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植物研究雑誌 J. Jpn. Bot. Originals 73: 119-124 (1998) Aneuploidy in Thelypteris laxa (Franch. & Sav.) Ching (Thelypteridaceae) Na rumi NAKATO Kokubunji High School，3- 2-5 Shin-machi，K okubu吋i-shi，Tokyo，1 85-0004 JAPAN (Recei ved on No vember 17，1 997) Chromosome number of Thelypteris laxa was determined，a nd two cytotypes， 2n=136 and 2n=134 were found. The former cytotype is regarded as tetraploid based on x=34，an d the latter as aneuploid (hypotetraploid) of the former. In Jap an，th e hypotetraploid seems to have aw ider distribution range than the eutetraploid. Thelypteris laxa is distributed in Japan， sporangium was counted to ascertain whether Korea (Cheju Is1.)， southern China and Tai- reproduction was sexual or apogamous. wan. In Jap an it is commoni n warmt emperate Voucher specimens are deposited in TI (the regions，o ccurring mainly in lightly shaded Botanical Gardens，Un iversity of Tokyo). places in lowlands (Iwatsuki 1995). Preceding chromosomal works have shown that this spe- Results and Discussion cies has two basic chromosome numbers，x= 34 Chromosome number Chromosome num- and 36; the former was suggested from mitosis bers were determined in 24 plants from 19 in root tip cells as 2n=1 36，a nd the latter from localities in Jap an (Table 1). Oft hese，3 p lants meiosis in spore mother cells as n=36，3 61V or from 210calities were found to be 2n= 136，an d 72 (see Table 1). As to Japanese plants，Ku rita the remaining 21 plants from 17 localities (1963) reported n=72 (x=36) from Kikugawa， were 2n=134 (Figs. 1& 2). The plants of Shizuoka Pref.，w hile Nakato (1996) counted 2n=136 and those of 2n=134 were indistin- 2n= 136 (x=34) from Tsukubasan，Ib araki Pref. guishable morphologically. In this paper，1 i ntend to report the results of a The basic chromosome numbers available further chromosomal survey on plants from 19 so far in the genus Thelypteris sensu Iwatsuki localities in Jap ani，nc luding an ewa neuploidal (1995) are diverse and are reported to be x=27， cytotype which widely occurs. 30，3 1，3 2，3 4，3 5 and 36 (Takamiya 1996). The 2n= 136 cytotype of T. laxa could safely be Materials and Methods regarded as tetraploid based on x34，agreeing 二 Localities of materials are listed in Table 1. with the observations of Nakato (1996)，a nd Plants were collected from native habitats，an d the 2n= 134 cytotype，ne wly found in this study， cultivated in pots. For the observation of so- as aneuploid (hypotetraploid) of the former. matic chromosomes，r oot tips were put in Chromosome numbers based on x=36，w hich 0.002 M 8-hydroxyquinoline for 4h ours，a nd was reported by Kurita (1963)，Ts ai and Shieh fixed in 45% acetic acid for 10 minutes. The (1983) and Weng (1985)，w ere not encoun- fixed roots were macerated with lNH Cl for 1 tered during the course of this study. minute at 60oC，a nd squashed in 29もaceto- Occurrences of aneuploids in ace rtain spe- orcein solution. The number of spores per cies have been often confirmed in various -119- 120 植物研究雑誌第73巻第3号平成10年6月 Table 1. Chromosome numbers of Thelypteris laxa (Franch. &S av.) Ching Chromosome number /Pl oidy and basic chromosome number presumed Locality /Re ference or voucher specimen (Nakato's No.) Previous study n=72，te traploid of x=36 Kikugawa，S hizuoka Pref. (Kurita 1963) n=36IV，te traploid of x=36 Lienhuachih，Na ntou，Ta iwan (Tsai and Shieh 1983，19 85) n=36，di ploid of x=36* Hangzhou，Zh ejiang，C hina** (Weng 1985) 2n= 136，te traploid of x=34 Tsukubasan，Ib araki Pref. (Nakato 1996) Present study 2n=136，te traploid of x=34 Sayama，Hi gashiyamatoshi，Tokyo Pref.，no . 2081 ** ・ Umegaya-toge，Om e-shi，T okyo Pre ，.fnos. 2142**，21 43 2n= 134，hy potetraploid of x=34 Hagurosan，H aguromachi，Yamagata Pref.，no . 2131 ** 田 Mizuho-machi，Ni shitama-gun，T okyo Pref.，no . 2097 Hinode-machi，Ni shitama-gun，T okyo Pref.，no . 1976 Hinohara-mura，Ni shitamルgun，Tokyo Pre ，.fnos. 1978，1 979** Takao-san，Ha chioji-shi，T okyo Pref.，no s. 2068，20 69 Hatsukari-machi，Ot suki-shi，Y amanashi Pre ，.fno. 2216 Tatsuruhama-machi，Ka shima-gun，Is hikawa Pre ，.fno. 1932** Shika-machi，Ha kui-gun，Is hikawa Pref.，no . 2226 Yugashima，Am agiyugashima-machi，Sh izuoka Pre ，.fnos. 2035*穴2036 Funabara，Am agiyugashima-machi，Sh izuoka Pre ，.fnos. 2062料，2063 Yadokoro，Ow ase-shi，M ie Pref.，no . 2125 Ouga，Sh ingu-shi，W akayama Pref.，no . 2105 Hikimi-cho，Mi no-gun，Sh imane Pre ，.fno. 2149料 Mochidagaura，Ha kata-ku，Fu kuoka-shi，Fu kuoka Pref.，no . 2113 Ikeda，Sa wara-ku，Fu kuoka-shi，Fu kuoka Pre ，.fno. 2127 Kumano，Bu ngotakada-shi，Oi ta Pref.，no . 2218 Fukata，Us uki-shi，Oi ta Pref.，no . 2222 * Octaploid (x=9) in the original paper. But n=36 should be assigned to diploid，be cause species with n=9 or 18 have not been reported in this genus. Specimensexamined for the number of spores per sporangium. All these have 64 good 料 spores m as porangmm. families and genera in Japanese ferns: Isoetes stricted areas. Ont he contrary，in T. laxa，th e japonica A.Br. and I. sinensis T.C. Palmer in hypotetraploids (2n= 134) seem to have aw ider Isoetaceae (Takamiya et al. 1994)，Le pisorus distribution range than the eutetraploid thunbergianus (Kaulf.) Ching in the (2n=136). The hypotetraploids may have ad- Polypodiaceae (Nakato et a .l1983)，P teris vantageous attributes that caused these exten- cretica L. in Pteridaceae (Nakato 1989)，et c. In sive distribution beyond the eutetraploids. these species，a neuploids have been found Comparison 01 chromosome length between rarely in euploid populations and/or in re- the two cytoηpes Although strict karyo- June 1998 Journal of Japanese Botany Vo .173 No. 3 121 Figs. 1，2. Somatic chromosomes of Thelypteris laxa. 1. 2n=136，e utetraploid，no . 2142. 2. 2n=134， hypotetraploid，no . 1978. Scale bar =5 μm. logical analysis was difficult from nuclear mosome number in the hypotetraploid might plates obtained in this study，th e sizes of chro- not arise from simple loss but from fusion of mosomes could be measured and comped chromosomes. 訂 between those of the eutetraploid and the Reproductive system The spore numbers per hypotetraploid. Figures 3a nd 4s how somatic sporangium could be counted in 2p lants of the chromosome complements arranged by tetraploid and 6p lants of the hypotetraploid. length，ea ch corresponding to those of Figs. 1 In all of these specimens，64 good spores were and 2，re spectively. counted in each sporangium，i ndicating that In the eutetraploid of no. 2142 (Fig. 3)，th e they are sexual in reproduction (Figs. 5& 6). variation in chromosome length was gradual; This finding is in accordance with Momose the chromosomes range in length from 1.9μm (1967) from the studies of prothallia. to 0.9μm with the mean of 1.3μm; and the Cytotaxonomy 01 T. laxa and related species longest chromosome was about 2.1 times as Iwatsuki (1995) classified the genus Thelypteris long as the shortest. On the contrary，in the (37 spp.) of Japan into eight subgenera and hypotetraploid of no. 1978 (Fig. 4)，t he first eight sections and placed T. laxa in subgenus two chromosomes were much longer than the Thelypteris，se ction Metathelypteris，to gether others，be ing 2.4μmi n length; the remaining with T. gracilescens (Blume) Ching，T . chromosomes showed ag radual decrease in uraiensis (Rosenst.) Ching and T. hαttorii (H. length from 1.8μmtoO.9μm; the mean length Ito) Tagawa. This section is referable to the of all chromosomes was 1.2μm; and the length genus Metathelypteris recognized by Ching of the longest chromosomes was about 2.4 (1963). The diagnostic characters of this group times that of the shortest. The two larger chro- are: free veins，in dusiate sori，ab sence of aero- mosomes were also observed in the comple- phore，ve inlets that do not reach the margin of ments of other hypotetraploids. These results segments，a nd bipinnatifid to tripinnatifid suggest the possibility that reduction of chro- fronds. Other Asian species allied to T. laxa 122 植物研究雑誌第73巻第3号平成10年6月 S 都議襲撃襲撃襲撃襲撃護審議事事事事議議事襲撃議融事襲撃事職事事審議毒事馨事番難審議亀韓併事農事馨事議議事撃事輔韓議著書轟襲撃事襲撃韓議事事襲撃事事事韓議事著書襲撃事事轟議事襲撃事事事襲撃襲撃事事 3 特輯襲撃事事時鱒襲駒輔輸事議事馨綿襲撃幹馨襲撃轟議 6 議事事事事襲撃事事告書意義襲撃襲撃事事事事事事事事護憲講義襲撃韓議事事事事事事襲撃事事事事事事襲撃事事事事事審議事事事腕襲撃事事 E 1 踏襲撃講義講義鞠撃事時鵠持議審議事師事車場事場事事事輔 4 1 絹綿綿綿幹事事襲撃鵠鵠講義襲撃事事襲撃事鵜綿払 Figs. 3，4. Somatic chromosome complement arranged by length. 3. 2n=136 chromosomes from Fig. 1，n o. 2142.4. 2n==134 chromosomes from Fig. 2，no . 1978. Scale bar =5 μm. Figs. 5，6 . Sixty-four spores in one sporangium. 5. Eutetraploid，no . 2142. 6. Hypotetraploid，no . 2131. Scale bar =1 00μm. June 1998 Journal of Japanese Botany Vol. 73 No. 3 123 are T. adscendens Ching，T. decipience Ching， Phytotax. Sin. 8: 289-335. T. flaccida Ching and T. singalanensis Ching Hirabayashi H. 1969. Chromosome numbers in several species of Aspidiaceae. 1. Jpn. Bot. 44: 113-119. (Ching 196 ，3，Iwatsuki 1965). Iwatsuki K. 1965. Taxonomy ofthe Thelypteroid ferns， Among these species，t he following four with special reference to the species of Japan and species have been examined cytologically: T. a司jacentregions. IV. Enumeration of the species of flaccidα(n=35 and n=70，M anton and Sledge Japan and adjacentregions. Mem. Col .1Univ. Kyoto， B，31 : 125-197. 1954)，T. singalαnensis (n=70，Ma nton 1954)，一一一一一 1995.Thelypteridaceae. In: Iwatsuki K.， T. uraiensis (n=62，Hi rabayashi 1969; n=36IV， Yamazaki T.，Bu ffordD.E. andOhbaH. (eds.)，Flora Tsai & Shieh，1 983) and工laxa(see Table 1). of Japan，V o .11，pp . 174-194. Kodansya，To kyo. From these data，it is assumed that the section Kurita S. 1963. Cytotaxonomical studies on some Methathelypteris sensu 1wa tsuki (1995) (or leptosporangiate ferns. 1. Coll. Arts Sci. Chiba Univ. 4: 43-52. the genus Metathelypteris sensu Ching，19 63) Manton I. 1954. Cytological notes on one hundred spe- is characterized by polybasic chromosome cies of Malayan ferns. In: Holttum R. E.，A R evised numbers such as x=31，3 4，3 5 and 36. These Flora of Malaya，V o .1H，pp . 623-627. Gov. Print. clearly show that aneuploidy as well as poly- Office，Si ngapore. 一一一一-and Sledge，W . A. 1954. Observations on the ploidy has played an important role in the cytology and taxonomy of the Pteridophyte flora of evolution of this group. Ceylon. Philos. Trans. Roy. Soc. London，B ，2 38: As statedearlier，T. laxais distributed widely 127-185. in warm temperate areas ofE. Asia. However， Momose S. 1967. Prothallia of the Japanese ferns data available from outside Jap an are only (Filicales). Univ. Tokyo Press，T okyo. Nakato N. 1989. Distributions and mo中hologicalvaria- those recorded by Tsai and Shieh (1983，19 85) tions ofdiploid and triploidcytotypes of Pteris cretica and Weng (1985)，e ach based on as ingle L. in the Kanto District. J. Phytogeogr. Taxon. 37: material from Taiwan and China，re spectively， 113-119. and reporting the base number of x=36. It is 一一一一 1996. Notes on chromosomes of Japanese pteridophytes (4). 1. Jpn. Bot. 71: 163-167. noted that Tsai and Shieh (1983，1 985) re 同一一一-and Masuyama S. and Mitui K. 1983. Studies ported the number of n=36IV (36 quad- on intraspecific polyploids of the fern Lepisorus rivalents) at meiosis with the normal produc thunbergiαnus (1). Their distributional patterns in 向 tion of 64 spores per sporangium. Extensive Kanto districts and the occuenceof new cytotypes. 町 chromosomal studies，es pecially from outside 1. Jpn. Bot. 58: 195-205. Takamiya M. 1996. Index to chromosomes of Japanese Jap an，ar e needed to understand the cytologi 町 Pteridophyta (1910-1996). Japan Pteridological So- cal evolution in this species. ciety，T okyo. 一一一一，Watanabe M. and Ono K. 1994. Biosystematic 1t hank Dr. D. M. Britton，U niversity of studies on the genus Isoetes in Jap an I. Variations of the somatic chromosome numbers. J. Plant Res. 107: Guelph，fo r his kind linguistic review of the 289-297. manuscript. 1a lso thank Mr. Koichi Ohora Tsai 1.-L. and Shieh W.-C. 1983. A cytotaxonomic (WakayamaPref.)， Mr. Takeshi Suzuki (Hyogo survey of pteridophytes in Taiwan. 1. Sci. Engin. 20: Pref.) and Mr. Sadao Tsutsui (Fukuoka Pref.)， 137-158. 一一一-and一一一一， 1985. A cytotaxonomic survey of for supplying some materials. the fern family Aspidiaceae (sensu Copeland) in Taiwan.1. Sci. Engin. 22: 121-144. References Weng R.-F. 1985. Observation on chromosomes of Ching R.-C. 1963. A rec1assification of the family some Chinese ferns. J. Wuhan Bot. Res. 3: 367-370. Thelypteridaceae from the mainland of Asia. Acta 124 植物研究雑誌第73巻第3号平成10年6月中藤成実:ヤワラシダの異数性ヤワラシダにはx=34の4倍イ本，および、x=36の2 体は正4倍体から単純に2本の染色体が欠失したも倍体と 4倍体が報告されている.今回，日本の 19 のではなく，正4倍体の染色体が融合したために生産地から採取した24個体について染色体を観察しじたのではないかと推察された.岩槻 (1995)はたところ， 2産地からの3個体は2n=136，17産地ヤワラシダを， Thelypteris亜属Metathelypteris節にからの21個体は2n=134であった.前者はx=34の分類した.この仲間の染色体数は現在までヤワラ正4倍体，後者は異数体で低4倍体と考えられる. シダを含め4種で調べられており，染色体基本数に本研究においてはx=36に基づく染色体数は観察でついてx=31，34， 35， 36，倍数性では2倍体と4倍きなかった.正4倍体と低4倍体の染色体の長さを体が知られている.これはこの仲間の細胞学的測定したところ，正4倍体では136の染色体長は漸進化には倍数性ばかりでなく異数性が重要な役割次変化するが，低4倍体では134の染色体のうち2 を演じたことを示している. 本の染色体が特に長かった.このことから，低4倍 (東京都立国分寺高等学校) 第73巻2号正誤 Erratain Vol. 73 No. 2 頁(Page) 行(Line) 誤(For) 正(Read) 104 trilobed or tri10bed to 106 ↓ 2 var. himalaica Hara var. himalaica H.Hara 108 ↑ 4 (Hayata ex Honda) (Hayata) 108 ↑ 6 Hayata ex Honda Hayata in Honda 108 Hayata ex Honda Hayata 109 ↓22 Hayata ex Honda Hayata 110 ↓10 Nakai ex Honda Nakai in Honda 111 ↑ 6 (Na kai) Hara (Nakai) H.Hara 111 ↑16 (Nakai) ex Honda Nakai in Honda 111 ↑21 Thunb.8)e x Murray Thunb.8)i n Murray 112 ↑24 Nakai ex Honda Nakai 113 ↑24 Heloniopsis Heleniopsis 114 ヤマシマショウジョウノてカマヤクシマショウジョウノくカマ