl. typ.

Folia ^caulina oblongo-lanceolata,

basi

cuneata, sessilia, paginisadaxialibus sparse

pilosa,pilisplerumque

brevibus

cum aliquot pilis

longis.

Type:

Kyushu: Oita Pref., Usa-gun, Ajimu T., Higashi-shliya, Oct. 31, 1989, Soojima

89IO08

(MAK).

This_variety

is distinguished from

var.

Ieiophyllus by hair density

on the

leaf beneath.

The Japanese Society for Plant Systematics

The JapaneseSociety for Plant Systematics

August 1993

SOEJIMA:

^Asterieiqpltyllus

in

Kyushu

47

In

^contrast tothe glabrous

leaf

of var.

IeiophyUus,

^var. intermedius

has

much short hairon the

leaf beneath.

Perennialsufuutescent rhizomatous herbs. Stems erect ca. 50cm tall,denselypilose.

Leaves

scarcely pubescent^{on abaxial side of}

leaf,

^{mostly short hairswith a}

few long

^ones.

Fig. 10 TypespecimenofAstersatsumensis Soeiima