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A Revision of Cardamine loxostemonoides O. E. Schulz (Cruciferae) PDF

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植物研究雑誌 J. Jpn. Bot. 68: 199-208 (1993) AR evision of Cardamine loxostemonoides O. E. Schulz (Cruciferae) Abdur RASHIDa,ba nd Hideaki OHBAa aDepartment of Botany,Un iversity Museum,Un iversity of Tokyo,7- 3-1 Hongo,T okyo 113 JAPAN; bDepartment ofBotany,Un iversity ofPeshawar,Pe shawar,P AKISTAN Cardamine loxostemonoides (アブラナ科)の分類学的研究 A.ラシッドa,b,大場秀章a a東京大学総合研究資料館生物系研究室 113東京都文京区本郷7-3-1 bノfキスタン ペシャワール大学植物学教室 (Received on Janu訂y9,1 993) Cardamine loxostemonoides O. E. Schulz is revised. Based on the discussion on gross mo中hology,it is clarified that the so called C.loxostemonoides can be distinguished into three speci白;C.loxostemonoides sensu stricto,C. luxuriαns and C. tibetana. Chromosome number for C.luxurians and C.loxostemonoides is reported forthe first time. Cardamine tibetα,na,fr om S. E. Tibet is described in this paper; an ew combination C.luxurians is proposed. Schulz (1927) described Cardamine loxo- RAWa nd TI were used for the study. Fifteen quanti- stemonoides in the section Macroca中ousbased on tative characters were rneasured in eachindividual in Duthie's collection in the 1880's from N. W. India addition to the qualitative characters (Table 1). The and W. Nepal. Jafri (1973),H ara (1979) and Polunin number ofleaflets were counted in the second cauline and Stainton (1989) consider this as ad istinct species leaves from the base of each individual for the char- ranging from Pakistan to Bhutan in elevations 3000 to acter 3 and terminal leaflet of the same leaf was 5000 m (Fig. 1). Lan and Cheo (1981) transferred it to measured for the characters 5a nd 6. Minimum,ma xi- the genus Loxostemon. mum,a verage and standard deviation for each vari- Regarding our study of the genus Cαrdα,mine and able was calculated. For the qualitative characters other allied genera recently weh ave realized that C. sorne terms such as coral bulbils,g lobular bulbils, loxostemonoides has not been critical1ye xarnined. reduced leafy bulbils,di gitate appendages and cres- The present study aims to clarify vegetative and cent scaly leaves were used based on morphological reproductive features and to provide chrornosorne appearances. numbers for the taxonomic revision. Chromosome counts were rnade from the root tips. The rnaterials were col1ected from Pakistan and Ne pal Materials and methods and treated in 0.1 % colchicine for 2h ours; fixed in Herbarium specimens preserved in BM,E ,K , Newcomer's solution; rnacerated in 1N HCl for 10 -199- 200 植物研究雑誌第68巻第4号 平成5年8月 二二〉 Fig. 1. Distribution map of C. luxurians (圃),C. loxostemonoides (企)and C. tibetana (・). minutes at 600C and squashed after staining for 2 composed of fleshy scaly leaves (Figs. 2,3). The scaly hours in 2% aceto-orcein. The voucher specimens leaves are somewhat whitish in co.lour and fleshy at (Ohbaetal. no. 393 and Miyamoto etal. no.147) were the base tapering into apical digitate appendages. The deposited in TI and PUH. specimens collected in S. E. Tibet have delicate rhi- zome and both coral and reduced leafy bulbils瓜the Observations axils of stolons (Figs. 3-6). Thec oral bulbils are 1. Morphology composed of crescent shaped scaly leaves which are Cardamine loxostemonoides,g rowing in moist aangedon the reduced axis ending into apical bud. 打 habitats among the loose boulders,ro ck crevices and Some of the crescent scaly leaves also bear axillary screes,s hows great variations in rhizome morphol- bulbils. Reduced leafty bulbils ecomposed of re- 訂 ogy,pl ant height,n umber of cauline leaves,n umber duced leaves. They eclose to the soil surface and 訂 of leaflets,le aflet size,pu bescence,n umber of flow- usually terminating into aerial shoots. ers per raceme,a nd shape of petals among different b. Cauline Leaves: The Pakistan plants have 5-11 localities (Table 1). petiolulate,s uborbicular-oblong,l obulate and gla- a. Rhizome: The plants from Pakistan show vigor brous leaflets. Those from Ne pal have 7-13,pe tiolulate 同 ous growth of stolons but without any bulbils. The or sessile,o bovate,s uborbicular or linear oblong, stolons produce secondary stolons from the axils of lobulate and glabrous to hairy leaflets. Those from the membranous scaly and the radical leaves. The S. E. Tibet are 7-9 sessile,ob ovate leaflets with hairy scaly leaves are papery with wide base and tapering margms. mto acute to acuminate apex. c. Flowers: Theo vules訂eanatropous,v alves The plants from Nepal have slender rhizomes and veinless,p lacentae uniform,f unicles filiform and delicate aerial stem. The rhizome bears globular bulbils septa nerveless. Generally C. loxostemonoides has a August 1993 Journal of Japanese Botany Vol. 68 No. 4 201 Table 1. Mean,st andard deviation and range for quantitative characters in the three species. c. luxurians C. tibetana C.loxostemonoides 1. Plant height (cm) 11-25 8-12 5-17 (17.2:t4.07) (9.8:t1.63) (10.3:t3.48) 2. Number of cauline 3-5 2-4 1-3 leaves (4:t0.77) (3:t0.89) (2.25土0.62) 3. Number of leaf1ets 5-11 7-9 7-13 (8.4:tl.90) (8.2:t0.97) (9.2:t1.66) 4. Number of f10wers ι20 2-6 1-12 (11.6:t3.89) (4.4:t1.35) (5.9:t3.16) 5. Length of terminal 13-16 5-8 5-11 leaf1et of 2nd (15.07士0.92) (6.3士1.07) (8.4:t2.45) cauline leaf (mm) 6. Width of terminal 7-12 3-6 1.5-5 leaf1et of 2nd (8.68:t1.23) (4士1.09) (2.93:t0.94) cauline leaf (mm) 7. Length of sepals (mm) 3-3.6 3.4-3.6 2.5-3.5 (3.29:tO.17) (3.48:t0.07) (2.94:t0.28) 8. Width of sepals (mm) 1.5-2.1 1.5-1.8 1.5-2 (1. 87:t0. 17) (1.68土0.11) (1.76:tO.18) 9. Length ofpetals (mm) 10-12.2 10-12.1 10-12 (11.51:tO.68) (10.82土0.78) (10.82:t0.66) 10. Width of petals (mm) 5-6.1 8.3-8.5 5-6.6 (5.78士0.38) (8.4土0.06) (5.87:t0.43) 11. Length of short 5-5.5 6ル6.2 5-5.4 stamens (mm) (5.24:t0.15) (6.08土0.07) (5.21土0.12) 12. Length of short 0.9-1.2 1.1-1.2 1-1.2 stamen's anther (mm) (1.02:t0.07) (1.16:t0.04) (1. 12:t0.07) 13. Length of long 6.8-7 6.仏6.3 6-6.5 stamen (mm) (6.93:tO.07) (6. 16:t0.10) (6.18土0.19) 14. Length oflong 0.9-1.2 1.0-1.2 0.9-1.2 stamen's anther (mm) (1.0l:t0.07) (1.12:t0.07) (1.06:t0.l0) 15. Length of pistil (mm) 5.2-6.2 6.3-6.5 5.8-6.1 (5.9:t0.27) (6.42:tO.07) (5.95士0.09) racemose inflorescence. The number of flowers per in central Nepal (Figs. 11,12 ). inflorescence is different among localities (Table 1). The flowers from Pakistan have ovate sepals with Taxonomic comments and discussion visible veins,s pathulate petals,a nd 16-20 ovules Schulz (1903) defined one monotypic section (Fig. 8). Those from Nepal have ovate sepals with Macrocarpous from Estrecho de Magallanes,S. Chile prominent veins,ob ovate petals and with 14-20 ovules based on C. geraniifoliα(Poiret) DC. Schulz (1927) (Fig. 9). Those from Tibet are ovate sepals with described C. loxostemonoides from Himalaya and obscure veins; widely obovate petals and 14-18 ovules then placed it in the section Macrocarpous (Schulz (Fig. 10). 1936). Both the species are geographically quite f.訂 2. Chromosome number from one another in addition to different morphologi- Thirty two (2n=32) somatic chromosome number cal characters. The present study does not support the was observed in 10 individuals from two populations homogenous nature of the section Macrocarpous and in N. Pakistan and 6i ndividuals from one population is difficult to consider them as am onophyletic one. 202 植物研究雑誌第68巻第4号 平成5年8月 Figs.2-3. C.loxostemonoides,2. rhizome,Ba r: 1m m,3. globul紅 bulbil,Bar: 0.5 mm. Figs. 4ー7.C. tibetana,4. rhizome,Ba r: 1m m,5. young coral bulbil,Ba r: 0.5 mm,6. coral bulbil,Ba r: 0.5 mm,7. reduced leaves at the base of stem,Ba r: 1m m,st: stem,pt: petiole of radicalleaf,sc : fleshy scaly leaf,dt : digitate apex of the fleshy scaly leaf,cr : crecent shaped scaly leaf. Moreover,t he present finding reveals some more α!ppendiculata weh ave observed growth of plantlets reliable characters such as types of bulbils,t exture from the leaflets and petioles. Lifeform accompanied and apex of scaly leaves which can easily be observed by rhizomes,s tolons and bulbils are found widely even in the dry herbarium specimens. throughout in the genus Cαrdamine for regeneration. Eighty five percent species of Caraαmine,a nd Hooker and Thomson (1861) established the the species of Loxostemon are perennial and repro- genus Loxostemon by the winged long filaments bent duce vegetatively from rhizome. Cardaminep ratensis outside (reflexed) in the upper part. Cardamine reproduces from foliage leaves (Lovkvist 1956). In C. loxostemonoides differs from Loxostemon in having August 1993 Joumal of Japanese Botany Vol. 68 No. 4 203 8 c 9 命 川 W c 10 ハ U c Fig.8. Cardamine luxurians,F ig. 9. C. loxostemonoides,F ig. 10. C. tibetana,a: sepal,b: petal,c: short and long stamens,d: pistil. Bar: 1m m. 204 植物研究雑誌第68巻第4号 平成5年8月 亡3 12 Fig. 11. Somatic chromosome number in C.loxostemonoides (2n=32). Bar: 10μm;Fig. 12. Il1ustration of fig. 11. wingless erect long filaments but possessing the fila- loxostemonoides) is characteristic in globular bulbils ments slightly curved towards the stigma,wh ich e and the Tibetan one (C. tibetana) is delimited by 訂 in coincidence with the genus Cardamine. possessing coral and leafy bulbils. The later two have Other characters used for the delimitation of genus smallleaflets. Loxostemon such as few flowers,f ew leaves,f ew Usually the genus Cardamine possesses the basic leaflets and weak stem (Hooker and Thomson 1861) number 8,t herefore C. luxurians (2n=32) and C. enot reliable and inconsistent. For example,C . loxostemonoides (2n=32) are considered as tetraploid. 訂 loxostemonoides,is 5-25 cmt all and has 1-5 cauline This is also the first record of chromosome counts leaves,5- 131eaflets and 1-20 flowers. Therefore,it from Himalayan region in the genus Cardamine. must be treated under the genus Cardamine as no evidence exists to put it under the genus Loxostemon. Taxonomic treatment Cardamine and Loxostemon are unique in the Key to the species Cruciferae in having the characters of explosive fruit 1. Bulbils absent; scaly leaves membranous with with marginate narrow replum and elastic de- acute to acuminate apex; leaflets suborbicular, hiscence by spiral coiling of the valves. However, glabrous,la rge; petals spathulate.…… Loxostemon is distinguished from Cardamine by the ..1. C. luxurians triangular obovate filaments of the long stamens. 1. Bulbils present; scaly leaves,f leshy,c rescent Cardamine loxostemonoides varies in rhizome shaped or fleshy with digitate apex; leaflets bulbils morphology. Reduction in the main axis of suborbicular-obovate,li near-oblong,gl abrous to rhizome and change in the texture and shape of scaly hairy; petals wide obovate,o bovate-oblong leaves have resulted in variation of mo中hology. 2. Globular bulbils present and leafy bulbils ab- Cardamine loxostemonoides in the Himalayan sent; scaly leaves digitate at the apex; reduced region exhibits different characters and can be di- leaves absent at the base of the stem,pe tals vided into three species. The Pakistan species (c. obovate oblong …・・...2.C. loxostemonoides H H luxurians) is characterised by vigorous stoloniferous 2. Coral and leafy bulbils present; scaly leaves rhizome without bulbils; the Nepal one (C. crescent shaped; reduced leaves present at the August 1993 Journal of Japanese Botany Vol. 68 No. 4 205 base of stem; petals widely obovate ....・ ・.. 2. Cardamine loxostemonoides O. E. Schulz in H ・... ・..……………...・ ・.....・ ・・・...3.C. tibetαnα Notizbl. Bot. Gard. Berlin 9: 1069 (1927); Jafri in Fl. H H H H H , Pak. 55: 168 (1973); Hara in Enum. Flow. Pl. Nepal 1. Cardamine luxurians (0. E. Schulz) Rashid 2: 40 (1979); Polunin and Stainton,Fl ow. Himal. 41 et H. Ohba,co mb. et stat. nov. (1989). Cardamine loxostemonoides O. E. Schulz var. Loxostemon loxostemonoides (0. E. Schulz) Y. C. luxuriansO. E. SchulzinNotizbl. Bot. Gard. Berlin9: Lan et T. Y. Cheo in Bull. Bot. Res. N. E. Forest. lnst. 1070 (1927). 1: 54 (1981); T. Y. Cheo inFl. Reip. Pop. Sin. 33: 233 Plants perennial with branched stoloniferous rhi- (1987). zome,bu lbils absent,sc aly leaves membranaceous, Plants perennial with stoloniferous branched rhi- apex acute to acuminate somewhat greyish in colour. zome,bu lbils present,sc aly leaves f1eshy with apical Stem suberect to procumbent,1 1-25 cm long,gl a- digitate appendages somewhat white in colour; stem brous. Radical leaves present; cauline leaves 3-5, suberect or procumbent,5 t o 17 cm long,gl abrous, long petiolate,pi nnatisect,us ually leaf1ets 5-11; leaf- sometimes hairy in the upper portion. Radicalleaves lets petiolulate,s uborbicular,m argins entire to present; cauline leaves 1-3,l eaves long petiolate, trilobulate,13 -16 mmlo ng,7 -12mmb road; terminal pinnatisect,us ually leaf1ets 7-13,va riable in size and one slightly biggerthan the lateralleaf1ets. Raceme 6- shape; leaf1ets petiolulate or expetiolulate,su borbicular 20 f1owered. Flowers 9-14 mma cross,s howy,pu r- to linear oblong,5 -11 mml ong,1. 5-5 mmb road, ple; pedicels 11-19 mmlo ng; sepals 3-3.6 mml ong, terminallobe scarcely differentiated from the lateral 1.5-2.1 mmbr oad; petals 4t imes longer than sepals, ones,en tire to lobulate,ma rgins hairy. Raceme 1-12 10-12.2 mml ong,5 -6.1 mmb road,v enation dark f1owered. Flowers 8-14 mma cross,s howy,pi nk or violet; glands 4i. e. 2r ing shaped at the bases of short pu叩le;pedicels 10-20 mmlo ng; sepals 2.5-3.5 mm stamens,2 l ateral glands at the dorsal side of long long,1 .5-2 mmb road; petals 4t imes as many as stamens; long stamens 6.8-7 mml ong,sh ort stamens sepals; 10-12 mml ong,5- 6.6 mmb road,as cending, 0.9-1.2 mmlo ng. Anther dehiscence introrse,al ong obovateoblong,ha rdly clawed,da rk violetly veined; ‘ the longitudinal slits. Siliques linear compressed 2.5- long stamens 5-5.4 mml ong,s hort stamens 6-6.5 3.3 mmlo ng,1. 5 mmbr oad,st raight,gl abrous,se ptum mmlo ng; anther dehiscence introrse along the longi 田 and valves veinless; ovules 16-20,pl acenta uniform, tudinal slits; Pisti15 .8-6.1 mm,ov ules 16-20,se ptum funicle filiform. Pollen grains mostly elliptic to spheri- and valves veinless,pl acenta uniform,fu nicle filiform. cal,1 5.6-30μm long,1 5-17.5μm broad,is opolar, Siliques linear oompressed 2.4-3.2 mml ong,1 .4 mm radiosymmetrical,tr icolpate,su rface sculpturing re- broad,st raight,gl abrous. Pollens 17.5-39.4μmlong, ticulate. Somatic chromosome number 2n=32. 16.9-21.9μmb road,mo stly elliptic,is opolar,ra dio- Holotype: N. W. lndia (Tihri-Garwal): Nila Val- symmetrical,t ricolpate,s urface sculpturing reticu- ley,1 6 August,1 883 (Duthie 912,pr obably DD,no t late. Somatic chromosome number 2n=32. seen,ph oto in K; isotype in K). Lectotype: N. W. lndia (Kunawer): Nipchang val- Other specimens examined: Pakistan: Northem ley alt. 4300-4600 m (Duthie 2724,31 August 1884, Areas,D istrict Gilgit,A store,A bove Rama Lake, K,t he lectotype is here selected; isotype in ?DD). (Webster & Nasir 6450,1 2 August 1955,R AW); Syntypes: Lebung Pass (Duthie 5330,1 A ugust 1886); RamaL ake,(H . Ohba et a ,.l393,20 August 1992,TI , Tihri-Garwal: Chinal opposite Bandrapunch (Duthie PUH). 911,9 13,2 5 August 1883); West Nepal: Opposite 206 植物研究雑誌第68巻第4号 平成5年8月 Budhi village (Duthie 5330,18 July 1886); Nampha 1973,K ); Thumu Khola,K humbu (Bowes Lyon Gadh,(D uthie 5330,2 7 July 1886),al l probably in 2211,2 0 June 1964,B M); Hongu Khola,le ft bank DD,no t seen. near Mera Kharka (Mc Cosh 360,3 J uly 1964,BM ). Specimens examined: W. Nepal: Dozam Khola ne紅白mikot(Polunin,S ykes &W illiams 4284,1 3. Cardamine tibetana Rashid et H. Ohba,sp . June 1952,B M,TI ); near Pudamigaon,ne ar Suli Gola nov. (Fig. 13) (Polunin,Sy kes & Williams 2253,19 June 1952,TI ); Cardamine loxostemonoide O. E. Schulz similis Padmara Lagna (Polunin,S ykes & Williams 4372, sed bulbilos coralliforme et illos foliaceos et folia June 1952,B M,TI ); Phoksumdo Tal (Stainton 4296 rudimentalia habens. 12 June 1963,B M,TI ); Phoksumdo Khola (Stainton Plants perennial with stolonoiferous rhizome,tw o 4316,1 7 June 1963,E ); Porakya Lagna,K amali types bulbils present i. e. cor叫 bulbilsand leafty (Stainton 4878,23 May 1965,B M,TI ); Tingjegaon bulbils; scaly leaves fleshy crescent shaped,sp irally (Polunin,Sy kes &W illiams 1155,19 June 1952,B M, arranged,so mewhat whitish in colour. Stem suberect TI); BalangraPass (Polunin,Sy kes &W illiams 1043, to erect,8 -12 cm long,g labrous. Radical leaves 30 May 1952,TI ); Sisne Himal (Stainton 6342,2 7 present; cauline leaves long petiolate 2-4, leaflets June 1968,B M,TI ); Sisne Himal (Stainton 6347,29 expetiolulate obovate-oblong with entire m gins, 訂 June 1968,B M,T I); Tarap-she (Dolpa District) hairy usually along the margins; 5-8 mml ong,3- 6 (Yoshida 1035,9July 1991,TI) ; Chaurpmi(Dobremez mmb road,al l the leaflets in the same leaf uniform. 2301,1 7 May 1973,B M); 3.5 miles E of Saipal Raceme panicle,2 -6 flowers. Flowers 8-11 mm (Amold 306,31 August 1954,BM ); 5m iles,N. E. of across,s howy,vi olet; sepals 3.4--3.6 mml ong,1. 5- Saipal (Amold 233 & 230,2 5 August 1954,B M); 1.8m mb road,ve nation obscure; petals 10-12.1 mm Chhairogaon,N . of Tukucha (Stainton,S ykes & long,8. 3-8.5 mmbr oad,vi olet,wi dely obovate; glands Williams 859,1J une 1954,BM ); Sialgarki (Polunin, 4,i. e. 2r ing shaped glands at the base of the short Sykes & Williams 991,2 2 May 1952,B M); Jangla stamens,21 ateral glands on dorsal side of the bases of Banyang (Einsson,Skarby & Wetterhall 838,1 7 long stamens; short stamens 2; long stamens 4; short 訂 June 1963,BM ); C. Nepal: 3m iles S. E. of百lOrong stamens 6-6.2 mml ong; long stamens 6-6.3 mm pass (Epstein 5,2 5 June 1977,B M); Khung Khola long; anther dehiscence introrse,lo ngitudinally along (Gray-Wilson & Philips 646,1 6 August 1973,K) ; the slits; pistil 6.3-6.5 mmlo ng,st igma flat,sl ightly Langtang (Polunin 188,8 J une 1949,BM ); Langtang bifid; ovules 14ー20,sep佃m and valves veinless, valley,fa cingN. across river (Polunin 654,June 1949, placenta uniform,fu nicle filiform; siliques linear com- BM); Langtang valley (Miyamoto et al. 147,A ug. pressed,gl abrous. Pollen grains 19.4--27.5μm long, 1992,TI ); Tinkar Khola (Tyson 60,2 4 June 1953, 16.3-22.5μm broad,s pherical,i sopolar,r adio- BM); Nampa Khola,(T yson 37,14 June 1953,BM ); symmetrical,tr icolpate,sp herical to elliptical in shape, N. E. of Chalike Pakar (Stainton Sykes & Williams surface sculpturing reticulate. 3115,14 June 1954,B 加1);Jargeng Khola (Lowndes Type: S. E. Tibet,T se La,L angong,9 4000'N. 1045,21 June & 2J uly 1950,B 民I[);E. Nepal: Arum 28045 'E,alt . 14--15000 ft (Ludlow and Sheπiff5618, valley,Pi ling Khola,N. E. of Chyamtang (Stainton 21 June 1938,ho lotype,TI : isotype,BM ). 406,23 May 1956,K) ; east oflower Sangdah (Grey- Wilson & Philips 411,2 7 July 1973,K ); north of Wea re thankful to the directors and curators of Chung See (Grey-Wilson & Phillips 613,12 August BM,E,Kf ortheloanofspecimens; Dr. F. Miyamoto, August 1993 Journal of Japanese Botany Vol. 68 No. 4 207 Fig. 13. Cardamine tibetana Rashid et H. Ohba (Ludlow and Sherriff 5618,Ty pe). Tokyo University of Agriculture,fo r collection of (Field Research),No . 04041038 in 1992 to H. O. root tips for this study,Dr . A. R. Beg and Dr. F. References Hussain,Un iversity of Peshawar for help in the field. (Other references are in the taxonomic treatment) This study was supported by ag rant from the Lovkvist B. 1956. The Cardamine pratensis complex,ou tlines ofits cytogenetics and taxonomy. Symb. Bot. Upsal.14(2): Monbusho Intemational Scientific Research Program 208 植物研究雑誌第68巻第4号 平成5年8月 1-131. 数はパキスタンとネノマール産で2n=32であった. Hooker J. D. and Thomson M.D . 1861. Praecursores ad Floram 形態学的形質にはノマキスタン(cid:631)北西インド(クマ Indicam. Jour. Linn. Soc. 5: 128-181. Schulz O. E. 1936. Cruciferae. In Engler A. and Prantle K. A. E., オン), ネパール, 南東チベットの間で不連続な Die naturlichen Pflanzenfamilien,2 nd ed.,17 b: 227-658. 差異が見いだされた.それらは地理的にもまと まっており,独立の種として認識するのがよい. 要旨 なお,周太炎と藍永珍 (1981)はこの種をLox- Cardamine loxostemonoidesはヒマラヤ高山帯に ostemon属に移した. 同属の種とC.loxostemono- 広く分布するがまれな植物で, これまで標本も少 idesは花の構造が違う.Loxostemon属は成熟す なし不明の点が多かった.ネパールとパキスタ るとらせん状に巻く果皮をもつことで,タネツケ ンで現地観察をし,可能な限り標本を借用して, バナ属に類似するが,長雄蕊は逆三角状卵形の花 形態学的形質と染色体の観察を行なった.染色体 糸をもつことで異なる.

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