Table Of Content植物研究雑誌
J. Jpn. Bot.
80:271-277(2005)
A Cytotaxonomic Study of Five Species of Impatiens
(Balsaminaceae) in Jav a and Borneo，Ma lesia
Hiroshi IKEDA ，aShinobu AKIYAMAb，H irokazu TSUKAYA\
Maryati MOHAMEDd and Dedy DARNAEDle
"Department of Biosphere-Geosphere System Science，Fa culty of Informatics，
Okayama University of Science，Ri dai-cho 1-1，Ok ayama，70 0-0005 JAPAN;
E-mail: [email protected]
bDepartment of Botany，Na tional Science Museum，To kyo，
Amakubo 4-1-1，Ts ukuba，Iba raki，30 5-0005 JAPAN;
CNational Institute for Basic Biology，O kazaki Center for Integrative Bioscience，
Nishigonaka 38，My odaiji-cho，Ok azaki，44 4-8585 JAPAN;
The Graduate University for Advanced Studies，Sh onan Village，Ha yama，Ka nagawa，24 0-0193 JAPAN;
Graduate School of Science，Ky oto University，Ky oto，60 6-8502 JAPAN;
dlnstitute for Tropical Biology and Conservation，Un iversiti Malaysia Sabah，
Locked Bag 2073，88 999 Kota Kinabalu，Sa bah，M ALAY SIA;
CBogor Botanical Garden，J .I Ir. H. Juanda No. 13，Bo gor，16 122 INDONESIA
(Received on March 5，2 005)
Somatic chromosome numbers of five species belonging to the genus Impatiens
(Balsaminaceae) collected from Java and Bomeo，M alesia are reported. Impatiens
chonoceras Hassk. was found to be 2n = 14，1. javensis Steud.，1. platypetala Lind ，.land
1. radicans Zoll. were 2n = 16，a nd 1. kinabaluensis S. Akiyama & H. Ohba was 2n =
12. Chromosome numbers of 1. chonoceras，1. javensis，1. radicans，an d 1. kinabaluensis
were examined for the first time. Karyotypes of 1. javensis，1. platypetala，a nd 1. radicans
are similar and are thought to be closely related to each other. Taxonomical notes for
each species are given.
Key words: Chromosome，cy totaxonomy，Im patiens，k aryomorphology，Ma lesia.
Impatiens (Balsaminaceae) consists of 450 polyploidy and aneuploidy with different
to 850 species distributed in tropical and basic chromosome numbers might play an
temperate regions in the northern hemisphere important role in the speciation of
(Grey-Wilson 1980， Cronquist 1981). Impωiens.
Southeast Asia including Malesian region is Karyological studies have been made for
one of the centers of diversity for Impatiens species of Impatiens in several regions. For
(Grey-Wilson 1980). example，I ndia (Govindarajan and Subrama-
Chromosome numbers of Impatiens have nian 1986，R ao et al. 1986)，t he Himalayas
been examined by many authors and various (Akiyama et al. 1992)，a nd Southwest China
somatic chromosome numbers，2 n =6 ，8 ，1 2， (Sugawara et al. 1994，1 997). For Southeast
14，1 5，1 6，1 7，1 8，1 9，2 0，2 6，2 8，3 2，3 4，3 6， Asian species of Impatiens，a lthough several
40，4 4，4 8，5 0，5 6，ca . 60，h ave been reported chromosome counts have been reported
(see Federov 1969，G oldblatt 1981，1 984， (Jones and Smith 1966，A risumi 1973，1 987，
1985，1 988，G oldblatt and Johnson 1990， Shimizu 1979，L arsen 1981，O kada 1989)，
1991，1 994，1 996，1 998，2 000，2 003). Thus no karyological study has been done.
-271-
272 植物研究雑誌第80巻第5号 平成17年10月
Table 1. Species examined，lo calities and chromosome numbers of five species of Impatiens
Chromosome
Species Localities (voucher specimens and their deposition)
number (2n)
1. chonoceras Hassk. Indonesia. West Java (Jawa Barat); Mt. (Gn.) Papandajan，1 950- 14*
2470 m al .t(Tsukaya & a .l104，9 J uly 2003，T NS，B O)
よjavensisSteud. Indonesia. West Java (Jawa Barat): Gn. Gede-Pangorango Natio- 16*
nal P訂k，1840 m alt. (Tsukaya & a .l235，6 M arch 2003，T NS BO)
よplaη'PetalaLindl. Indonesia. West Java (Jawa Barat); Gn. Gede-Pangorango 16
National Park，2 120 & 1840 m alt. (Tsukaya & al. 259，2 60，8
March 2003，T NS，B O)
よradicansZoll. Indonesia. West Java (Jawa Barat): Gn. Gede-Pangorango 16*
National Park，G n. Gede/Gn. Pangorango，2 400 m alt. (Tsukaya &
al. 240，7 M arch 2003，T NS，B O)
1. kinabaluensis Malaysia. Sabah (Borneo Island); Kinabalu National Park，M t. 12*
S. Akiyama & H. Ohba Kinabalu，T impohon Gate-Layang-Layang，1 770 m alt. (Tsukaya
& al. 0403201，1 4 March 2004，T NS，B ORH**)
*New count for the taxon.
**Herbarium of Institute for Tropical Biology and Conservation，U niversiti Malaysia Sabah.
This paper reports the chromosome num- was 2n = 14，1. javensis，1. platypetala，a nd
bers and karyotypes of five species of 1. radicans were 2n = 16，a nd 1. kinabalu-
Impatiens collected from Java and Borneo， ensis was 2n = 12，r espectively (Table 1).
Malesia. Among them，c hromosome numbers of 1.
chonoceras，1. javensis，1. radicans，a nd 1.
Materials and Methods kinabaluensis were counted for the first time.
Materials were collected from their native
habitat in West Java (Indonesia) and Mt. 1. Impatiens chonoceras Hassk. (Fig. 1a)
Kinabalu，B orneo (Sabah，M alaysia) in 2003 The somatic chromosome number of I.
and 2004. Localities and voucher specimens chonoceras was 2n = 14. This number is
are listed in Table 1. considered to be diploid，w ith basic chromo-
For observation of somatic metaphase some number x= 7. Chromosomes are 1.5-
chromosomes，r oot tips were pretreated in 3.1μm long. Karyologically，t he comple-
the field with a2 m M8 -hydroxyquinoline ment is monomodal，g radually reducing in
solution for about 3 hours and fixed with size. Satellites were observed in two pairs of
Newcomer's fluid (see Sharma and Sharma chromosomes: in the longest and second
1980，W akabayashi 1988). The root tips longest pairs.
were stained by Feulgen's nuclear reaction， Akiyama et al. (1992) reported chromo-
and macerated in am ixture of 2% pectinase some numbers of 16 species in the
and 2% cellulase for 1.5-2 hours. They were Himalayas and reported those of 1. falcifer
again stained with 2 % lacto-propionic and 1. serrata as 2n =1 4. Sugawara et al.
orcein and then squashed and observed using (1997) reported 15 species of. Impatiens in
al ight microscope. Yunnan，S WC hina and chromosome num-
bers of 1. kamtilongensis Toppin and 1.
ResuItsa nd Discussion wuchengyihu S. Akiyama，H . Ohba & S. K.
Chromosome number of 1. chonoceras Wu as 2n = 14. All the species mentioned
October 2005 Journal of Japanese Botany Vo .l80 No. 5 273
above are similar in having characteristic flo- area.
ral morphology; the shape of the lower sepal
is navicular tapering to the spur and the in- 2. Impatiens javensis Steud.，1. platypetala
florescence usually has one or two flowers Lindl. and 1. radicans Zoll. (Figs. 1b- e)
(Akiyama et al. 1991，1 996). Morphologi- The somatic chromosome numbers of 1.
cally 1. chonoceras is similar to these spe- javensis，1. platypetala，a nd 1. radicαns were
cies. Impatiens chonoceras has al ower sepal 2n = 16. These numbers are considered to be
with pubescence on the outer surface，li ke 1. diploid，w ith basic chromosome number x=
kamtilongensis and 1. wuchengyihii，w hile 8. Chromosomes are 0.9-1.8μm long in 1.
the lower sepals of l. falc俳randよserrata javensis，1 .0-1.5μm in 1. platypetala，a nd
are glabrous on the outer surface. The inflo- 1.1-1.7μm in I. radicαns. Karyomorpho-
rescence ofよ chonocerαsis axillary and logically they showed similar karyotypes.
，
pedunculate，a nd usually consists of two The complements were monomodal，g radu-
flowers with ab asal bract. These two flowers ally reducing in size. Satellites were ob-
appear in opposite. The inflorescences of 1. served in one pair of chromosomes.
kamtilongensis and 1. wuchengyihii are Van Steenis (1948) recognized three sub-
racemose with two flowers (Akiyama et al. species in 1. plaη'Petala: subsps. platypetala，
1996)，wh ile those ofよfalciferand l. serrata aurantiaca，a nd nematoceras. Among the
are unique and have a middle bract subspecies，s ubsp. platypetala has the widest
(Akiyama and Ohba 2000). It is notable that distribution range， throughout Jav a and
よchonocerashas the apparently most primi- Sumatra. Zinov' eva-Stahevitch and Grant
tive form of inflorescence among Chinese (1984) reported chromosome numbers of 1.
species (i. e.， 1. kamtilongensis and 1. plaη'Petala subsp. platヲ'Petalaas n= 7 and
wuchengyihii) and Himalayan species (i. e.， 2n =1 4 (from Mt. Gede， Java)， subsp.
よfalciferandよserratα). aurantiaca 2n =1 4 (from Sulawesi)， and
Cytologically，a ll the species mentioned subsp. nematoceras n= 8a nd 2n = 16 (from
above have similar karyotypes. Impatiens Indonesia). Arisumi (1987) reported the
chonoceras， 1. kamtilongensis， and 1. chromosome number of 1. platypetαlα
，
wuchengyihii have somatic chromosome (subsp.) aurantiaca as 2n =8. Okada (1989)
numbers 2n = 14，w ith monomodal comple- counted chromosome numbers of 1.
ments. Impatiens falcifer and 1. serrata also plaη'Petala from Sumatra，w hich may be
have somatic metaphase chromosomes 2n = subsp. platypetala based on distribution，a s
14，a nd they show monomodal complements 2n =1 6. Our count for 1. platypetala，r efers
as judged from the plates (Akiyama et al. to subsp. plaη'Petala， coincides with the
1992). number counted by Okada (1989)，bu t differs
Impatiens chonoceras was collected in Mt. from the number given by Zinov'eva-Stahe-
Papand司jan，af amous volcano located in SW vitch and Grant (1984) for ac ollection from
Jav a. From the morphology and cytology，1. the same locality. For confirming the chro-
chonocer仏，5・isconsidered to be closely re- mosome numbers of the species and the
lated to the species of the Himalayas and infraspecific taxa of 1. plaη'Petala，ex amina-
adjacent regions. Potentilla polyphylla Wall. tion of ample materials collected from the
ex Lehm. (Rosaceae)，d istributed mainly in entire range of distribution is necessary.
the Himalayas and a匂acentregions，w as Impatiens platypetala，1. javensis，a nd 1.
collected in this mountain (Kalkman 1968， radicans are closely related species judging
1993)，s uggesting that Mt. Papandajan had from floral mo叩hologyand kaηotypes. In
been ar efugium for Himalayan plants in this Gunung (Mt.) Gede-Pangrango National
274 植物研究雑誌第80巻第5号 平成17年10月
嫌 事際調惨事襲
1
聖書
a臨調畢
Fig. 1. Somatic metaphase chromosomes of five species of Impatiens in SE Asia. a: 1. chonoceras (2n =
14). b: 1. javensis (2n =16 ). c，d: 1. platypetala (2n =16 ). e: 1. radicans (2n =16 ). f: I. kinabaluensis
(2n = 12). Arrows indicate satellite chromosomes. Bar indicates 5ドm.
October 2005 Journal of Japanese Botany Vo .l80 No. 5 275
Park， they inhabit different altitudes: 1. probab1y a1so occurs in Borneo Su1awesi
ヲ
plaηpetala at the 10west a1titudes，1 ess than and the Lesser Sunda Is1ands". Masamune
1800 m，1. javensis between 1800 m and (1942) 1isted 1. platypetala among Bomean
2400 m，a nd 1. radicans above 2400 m. Impαtiens.
Impatiens plaηpetala is distributed wide1y in After examining the morpho1ogy of
Ma1esia whi1e l. javensis is found in Java Impatiens collected in Mt. Kinabalu，B orneo
and probab1y in Sumatra，a nd 1. radicans is (Tsukaya & al. 0403201)，it is concluded that
endemic to West Jav a (as far as known，f rom this is not 1. platypetala，b ut an ew species，
Mts. Pangrango，T jikurai and Papandjan) named 1. kinabaluensis (Akiyama et a .l
‘
(van Steenis 1972). Impatiens javensis and 2005)
I. radicans are genetically close and it is
known that they produce interspecific hybrid Wet hank Prof. Hiroshi Okada，O saka City
(Tsukaya 2004). It may be thought that 1. University，fo r giving information about cy-
javensis and 1. radicans might be differenti- tology of Impαtiens in Southeast Asia. This
ated in dip10id 1eve1 after altitudina1 iso1ation study was partly supported by aG rant-in-Aid
from an 1. plαηpetala-1ike ancestor having for Scientific Research (B) for the Jap an
16 somatic chromosomes in Jav a Is1and. Society for the Promotion of Sciences，n o.
1440517 (to H. T.).
3. Impatiens kinabaluensis S. Akiyama & H.
Ohba (Fig. 1f) References
The somatic chromosome number of 1. Akiyama S. and Ohba H. 2000. Inflorescences of the
kinabaluensis was 2n =1 2. The number is Himalayan species of Impatiens (Balsaminaceae).
considered to be dip1oid，w ith basic chromo- J. Jpn. Bot. 75(4): 226-240.
some number x= 6. Chromosomes are 2.4- 一一一，一一-and Wakabayashi M. 1991. Taxonomic
notes of the east Himalayan species of Impatiens
3.8μm 10ng. Karyo1ogically，t he comp1e-
(Balsaminaceae). In: Ohba H. and Malla S. B.
ment is monomoda1，g radually reduced in (eds.)，T he Himalayan Plants 2: 67-94. University
size. Satellites were not observed. of Tokyo Press，T okyo.
Chromosome numbers of Impαtiens in 一一一， 一一一 andWu S.-K. 1996. Further notes of
Southeast Asia including Ma1esia have been Impatiens (Balsaminaceae) from Yunnan，C hina.
Bull. Natn. Sci. Mus.，T okyo，S er. B，22 (4): 135-
fragmentari1y reported (Jones and Smith
144.
1966，A risumi 1973， 1987，S himizu 1979， 一一一， Wakabayashi M. and Ohba H. 1992.
Larsen 1981， Zinov'eva-Stahevitch and Chromosome evolution in Himalayan Impatiens
Grant 1984，O kada 1989)，w ith those of (Balsaminaceae). Bot. J. Linn. Soc. 109: 247-257.
about 20 species so far revea1ed. Among 一一一， Ohba H.，I keda H.，T sukaya H. and Mohamed
M. 2005. Anew species of Impatiens
these species，t wo species were reported as
(Balsaminaceae) from Mt. Kinabalu，B orneo. J.
2n = 12; I. chiangdaoensis T. Shimizu from
Jpn. Bot. 80: 266-270.
Thai1and (Shimizu 1979，L arsen 1981) and Arisumi T. 1973. Chromosome numbers and inter-
“1. harlandii Dransfie1d ined." from Borneo specific hybrids among New Guinea Impatiens spe-
(Zinov'eva-Stahevitch and Grant 1984) a1- cies. 1. Hered. 64: 77-79.
though we cou1d not find any 1iterature on 一一一 1987.Cytology and morphology of ovule cul-
ture-derived interspecific Impatiens hybrids. 1.
“1. harlandii" ， and cou1d not determine
Amer. Hort. Sci. 112(6): 1026-1031.
whether this is the same species as 1.
Cronquist A. 1981. An Integrated System of Classifi-
kinabaluensis or not. cation of Flowering Plants. Columbia University
Van Steenis (1948) noted the distribution Press，N ew Yo rk.
of 1. plaηpetala subsp. plaηpetala as Federov A. A. 1969. Chromosome Numbers of
Flowering Plants. Komar. Bot. Ins ，.tAcademy of
“throughout Jav a，f urther in Sumatra and
276 植物研究雑誌第80巻第5号 平成17年10月
Science，L eningrad. J. J. O. (eds.)，F lora Malesiana ser. 1，1 1(2): 227-
Goldblatt P. (ed.) 1981. lndex to Plant Chromosome 351.
Numbers 1975-1978，p. 143. Missouri Botanical Larsen K. 1981. Chromosome numbers in Impatiens
Garden，S t. Louis. from Thailand. Nord. J. Bot. 1: 43-44.
一ー一(ed.)1984. lndex to Plant Chromosome Numbers Masamune G. 1942. Enumeratio Phanerogamarum
1979-1981，p. 109. Missouri Botanical Garden，S t. Bomearum， p. 433. Taiwan-sotokufu-gaijibu，
Louis. Taihoku.
一一一(ed.)1985. lndex to Plant Chromosome Numbers Okada H. 1989. Chromosome counts of some plants
1982-1983，p. 60. Missouri Botanical Garden，S t. collected from West Sumatra. Occasional papers，
Louis. Kagoshima University Research Center for the
一一一(ed.)1988. lndex to Plant Chromosome Numbers South Pacific (16): 11-14.
1984-1985，p p. 67 & 68. Missouri Botanical Rao R. V. S.，R angaswami Ayyangar K. and Sampath-
Garden，S t. Louis. kumar R. 1986. On the karyological characteristics
一一一andJohnson D. E. (eds.) 1990. lndex to Plant of some members of Balsaminaceae. Cytologia 51:
Chromosome Numbers 1986-1987，p p. 54 & 55. 251-260.
Missouri Botanical Garden，S t. Louis. Sharma A. K. and Sharma A. 1980. Chromosome tech-
一一-and --(eds.) 1991. lndex to Plant Chromo- niques. Theory and Practice， 3rd ed.， p. 55.
some Numbers 1988-1989， p. 71. Missouri Butterworths，L ondon.
Botanical Garden，S t. Louis. Shimizu T. 1979. A comment on the limestone flora of
and一一一 (eds.)1994. lndex to Plant Chromo- Thailand，w ith special reference to Impatiens. Acta
some Numbers 1990-1991， p. 63. Missouri Phytotax. Geobot. 30: 180-187.
Botanical Garden，S t. Louis. van Steenis C. G. G. J. 1948. The species of Impatiens
一一-and一一一 (eds.)1996. lndex to Plant Chromo- indigenous to Java (Bals.). Bull. Jard. Bot.
some Numbers 1992-1993， p. 73. Missouri Buitenzorg，se r. 3，1 7(4): 384-387.
Botanical Garden，S t. Louis. 一一一 1972. Balsaminaceae. In: Hamzah A.，
一一-and --(eds.) 1998. lndex to Plant Chromo- Moehamad T. and van Steenis C. G. G. J.，T he
some Numbers 1994-1995，pp . 51 & 52. Missouri Mountain Flora of Java，P l. 6. E. J. Bril1， Leiden.
Botanical Garden，S t. Louis. Sugawara T.，A kiyama S.，M urata 1. and Yang Y. P.
一一-and一一一 (eds.)2000. lndex to Plant Chromo- 1994. Karyology of ten species of Impatiens
some Numbers 1996-1997，pp . 38 & 39. Missouri (Balsaminaceae) from SWC hina. Acta Phytotax.
Botanical Garden，S t. Louis. Geobot. 45: 119-125.
一一一and一一一 (eds.)2003. lndex to Plant Chromo- 一一二一一一， Yang Y. P. and Murata J. 1997.
some Numbers 1998-2000， p. 75. Missouri Karyological characteristics of Impatiens
Botanical Garden，St . Louis. (Balsaminaceae) in Yunnan，C hina. Acta Phytotax.
Govindarajan T. and Subramanian D. 1986. Karyo Geobo .t48: 7-14.
・
taxonomy of south lndian Balsaminaceae. Cyto- Tsukaya H. 2004. Gene flow between Impatiens
logia 51: 107-116. radicans and I. javensis (Balsaminaceae) in
Grey-Wilson C. 1980. Impatiens of Africa. A. A. Gunung Pangrango，Ce ntral Java，ln donesia. Amer.
Balkema，R otterdam. J. Bot. 91(12): 2119-2123.
Jones K. and Smith J. B. 1966. The cytogeography of Wakabayashi M. 1988. Present situation of cyto-
Impatiens L. (Balsaminaceae). Kew Bull. 20: 63- taxonomy of Himalayan plants. Newslett. Himala-
72. yan Bot. (3): 8-11.
Kalkman C. 1968. Potentilla， Duchesnea， and Zinov' eva-Stahevitch A. E. and Grant W. F. 1984.
Fragaria in Malesia. Blumea 16(2): 325-354. Chromosome numbers in Impatiens (Balsamina-
一一一 1993.Rosaceae. In: Kalkman C.，K irkup D. W.， ceae). Canad. J. Bot. 62: 2630-2635.
Nooteboom H. P.，S tevens P. F. and de Wilde W.
October 2005 Journal of Japanese Botany Vol. 80 No. 5 277
池田博，秋山忍 ，塚谷裕一<， M.マハメド ，
a h d
D.ダルナエデイ ジャワ島およびボルネオ島産
e.
ツリフネソウ属植物(ツリフネソウ科)の細胞分 ワ島付近に限られる 1.javensisと1.radicansは，
類学的研究 2n =1 6の染色体数を持つI.platypeωla類似の祖先
ジャワ島(インドネシア)およびボルネオ島キ 種から垂直的な棲み分けにより分化したものでは
ナバル山(マレーシア)で採集されたツリフネソ ないかと推定された.Impatiens kinabaluensis は，
ウ科ツリフネソウ属 5種の染色体数と核型を報告 形態、と核型から，これまで記載されていなかった
した. 新種であると考えられた.
Impαtiens chonoceras Hassk.は2n= 14，1 . javensis (a岡山理科大学総合情報学部
Steud.，1. platypetala Lind ，.l1. radicans Zoll.'ま2n= 生物地球システム学科，
16， l. kinabaluensis S. Akiyama & H. Ohba は2n 国立科学博物館植物研究部，
ニ b
12であった.Impatiens chonoceras，I. jαvensis，1. 自然科学研究機構・
C
radicans，1. kinabaluensisの染色体数は，今回が初 岡崎統合バイオサイエンスセンター;
めての報告である.Impatiens chonocerasは，花や 総合研究大学院大学先導科学研究科;
花序の形態と核型から，ヒマラヤや中国南西部に 京都大学大学院理学研究科
分布する種と類縁があると考えられた.Impatiens マレーシア・サパ大学，
d
javensis，1. platypeωlα，1. radicansの3種は，花の eインドネシア・ボゴール植物園)
形態と核型から互いに近縁と考えられ，分布がジャ
