Cytotaxonomic Study of Ferns from China II. Species of Fujian

10 Pages·2002·1.1 MB·English

by  Su-Juan Lin

・4ふふ市巴物bE1涜恥・ホ正也誌
，τd一司A1L山L
J'周，.ウB 判3nxuρAU nU ウM-EF
Cytotaxonomic Study of Ferns from China 11. Species of Fujian
Su-Juan LIN ，aHui-Ru ZHUANGb，K unio IWATSUKIc and Hai-Sheng Lub
aDepartment of Biological Science，Gr aduate School of Science，Un iversity of Tokyo，
Hongo 7-3-1，B unkyo-ku，T okyo，1 13-0033 JAPAN;
TheB ioengineering College，F 吋ianTeacher's University，F uzhou 35007，Fu jian，C HINA;
b
CThe University of the Air，W akaba 2-11，M ihama-ku，Ch iba，2 61-8586 JAPAN
(Received on August 8，2 001)
A cytotaxonomic study of ferns from Fujian Province，C hina，w as carried out. The
reproductive modes of 31 species in 18 genera were examined using ac onventional
method，a nd the chromosome numbers were counted for 24 species，in cluding 12 newly
examined in China. The chromosome numbers of both Lygodium microstachyum and L.
scandens are 2n =6 0. Pteris sem伊innatais as exual tetraploid with 2n =1 16 chromo-
somes，w hile P. esquirolii has 2n = ca. 90 chromosomes and is considered to be an
agamosporous triploid. Five species were observed in Dryopteris，al l of them being
agamosporous. Deparia japonica has been recorded from Jap an as ah exaploid，wi th n=
120 chromosomes but this study reports as exual tetraploid with 2n =1 64 chromosomes
from Fujian. Thelypteris glanduligera and T. parasitica are agamosporous triploids with
=
2n 108 chromosomes. (Continued from J. Jpn. Bot. 71: 214-222，1 996)
Key words: agamospory，F ujian Province，r eproductive mode，p teridophyta.
In studies of the biodiversity and Obviously，F ujian is also one of the prov-
systematics of Asian pteridophytes，cy tologi- inces with ar ich pteridophyte flora. Four-
cal information from Chinese ferns is indis- hundred and four species (including 25 en-
pensable. Although more than 300 Chinese demic) of ferns and fern-allies have been re-
pteridophytes have been studied cyto- corded in F吋ian[Lin et a ，.l 1982，f ollowing
logically by both Chinese and overseas re- Ching's system (1978)]. Approximately 37
searchers (Lin et al. 1996)，t his is only % of the total Fujianese species range to
around 10 % of the 3000 or so species found Jap an，a nd more than 12 % of the species to
in China. Lin et al. (1996) have previously Taiwan. As far as we know，t he F吋ianese
performed ac ytotaxonomic study of Chinese pteridophytes have never been investigated
ferns (Series 1) from Yunnan. As the con- cytologically，w ith the exception of af ew
tinuance，in the present study，w e carried out Dryopteris species (Weng 1989).
cytological observations on material from As af irst step in the systematic studies of
F吋ian. Fujianese pteridophytes，w e porthere the
問
Fuiian Province is located in the south- chromosome numbers of 24 species and re-
eastern coastal area of China. The variable productive modes of 31 species. The chro-
landforms and warm，h umid climate provide mosome numbers of some species are
Fujian with various ecosystems，w hich sup- counted for the first time in China. The pre-
port an abundance of plant diversity. sent study expects to contribute to our wider
-129-
130 植物研究雑誌第77巻第3号 平成14年6月
knowledge of the Chinese fems，a nd to make reproductive modes (Yoroi and Iwatsuki
possible a comparative study of the 1977，M urakami and Iwatsuki 1982，K ato et
pteridophyte floras of East China and Japan. al. 1992). The presence of 32 spores per spo-
rangium is supporting evidence for
島'Iaterialsand Methods distinguishing agamospory from the normal
As shown in Table 1，ma terials were col- sexual type with 64 spores per sporangium.
lected from the Linshi Forest center of In cases where spores escaped from the split
Fuqing，t he Forest Park of Fuzhou，a nd of sporangia of dry specimens，th e spore num-
the Gushan mountain in Fuzhou，so utheast of ber was estimated to be 64 based on practice
Fujian. Voucher specimens 'edeposited in counts of more than 32 in each sporangium.
紅
the Herbarium，t he Bioengineering Col1ege， On the other hand，if the practice count was
Fujian Teacher乍 University. lower than 32，it was considered to be 32-
Fresh root tips were pretreated with 0.002 spored.
M 8-hydroxyquinoline solution for 4-6 In the present study，t he chromosome
hours at 18-20oC，a nd then fixed in 3: 1 a ce- numbers of 12 species were newly reported
tic acid: alcohol (99 %) in the field，fo r ob- for China. Cytotaxonomically noteworthy
servation of somatic chromosomes. The species e'described and discussed below.
訂
observation method follows Lin et al. (1996).
Frond material was fixed in K'nop solution Lygodium About 40 species of Lygodium
for gametic observation and spore counting. are recorded worldwide (Iwatsuki 1992).
Dry specimens were also used for spore Only a few species have been studied
counts，w ith more than 10 sporangia being cytologically，t he basic chromosome num-
counted for each individual. bers of the genus being variously reported as
x= 28，2 9 or 30 (Mitui 1965，1 968，K urita
Results and Discussion 1967，T sai and Shieh 1983，R oy and Rao
The results are shown in Table 1a nd Figs. 1985，W alker 1985，S ingh and Roy 1988，
1-18. The somatic chromosomes of 24 spe- Na kato 1990， Ammal and Bhavanandan
cies belonging to 14 genera of 10 families 1992，T akamiya 1995).
were examined and the reproductive modes There are eleven species of Lygodium in
of 31 species of 18 genera were inferred by China，fo ur of which are found in F吋ian，but
spore counting. The frond materials were no previous cytological data were available
used for spore counts as the young spores in for them. In the present study，t wo species，
each unopened sporangium can be counted L. microstachyum Desv. and L. scandens (L.)
exactly. However，f ronds were too old to be Sw.，w ere observed cytologically. The chro-
used for gametic chromosome counting in mosome numbers of both species were re-
the present study. Dry specimens were used corded as 2n = 60. This coincided with the
for spore counts in those species for which numbers n= 3 0，2n =6 0 of L. scandens from
we did not have suitable frond material. In Nepal (Srivastava 1985)，bu t not with n= 29
most higher leptosporangiate fems， 64 in L. microstachyum of Taiwan (Tsai and
haplospores eproduced in each sporan- Shieh 1983). In Japan，L. microstachyum is
訂
gium in sexually reproductive species，w hile distributed in the south of the Ryukyus，a nd
32 diplospores are produced in each sporan- is considered to be av ariety of L. japonicum
gium in agamosporous species. A few sex- (Thunb.) Sw. (n =2 9，2n =58 ). To determine
ual，3 2-spored groups，s uch as lindsaeoid the correct base number and the relationship
fems，a re the exceptions (Lin et al. 1990). between L. microstachyum and L. japoni-
Thus，sp ore counts can be used for inferring cum， further research is encouraged.
June 2002 Journal of Japanese Botany Vo .l77 No. 3 131
Figs. 1-5. Photographs of somatic chromosomes. Scale bar =1 0μm. 1. Lygodium microstachyum 2n =60 .
2. Microlepia marginata 2n =c a. 164. 3. M. strigosa 2n =8 4. 4. Lindsaea orbiculata 2n =1 76. 5.
Sphenomeris chinensis 2n = ca. 190.
Microlepia Microlepia is a well-known Qiu (1988)，K ato et al. (1992)，N akato et al.
genus in Asia. About 81 % of the 70 known (1995) and Kato and Nakato (1999) in these
species have been recorded from China，fi ve studies，t he basic numbers were considered
of these being recorded from Fujian. Only to be x= 42 or 43.
four Chinese species were studied Two species， M. marginata and M.
cytologically by Weng (1985b)，W eng and strigosa， were collected for this study.
132 植物研究雑誌第77巻第3号 平成14年6月
Figs. 6-12. Photographs of somatic chromosomes. Scale bar =1 0 Ilm. 6. Pteris semipinnata 20 =11 6. 7.
Woodwardiajaponica 20 =6 8.8. Arachniodes exilis 20 =82 . 9. Dryopteris gushanica 2n =1 23. 10. D.
varia 2n =1 23. 11. Thelypteris glanduligera 20 =1 08. 12. T. acuminata 2n =72 .
June 2002 Journal of Japanese Botany Vol. 77 No. 3 133
Cytologically， triploids， te位aploids and ag ood explanation for the ex住emediversity
hexaploids have been reported in M. seen in this genus.
marginata from China，T aiwan and Japan Pteris esquirolii was cytologically ob-
(Mitui 1968，N akato and Serizawa 1981， served here for the first time in China. Based
Tsai and Shieh 1983，W eng 1985b，W eng on the chromosome number 2n = ca. 90，a nd
and Qiu 1988，N akato et al. 1995). For about thirty-two spores in each sporangium，
Chinese materials，t he chromosomes were it is suggested to be an agamosporous
counted as n =8 6 by Weng (1985b) and triploid in Fujian.
Weng and Qiu (1988) from Zhejiang
Province and 2n = 168 by Na kato et al. Woodwardia japonica Except for as ingle
(1995) from Jiangsu Province. Microlepia report (n = 17) from Zhejiang (Weng and
marginata is am orphologically variable and Qiu 1988)，W oodwardiajaponica has always
very common species in F吋ian.Our chro- reported as as exual，di ploid species with n=
mosome and spore counts showed 2n = ca. 34，2 n = 68 chromosomes in China and
164，a nd about 58 (64) spores in each spo- Japan (Mitui 1965，1 968，T akamiya et al.
rangium in M. marginata，so we suspect it to 1992，Na kato et al. 1995). The Fujianese ma-
be as exual tetraploid. terials were counted to be 2n =6 8 here，to o.
Microlepia strigosa is widespread in the Conventionally，th e basic chromosome num-
Asian tropics. In Jap an，i ts chromosome ber of Woodwardiαis considered to be x=
numbers have been reported as n= 43，2 n = 31，3 4. As Takamiya et al. (1992) pointed
86 by Mitui (1965， 1968， 1973， 1976)， out，if the lowest number is really 17，t hese
Sasamoto (1970) and Kurita (1972)，a nd species (including W. <Iαponicα) usually
印刷
2n = 84 by Kawakami (1979)，N akato and ferred to diploid should be considered as
Serizawa (1981). Tsai (1992) reported tetraploids.
n= 4311，86 11，2 n = 86，1 72 from Taiwan. In
the present observation，h owever，t he chro Dη'opteris About 300 species have been
闇
mosomes were counted as 2n =8 4，w hich is recorded for Dryopteris in China，o f which
an ew record from China. These results sug- about sixty species have been observed
gest it should be considered as exual，di ploid cytologically (Chiu 1981，W ang and Zhang
species according to both chromosome and 1981，W ang and Xia 1984，W ang 1985，
spore numbers (Table 1). Weng 1985a，1 989，N akato et al. 1995)，
most of which (more than 60 %) were
Pteris Pteris is al arge genus with about agamosporous triploids.
300 species in the world，i ncluding about As Table 1 shows， five species of
110 species in China. In Fujian，s ixteen spe- Dryopteris were observed here，t hree of
cies have been recorded. them enew cytological records for China.
訂
Cytological and systematic studies of All of them eagamosporous triploids，
紅
Chinese Pteris were performed by Chiu except for D. hwangii Ching，f or which，a n
(1981)，W ang (1989)，W eng (1985a，1 990)， estimated 80 chromosomes and 29-32 spores
Weng and Qiu (1988)，K ato et al. (1992)， in each sporangium suggest it was an
Nakato et al. (1995)，L in et al. (1996)，a nd agamosporous diploid species，n o matter de-
Kato and Nakato (1999). Although only spite the fact that the chromosome number
about 10 % of Chinese species are known could not be counted exactly. It was clearly
cytologically， the occuence of diploid， showed that agamosporous triploids epre-
町 訂
triploid and tetraploid cytotypes and sexual dominant in the evolutional diversity of
and agamosporous reproductive modes give Chinese Dηopteris.
Table 1. Materials and chromosome numbers of fern plants from Fujian，C hina ムw同一
Chromosome Sexuality Photo Chromosome Sexuality Photo
Family & species Voucher S/S Family & species Voucher S/S
no. (2n) & ploidy no. no. (2n) & ploidy no.
Schizaeaceae A. rhomboidea 97-48 56 sex.，4xワ
*LL.y gmoicdriousmt a(xc h=yu2m 9 ，D3e 0s)v 96-27 100-102 60 D，ツ(Wopatlelr.)i sCh(xi n=g 4 1) 49， 63 (64)
*L. scandens (L.) Sw 97-17 95 60 D. championi 97-3，8， 30，30， c. 123 apo.，3x
(Benth.) Ching 10，11，27， 28，30，30，
Dennsta巴dtiaceae 30，34， 30，30，
Microlepia (x =4 2，43 ) 41，42 32，30
M. hancei Prantl 97-19 64 sex. 本D.dehuaensis Ching 97-37 25 (32) ~120 apo.，3x
M. marginata (Houtt.) 97-20 58，58， c. 164 sex.，4x *D. gushanica Ching 97-24，28 32，29 123 apo.，3x 9
C. Chr. 97-36，44 58 (64) 傘D.hwargii Ching 96-7 29-32 c.80 apo.，2x
*M. strigosa (Thunb.) 97-38，45 55，55 84 sex.，2 x D. varia (L.) O.Kuntze 97-7，54 32，32 123 apo.，3x 10
Presl (64)
Thelypteridaceae
Lindsaeaceae Thelypteris (x =2 7，2 9，3 0，3 1，
Sphenomeris (x =4 7，4 8，4 9) 32，3 4，3 5，3 6)
S. chinensis Maxon 97-4 32 c，190 sex.，4x 事T.acuminata 97-39 49 (64) 72 sex.，2x 12 茜
Lindsaea (x = 44，4 7) (Houtt.) Morton 書
L. orbiculata (Lam.) Mett. 97-12 30，30 176 sex.，4x 4 T. glanduligera 97-29，32 28，28 108 apo.，3x 11 出
司
97-13 (32) (Kunze) Ching (32) 明
Sinopteridaceae 間
On(yTchhuinub.m) j aKpuonnziec um 97-9，35 64，64 sex ホT.p(aL.r)a sKit.Iicwaa ts 972-52 3， 262 (63，2) c. 108 apo.，3x 14 帯別什
*1ぺparvifoliaChing 97-5 64 c.124 sex.ワ 13
Adiantaceae 鴻
Adiantum (x = 30) Woodsiaceae ゴ
A. flabellulatum L. 97-15，16 61，61 sex Deparia (= A thyrium p.p.，x = 枇
(64) 40，41) ゆ
PPtteerriisd a(cxe a=e 2 9) 事D.j(Tahpuonnbi.c)a M.Kato 97-33 64 164 sex.，4x 15 論
P. ensiformis Burm 97-6，18 45，45 (64) sex. Diplazium (40，4 1) u
ホP.esquirolii Christ 96-4 30 (32) c.90 apo.，3x D. subsinuatum 97-60 64 sex. 叩h
P. semipinnaωL. 97-2，26 50，59 (64) 116 sex.，4x (Wall.) Tagawa
Aspleniaceae Polypodiaceae
Asplenium (x = 36，3 8，39 ) Lepisorus (x = 35，50 )
A. bullatum Wall. ex Mett. 97-50 64 注130 sex.，4x *L. thunbergianus 97-47 64 100 sex.，4x? 16
(Kaulf.) Ching
Blechnaceae Microsporum (x = 36)
Blechnum (x =3 1，3 3，3 4) M. fortunei 97-31 64 c.72 sex.，2x 17
B. orientale L. 97-1， 28 (32) apo. (Moore) Ching
Woodwardia (x = 31，3 4) Colysis (x =3 6)
W. japonica (L.f.) Sm. 97-14，46 40，40 68 sex.，2x C. pothifolia (Don) Presl 97-43 64 72 sex.，2x 18
(64)
Dryopteridacea巴
Arachniodes (x = 41)
A. exilis (Hance) Ching 9555f今ゐ弓い寸，勺，'p一ココ宍勺JUFNd8L244同 5044，8754，，0845 ，(806，，4 ) 82 sex.，2x 」N
The specimens of voucher number from 97-1 to 97-22 were collected from the Forest Park of Fuzhou; 96-and 97-23 to 97-50 from Mt. Gushan，F uzhou; 97-51 to 97-60f rom Linshi Forest center of Fuqing. 国何
S/S: Spore number in each sporangium. 回
品
( ): estimated spore number in each sporangium based on the practice count. 同
*: new records合omChina 明。
泊
June 2002 Journal of Japanese Botany Vo .l77 No. 3 135
Figs. 13-18. Photographs of somatic chromosomes. Scale bar = 10μm. 13. Thelypteris parvifolia 2n = ca.
124. 14. T. parasitica 2n =ca . 108. 15. Depariαjaponicα2n =1 64. 16. Lepisorus thunbergianus 2n =
100. 17. Microsporumfortunei 2n =ca . 72. 18. Colysis pothifolia 2n =72 .
136 植物研究雑誌第77巻第3号 平成14年6月
Thelypteris This is an extremely variable Lepisorus thunbergianus Lepisorus is an
genus cytologically. The base number of the extremely variable genus in terms of cytolo-
genus runs from 27 to 36 (Takamiya 1996). gy. Only two species have been observed
Four species were observed in this study. cytologically in China (Lin et al. 1996).
Thelypteris glanduligera has been reported Lepisonふ5・thunbergianusis cytologically
to be n=54 (as Parathelypteris glanduligera， complicated，it s chromosome numbers hav-
Weng 1985b)，a nd n= 68 in China (Weng ing been variously reported as n = 25，5 0，
and Qiu 1988)，a nd n = ca.72 (sex. 4x) in 2n =5 0，5 1，7 5，7 6，1 00，1 01，1 02 and 103，
Japan (Mitui 1968). In the present study，it including examples of polyploidy and
was confirmed to be 2n =1 08. Ona ccount of aneuploidy from Jap an (Mitui 1966a，b ，
the base number for the genus x= 2 7 [based 1968，1 971，1 975，T akei 1974，1 978，1 982，
on the count by Weng (1985b)，n = 5 4]，a nd Masuyama and Mitui 1983，M asuyama et al.
the spore number 32 in each sporangium，w e 1987，Mi tui et al. 1987). In the present study，
considered our specimen to be an agamospo- however，a c hromosome number of 2n =1 00
rous te仕aploid. If the basic number for was counted for Chinese materials.
Thelypteris is x= 3 6，o n the other hand，th is
individual should be considered an This study was supported by financial aids
agamosporous triploid. The co町ectconclu- from the Ministry of Education，C ulture，
sion is expected by the careful examination Sports，Sc ience and Technology for Oversea
of k yotype(chromosome genome set) and Scientific Program.
訂
meiotic division. The second species，T .
parasitica (Cyclosorus parasiticus) has been References
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林蘇絹a，荘恵如b，岩槻邦男c，慮海生d.中
国産シダ類の細胞分類学的研究11.福建省の種
中国福建省のシダ植物の細胞分類学的研究をお た結果， 4種が2n= 123の無配生殖の3倍体， 1種
こなった. 18属31種につき胞子数によって生殖様 が無配生殖の 2倍体であった.また，シケシダ
l
式を推定し， 24種の染色体数を明らかにした.そ Deparia japonica は日本ではn= 120の6倍体しか
のうち， 12種の染色体数は中国での初報告である. 報告されていないが，福建産のものは2n= 164の
特筆すべき知見は以下のようである. 有性生殖4倍体であることが今回の研究で、解った.
中国産のカニクサ属の染色体は未調査であった ハシゴシダ，Thelypterisglanduligeraとケホシダ T.
が，本研究により ，Lygodium microstachyumとL. parasitica 土日本ではどちらも n= 72の有性生殖
scandensの染色体数は2n= 60であることが明らか の4倍体であったが，福建省産のものは2種とも
になった.Pteris esquiroliiは2n= ca. 90の無配生 2n = 108で無配生殖の 3倍体であることがわかっ
殖3倍体種であり， P. semipinnataは2n= 116で た. (a東京大学大学院理学系研究科生物科学専攻，
有性生殖の4倍体であることが従来の n= 58の報 b福建師範大学生物工程学院，
告と一致した.オシダ属Dryopterisの5種を調べ c放送大学)