A Revision of Anemone L. (Ranunculaceae) from the Southern Hemisphere

2006·2.4 MB·English

by  Svetlana N. Ziman

植物研究雑誌
J. Jpn. Bo .t
Originals 81: 193-224(2006)
AR evision of Anemone L. (Ranunculaceae) from
the Southern Hemisphere
Svetlana N. ZIMAN ，aCarl S. KEENERb，Yu ichi KADOTA ，C
Elena V. BULAKHa and Olga N. TSARENKOa
aN. G. Kholodny Institute of Botany，Na tional Academy of Sciences，
2，Te reshchenkivska street，Ki ev，01 601 UKRAINE;
E-mail: [email protected]
b208恥，1uellerLaboratory，Pe nnsylvania State University，
University Park，Pe nnsylvania，1 6802 U. S. A.;
CDepartment of Botany，Na tional Science Museum，T okyo
4--1-1，Am akubo，Ts ukuba，30 5-0005 JAPAN
(Received on March 15，2 006)
The taxonomy of Anemone L. species distributed in the Southem Hemisphere was
re-evaluated on the basis of ac ritical morphological analysis of the available herbarium
material. A conspectus，ke y，de tailed morphological descriptions，to gether with figures of
flowers and fruits，a nd critical notes on all taxa are presented. Wea ccept 21 Anemone
species representing nine sections，a nd we confirm the validity of A. triternatα，A .
fanninii and A. peruviana which were not accepted by other authors. Wei nterpret the
mo叩hologicalsimilarities between A. thomsonii，A . caffra and A. janninii，A .
angustiloba，A. crassifolia，A. hepaticザ'oliaand A. rigida as corresponding to the section
level. Wed escribe one subsection within sect. Anemone and three series within sect.
Rivularidium. Within the southem hemisphere Anemone group the endelIuc taxa domi-
nate and all the monotypic sections eendemic. The majority of southem species e
紅 紅
components of non-tropical floras，li ke the northem taxa because they efound in high
ぽ
montane areas. Only af ew of them (viz.，A. thomsonii，A. angustiloba，A . peruviana and
A. jamesonii) 'eultra-oreophytes (growing at elevations of 2500-4000 m); A. decape-
訂
ωla and A. multifida occur alt. 100-3700 m. Five species (A. somaliensis，A. capensis，A.
antucensis，A .αssibrasiliana and A. moorei) can be regarded as hot-tropical plants，be ing
distributed alt. 300-1200 m.
Key words: Anemone，c omparative morphology，g eography，t axonomy，t he Southem
Hemisphere.
Anemone L. is one of the most remarkable The data on Anemone species are frag-
genera of the family Ranunculaceae and in- mented and sometimes debatable (Pritzel
c1uding about 150 species. The genus has a 1841，H ooker 1844，U lbrich 1906，L ourteig
worldwide distribution，w ith most represen- 1951，1 956，E ichler 1958，e tcふHowever，
tatives occu町ingmainly in the Northern the majority of the world's largest herbaria
Hemisphere. There are several Anemone spe- (viz.，K ，B M，E ，P ，W ，W U，L E，G H，U S，
cies disjunctively distributed in the Southern NY，et c.) include the herbarium specimens
Hemisphere，ma inly in montane regions with of these taxa，a nd we were lucky to examine
at emperate climate. most of these materials while visting Great
-193-
194 植物研究雑誌第81巻第4号 平成18年8月
Britain，A ustria，F rance，R ussia，U . S. A.， the components of the f10ras of the Southem
Vietnam and other countries. Hemisphere and they were included in three
Therefore，w e II_ltend to give our view on sections because most of them were moved
ap ossible differentiation and relations of the from sect. Anemonanthea to the recently
southem Anemone species on the basis of described sect. Rivularidium Jancz.
comparative morphology and taxonomy. We (Janczewski 1892) and sect. Eriocephalus
believe our inte叩retasionis valid because Hook. f. & Thoms. (Hooker and Thomson
the results of the analyses of molecular data 1855). Meanwhile four African species (A.
for several southem hemisphere Anemone capensis，A. alchimillifolia E. Mey. & Pritz.，
species were published recently (Hoot et al. A. fanninii Harvey and A. thomsonii Oliver)
1994， Ehrendorfer 1995， Hoot 1995， were included in sect. Pulsatilloides. Ulbrich
Ehrendorfer and Samuel 2001，S chuettpelz (1906) described 19 series，ei ght of which in-
et al. 2002)，a nd our taxonimic discussions cluded the southem species. Some of these
were informed by these studies. were recently p tof them were the basis for
紅
the recognition of several Anemone sections
Literature survey and even subgenera (Tamura 1995).
Three Anemone species from the floras of According to the most modem interpreta-
the Southem Hemisphere have been known tion of the genus Anemone and the family
since the 18th century (A. c(Jpensis， as Ranunculaceae (Tamura 1991， 1995)，
Atragene capensis-Linne 1753，A . decape- Anemone species from the f10ras of the
αtla-Arduino 1764 and A. triternαta-Vahl Southem Hemisphere (about 20) belong to
1794)，bu t most of these taxa were described five subgenera and seven sections: subgenera
in the 19th century (viz.，A. multifida-Poiret Anemone (sect. Anemone and Eriocephalus)，
1816，A . helleborifolia and A. tenuifolia- Rivularidium (Jancz.) Juz. (sect. Rivulari-
Candolle 1817，A. tenuifolia-Sprengel 1825， dium and Crassifolia Ulbrふ Hepaticifolia
A. antucensis-Poeppig 1833，A . hepatici- (Ulbr.) Tamura，R igida (Ulbr.) Tamura and
foliα-Hooker 1836 and others). In additim，a Pulsatilloides (DC.) Juz. (sect. Pulsatilloides，
few species (viz.，A . assibrasiliana -Kuhl Alchimillifolia (Ulbr.) Tamura and Kili-
1933， A. αngustiloba-Eichler 1958， A. mandscharica (Ulbr.) Tamura).
moorei-Espinosa 1940， A. somaliensis- Recently we re-examined the structure of
Hepper 1971) were described in the 20th the genus Anemone and we regard (Ziman
century. et al. 2002) that within it there e21
紅
1n the first system of the genus Anemone southem Anemone species belonging to nine
(Candolle 1817，1 824) the southem Anemone sections (because at present we do not recog-
species were placed in two sections (A. nize any Anemone subgenera).
decapetala，A tenuifolia，A. triternata and A. Here we present the conspectus of south-
multifida-sect. Anemonanthea DC.， A. em hemisphere part of the genus Anemone
capensis and A. helleborifolia-sect. (including nine sections and 21 species)，a
Pulsatilloides DC.)，a nd in the first mono- key for determination of species，t axonomic
graph of the genus Anemone (Pritzel 1841) analysis supported with detailed morphol-
they were included in four sections because ogic descriptions of all taxa，a nd the results
A. multifidαand A. helleborifolia were of discussion of their possible relationships.
placed in sect. Anemonospermos DC. and A.
hepaticifolia in sect. Homalocarpus DC.
1n the next monograph on Anemone
(Ulbrich 1906)，1 6 species were regdedas
紅
August 2006 Joumal of Japanese Botany Vol. 81 No. 4 195
Conspectus 2: 660 (1825)，p. p.
Anemone L.，S p. Pl. 538 (1753). Subsect. Longisηlae Ulbr. in Bot. Jahrb.
Type: A. coronariαL. 36: 204 (1905)，p. p.
Sect. 1. Anemone sensu Tamura in Acta Ser. 1. Multifidae Ulbr. in Bo .tJahrb. 36:
Phytotax. Geobot. 42: 180 (1991). 205 (1906) -Subsect. Multifidae (Ulbr.)
Genus Oriba Adans.，F am. Pl. 2: 459 Starodub.，V etrenytsy: 120 (1991).
(1763)，p. p. Type: A. multifidαPoir.
Sect. Anemonanthea DC.，S yst. Na .t1: 4. A. multがdaPoir.
212 (1817)，p. p. Sect. 3. Kilimandscharica (Ulbr.)
Sect. Oriba (Adans.) Spach，H ist. Nat. Tamura in Acta Phytotax. Geobot. 42: 180
Veg. 7: 250 (1839). (1991)-Ser. Kilimandschαricae Ulbr. in Bot.
Sect. Eriocephalus Hook. f. & Thoms.，F l. Jahrb. 36: 201 (1906).
Brit. Ind. 1: 20 (1855)，p. p. Type: A. thomsonii Oliver.
Subsect. 1. Somalienses Ziman，B ulakh & Subgenus Pulsatilloides (DC.) Juz.，F l.
Kadota，s ubsect. nov. URSS 7: 256 (1937).
Type: A. somaliensis Hepper. 5. A. thomsonii Oliver
Folia radicalia palmato-tripartita; involucri Sect. 4. Pulsatilloides DC.，S yst. Nat. 1:
phylla similia foliis folus radicalibus; 195 (1817)，p. p. -Subgenus Pulsatilloides
perianthii tepala persistentia， elliptico- (DC.) Juz.，F l. URSS 7: 256 (1937)，p. p.
oblanceolata， 15-30 mm longa， nervis - Genus Pulsatilloides (DC.) Starodub.，
basalbus 3-5，a nastomosibus 1-2; capitula Vetrenytsy: 124 (1991)，p. p.
carpellorum in fructu elongata; fructus Type: A. capensis (L.) DC.
ovoidei，j ugis lateralibus 0.2 mm latis， Subsect. Longisηlae Ulbr. in Bot. Jahrb.
stylodiis 1-1.2 mml ongis; grana pollinis 3- 36: 200 (1906)，p. p.
colpata. Ser. Pinnαt扮liaeUlbr. in Bot. Jahrb. 36:
Basal leaves palmately-triparted; involuc- 200 (1906).
ralleaves similar to basalleaves; tepals per- 6. A. capensis L.
sistent，e lliptic-lanceolate，1 0-15 mml ong， Sect. 5. Alchimillifolia (Ulbr.) Tamura in
basal tepal veins 3-5，w ith 1-2 anastomosing Acta Phytotax. Geobot. 42: 179 (l991)-Ser.
veins; fruiting heads elongate; achenes Alchimillifolia Ulbr. in Bot. Jahrb. 36: 200
ovoid，st yles 1.2-1.2 mml ong，m arginal ribs (1906)，p. p.
ca. 0.2 mmw ide; pollen grains 3-colpate. Type: A. c~所α(Ecklon & Zeyher)
1. A. somaliensis Hepper Harvey.
Subsect. 2. Carolinianae Starodub. in 7. A. c~所α(Ecklon & Zeyher) Harvey
Bot. ZhUffi. 74: 1345 (1989). 8. A. fanninii Harvey
Type: A. caroliniana Walter. Sect. 6. Rivularidium Jancz. in Rev. Gen.
2. A. decapetala Arduino Bot. 4: 251 (1892) -Subgenus Rivularidium
3. A. triternata Vahl (Jancz.) Juz.，F l. URSS 7: 255 (1937) -
Sect. 2. Eriocephalus Hook. f. & Thoms.， Subgenus Rivularidium sensu Starodub.，
Fl. Brit. Ind. 1: 20 (1855). Vetrenytsy: 119 (1991)，p. p.
Type: A. rupicola Cambess. Type: A. rivularis Buch.-Ham. ex DC.
Sect. Anemonanthea DC.，S yst. Na .t1: Genus Anemonidium (Spach) Holub in
212 (1817)，p. p. Folia Geobot. Phytotax. Praha 9: 272 (1974)，
Sect. Anemonospermos DC.，S yst. Nat. 1: p. p.
212 (1817)，p. p. Subgenus Meridium Starodub.，
Sect. D伊localymnataSpeng.，S yst. Veg. Vetrenytsy: 118 (1989)，p. p.
196 植物研究雑誌第81巻第4号 平成18年8月
Sect. Anemonospermos DC.，S yst. Nat. 1: Planta rhizomate， radicis adventitiae，
195 (1817)，p. p. caulisis sympodiale. Tepala 5-15，1 0-35 mm
Sect. Diplocαlymnαta Spreng.，S yst. Veg. longa，r ubra vel rubida，p ubescentia vel
2: 660 (1825)，p. p. glabra. Inflorescentiae pauciflore，s implici.
Ser. Rivulares Ulbr. in Bot. Jahrb. 36: 197 Petioli foliorum basalium basaliter abrupte
(1906)，e xc1. p. dilatati vel vaginati， laminae foliorum
Ser. 1. Angustilobae Ziman，B ulakh & basalium tematae， folioli.2-3-10bata vel
Kadota，se r. nov. p tita.
訂
Type: A.αngustilobαH. Eichler. Plants with rhizomes，a dventitious roots
Caudicis ramosi， radicis primariae， and sympodial scapes.Tepals 5-15，1 0-35
caulisis monopodiale. Tepala 5-7， 10-15 mm long，r ed or reddish， pubescent or
mm longa， pubescentia. Inflorescentiae subglabrous. Inflorescences few-flowered，
pauciflore， simplici. Petioli foliorum simple. Basalleaf petioles basally widely di-
basalium basaliter abrupte dilatati lated or vaginate，b asal leaf blades temate，
(auriculiforme)，l aminae foliorum basalium with 2-3-10bed or p紅白dleaflets.
tematae. 13. A. jamesonii Hook. f.
Plants with branched caudices，t ap-roots 14. A. sellowii Pritzel
and monopodial scapes.Tepals 5-7，1 0-15 15. A. assibrasiliana Kuhl. & Porto
mm long，p ubescent. Inflorescences few- 16. A. moorei Espinosa
flowered，s imple. Basal leaf petioles basally 17. A. antucensis Poeppig
sharply dilated (auriculate)，ba salleaf blades 18. A.tenuicaulis (Cheeseman) Parkin &
ternate. Sledge
9. A.angustiloba H. Eichler Sect. 7. Crassifolia (Ulbr.) Tamura in
10. A. sumαtrana de Vr iese Acta Phytotax. Geobot. 42: 178 (1991) -Ser.
Ser. 2. Mexicanae (Starodub.) Ziman， Crassifolia Ulbr. in Bot. Jahrb. 36: 199
Bulakh & Kadota， comb. nov. - Sect. (1906).
Mexicanae Starodub.， Vetrenytsy: 119 Type: A. crassifolia Hook. f.
(1989). 19. A .crassifolia Hook. f.
Type: A. mexicana Humb.， Bonpl. & Sect. 8. Hepaticifolia sensu Tamura in
Kunth. Sci. Rep. Osaka Univ. (16): 28 (1967) -Ser.
Plants with branched caudices，t ap-roots Hepaticifolia Ulbr. in Bot. Jahrb. 36: 197
and sympodial scapes. Tepals 4(-5)，6 -15 (1906) - Subgenus He pαticifoliα(Ulbr.)
mm long， white-yellowish， glabrous. Tamura in Acta Phytotax. Geobot. 42: 178
Inflorescences many-flowered，c ompound. (1991) - Genus Anemonidium subgen.
Basal leaf petioles basally sharply dilated Meridium Starod. sect. Meridium subsect.
(auriculate)，ba salleaf blades palmatifid with Hepaticifolia (Ulbr.) Starodub.，V etrenytsy:
pinnatisect leaflets. 118 (1991).
Subsect. Helleborifolia Stodub.， Type: A. hepaticifolia Hook. f.
訂
φ
Vetrenytsy: 119 (1989) -Ser. Hellebor lia Sect. Rivularidium Jan cz. in Rev. Gen.
Tamura in Acta Phytotax. Geobot. 42: 178 Bot. 4: 251 (1892)，p. p.
(1991)，ut ‘Helleborifoliae' . 20. A. hepaticifolia Hook. f.
11. A. helleborifolia DC. 9. Sect. Rigida (Ulbr.) Tamura in Sci.
12. A. peruviana Britton Rep. Osaka Univ. (16): 28 (1967) -Ser.
Ser. 3. Jamesonii Ziman， Bul広h & Rigida Ulbr. in Bot. Jahrb. 36: 199 (1906)
Kadota，se r. nov. -Subgenus Rigida (Ulbr.) Tamura in Acta
Type: A. jamesonii Hook. f. Phytotax. Geobot. 42: 178 (1991) -Genus
August 2006 Joumal of Japanese Botany Vo .l81 No. 4 197
Anemonidium subgen. Meridium Starodub. 0.5-1.0 mml ong，a chene hairs 4-5 mmlo ng;
sect. Meridium subsect. Rigida (Ulbr.) basal leaves 1-2-temate (subsect. Cぽoli-
Starodub.，V etrenytsy: 119 (1991). nianae) ....・ ・....・ ・...・ ・・・-…...・ ・.....・ ・...・...5
H H H H H H H
Type: A. rigidαGay. 5a. Tepals deciduous，8 -20 x2 -5 mm，w ith
，
Sect. Rivularidium Jan cz. in Rev. Gen. 5-7 basal veins and 1-2 anastomosing veins;
Bot. 4: 251 (1892)，p. p. filaments linear; inf10rescences few-
21. A. rigida Gay f1owered; basal and involucral leaves dis-
similar，b asalleaves dimorphic .…・
Key to species of Anemone from ..2. A. decapetala
the Southern Hemisphere 5b. Tepals persistent，1 0-15 x2 -3 mm，w ith
1a. Carpels and achenes more or less densely 3 basal veins and without anastomosing
pubescent， symmetrical， mainly ovoid， veins; filaments filiform; f10wers solitary;
sharply narrowed into substraight styles…2 basal and involucral leaves similar，. basal
1b. Carpels and achenes glabrous or leaves monomo中hic....・ ・...3. A. triternata
H
subglabrous，m ainly assymetrical，o blong- 6a. Carpels and achenes distinctly stalked，
ovoid，g radually narowed into curved or cylindroid，c overed with hairs 3-4 mml ong;
hooked styles ・... ・.....・ ・.....・ ・・・.....・ ・..…・ 9 styles 1-2 mml ong; stigmas capitate; tepals
H H H H H H
2a. Carpels and achenes with dense hairs， having few anastomosing veins，s parsely pu-
mainly longer than the bodies; tepals bescent; basal leaf petioles basally sha中ly
monomo中hic，densely pubescent; above- dilated (auriculate)， basal leaf blades
ground shoots sympodial or monopodial glabrous; plants with short rhizomes and ad-
…3 ventitious roots (sect. Kilimandscharica) .
2b. Carpels and achenes covered with hairs …… 5. A. thomsonii
shorter than the bodies; tepals monomorphic 6b. Carpels and achenes sessile，s pindle-like
or dimorphic，p ubescent or glabrous; above- or oblong-ovoid，c overed with hairs 2-3 mm
ground shoots sympodial ....・ ・-…...・ ・...・ ・.6 long; styles 6-10 mml ong; stigmas linear;
H H H
3a. Carpels and achenes ovoid-globose; tepals having more than 5( sometimes more
tepals 10-18，d eciduous or persistent; above- than 10) anastomosing veins，d ensely pubes-
ground shoots sympodial; plants with tuber- cent; basal leaf petioles basally slightly di-
ous rhizomes and adventitious roots (sect. lated，b asal leaf blades pubescent; plants
Anemone) .…… ・・...・ ・・・...・ ・....・ ・...・ ・...4 with caudices and tap-roots ・... ・.....・ ・..…..7
H H H H H H H H H H
3b. Carpels and achenes subellipsoid; tepals 7a. Carpels and achenes spindle-like; tepals
6-10， persistent; above-ground shoots 15-25，2 5-50 mml ong，l inear-lanceolate，
monopodial; plants without tuberous rhi- densely pubescent，h aving 3-9 anastomosing
zomes but having branched caudices and tap- veins; leaves 子temate，subglabrous; non-
roots (sect. Eriocephalus)…… 4. A.multifida rosetteous semi-shrubs (sect. Pulsatilloides)
4a. Tepals elliptic-obovate，gl abrous; ca叩els ..6. A. capensis
and achenes ovoid and not compressed; 7b. Carpels and achenes oblong-ovoid; tepals
ca叩eland achene styles 1-2 mm long， 10-12，2 0-40 mm long，w ide-lanceolate，
achene hairs 3-3.5 mml ong; basal leaves sparsely pubescent，h aving more than 10
palmately 3-parted (subsect. Somaliense) .... anastomosing veins; basal leaves palmately
.1. A. somαliensis lobed，v illous; semirosetteous herbaceous
4b. Tepals linear-oblong or lanceolate， plants (sec .tAlchimillifolia) ....・ ・.....・ ・-….8
H H
densely pubescent; ca中els and achenes 8a. Carpels and achenes 4-5 mm long，
subglobose，d istinctly compressed and with covered with dimorphic hairs 1-5 mml ong;
visible lateral ribs; cpeland achene styles tepals monomorphic; filaments distinctly
紅
198 植物研究雑誌第81巻第4号 平成18年8月
dilated; basal leaf blades 5-7 cm wide .. 13a. Tepals 6-12 mm long， yellowish，
…-.7. A. C~伊α achene styles basally curved，2 -3 mml ong;
8b. Carpels and achenes 5-10 mml ong，co v- basal leaf blades 8-15 x1 2-18 cm，b ract
ered with monomorphic hairs 3-6 mml ong; petioles wide，3 -5 mml ong .
tepals dimorphic; filaments slightly dilated; ・... ・11.A. helleborifoliα
H
basalleaf blades 12-20 cm wide ..… 13b. Tepals 12-15 mml ong，w hite; achene
…. 8. A. fanninii styles apically uncinate，1 -2 mml ong; basal
9a.Cぽpelsand achenes oblong-ovoid (rarely leaf blades 6-9 x7 -15 cm，b ract petioles
subcylindroid)，m ainly 3-7 mml ong; tepals narrow，1 -3 mml ong ・…… 12.A. peruviana
(5-)6-15 (Sect. Rivularidium)…....・ ・-…..10 14a. Tepals with naowbases and wide api-
町
H
9b. Carpels and achenes shortly-ovoid or ces，a nastomosing veins absent; basal leaf
cylindroid，2 -4(-5) mml ong; tepals (4-)5-7 petioles basally sha中lydilated .…...・ ・-… 15
H
.1 9 14b. Tepals with naowbases and apices，
町
10a. Tepals 5-7，wh ite or pinkish，pu bescent; having or not having vein anastomose; basal
basal leaf petioles basally sharply dilated， leaf petioles basally vaginate ・... ・・・"…・ 17
H H H
basal leaf blades ternate; plants with 15a. Tepals 5，8-12 mml ong，m onomo叩hic，
caudices，t ap-roots and monopodial scapes sparsely puberulent along central vein;
.11 C紅pelsand achenes 2-4 mm long，w ith
10b. Tepals 5-15， red or yellowish， styles ca. 1m ml ong; cymes 2-3-flowered;
subglabrous; basal leaves various; plants basal leaf blades 2- or 3-ternate， not
various underground shoots and roots but coriaceous; bracts shortly petiolate，bi ternate
with sympodial scapes ....・ ・-……....・ ・...… 12 13.A. jα:mesonii
H H 日
l1a. Tepals 5-7，1 5-20 mml ong，w hite， 15b. Tepals 8-16，1 5-35 mml ong，d imor-
with 3-5 anastomosing veins; cymes 1-2- phic，g labrous or subglabrous; ca中elsand
flowered; achene styles uncinate，1 -1.5m m achenes 3-9 mml ong，w ith syles 2-6 mm
long ・... ・....・ ・....・ ・...・ ・-….9. A. angustiloba long; cymes 1-2-flowered; basalleaf blades
H H H H
llb. Tepals 5，1 0-15 mml ong，pi nkish，wi th ternate，co riaceous，br acts sessile，en tire (dis-
7-9 anastomosing veins; cymes 3-5- tally dentate) ・... ・・・......・ ・......・ ・..… ・.・16
H H H H H H H
flowered; achene styles hooked，0 .3-0.5 mm 16a. Achenes 6-9 mml ong，w ith styles 4-6
long ....・ ・-…....・ ・....・ ・...… 10.A. sumatrana mml ong; basal leaflets 2-3-parted (lobed)
H H H
12a. Tepals 4(-5)，6 -15 mml ong，w hite- .14 . A. sellowii
yellowish，w ith solitary anastomosing veins， 16b. Achenes 3-5 mml ong，wi th styles 2-3
glabrous; inflorescences many- flowered， mml ong; basal leaflets entire .
compound; leaflets of basal leaf blades …… 15. A.αssibγαsilianα
pinnatisect，g labrous; plants with caudices， 17a. Tepals 20-35 mm long， with 5-7
tap-roots and widely dilated bases of basal anastomosing veins， sp sely pubescent
紅
leaf petioles (Ser. Mexicanae) ....・ ・...・ ・..13 along central vein; cpelsand achenes not
紅
H H
12b. Tepals 5-15，1 0-35 mml ong，m ainly compressed， basally sparsely pubescent;
red or reddish，h aving or not having anasto- styles substraight; basal leaf blades 15-35
mosing veins，p ubescent or subglabrous; cm long，gl abrous .…....・ ・...・...16. A. moorei
H
inflorescences mainly few-flowered，s imple; 17b. Tepals 4-15 mm long， without
leaflets of basal leaf blades 2-3-10bed or anastomosing veins， subglabrous; carpels
p d，sp selypuberulent; plants with rhi- and achenes compressed，h aving lateral ribs，
紅白 紅
zomes，ad ventitious roots and widely dilated glabrous; styles curved; basal leaf blades 3-
or vaginate bases of basal leaf petioles (ser. 8c m long，s p selypubescent ....・ ・.，.・ .・18
紅
H H
Jamesonii) ・…・・ ・・..… ・・...・ ・.....・ ・...・ ・...14 18a. Tepals 7-15 mml ong，el liptic，pi nkish-
H H H H H H H
August 2006 Joumal of Japanese Botany Vo1. 81 No. 4 199
white; achenes shortly stalked; styles 3-5 30-40o btuse ultimate lobules，gl abrous (Fig.
mml ong; inflorescences 2-3-flowered; basal la). Scapes 7-18 cml ong，1 -2-flowered (lat-
leaf blades 3-parted or 3-10bed; plants with eral flower frequently under-developed)，
short rhizomes…・ ・・...・ ・.17.A.αntucensis appressed-pubescent. Involucral leaves 3
H H H
18b. Tepals 4-6m ml ong，li near-lanceolate， (-4)， sessile，ba sally connate，si milar to basal
red; achenes sessile; styles 1-2 mml ong; leaves，bl ades 2-3-p紅白dor lobed，w ith 10-
flowers solity;.basal leaf blades ternate or 15 obtuse ultimate lobules or teeth，1. 5-2.5
征
bitemate; plants with short nodulose and cm long，su bglabrous. Pedicels 2-4 cm long，
long stolon-like rhizomes …. densely pubescent under flowers. Tepals 10
.18 . A. tenuicaulis -18， persistent，- monomorphic， elliptic-
19a. Achenes 2-3 mml ong; tepals 12-18 obovate，w ith wide bases and obtuse dentate
mm long，w ith 3-5 anastomosing veins; apices，b lue or mauve，1 0-15 x3 -5 mm，
flowers solitary; plants with tuberous and with 3-5 basal veins and few anastomoses，
stolon-like rhizomes (sect. Crassifolia)… glabrous. Stamens 3-4 mml ong，w ith linear
.19 . A. crassifolia filaments，e llipsoid anthers and narrow con-
19b. Achenes 4-5 mml ong; tepals 15-35 nectives. Carpels ovoid，no t compressed，ca .
mml ong，w ith more than 10 anastomosing 1m ml ong，de nsely covered with hairs 3-3.5
veins or without them; inflorescences com- mml ong，st yles straight，1 -2 mml ong; stig-
pound，ma ny-flowered; plants without tuber- mas linear. Fruiting heads elongate. Achenes
ous and stolon-like rhizomes….....・ ・...・...20 ovoid，1. 6-2.0 x1 -1.2m m，la nate，m arginal
H
20a. Tepals pinkish-white，w ithout anasto- ribs ca. 0.2 mmw ide; hairs 3-3.5 mml ong;
mosing veins， sparsely pubescent; anther styles straight，1 -2 mml ong，s tigmas linear
connectives wide，w ith projections; basal (Fig.2a).
leaf petioles basally sharply dilated and their Chromosome number: unknown.
blades 3-5-10bed; plants with short rhizomes Pollen grains: tricolpate (present paper:
and adventitious roots (sect. Hepaticifolia) voucher specimen Thulin & Warfa s. n.，
…20. A. hepaticifolia UPS-K).
20b. Tepals red， with more than 10 Distribution and habitat: E. Africa，
anastomosing veins，g labrous; anther con- Somalia，n a町owendemic of Al Medo Hills;
nectives narrow，w ithout projections; basal in stony ground or limestones，t ogether with
leaf petioles basally vaginate and their blades species of Buxus，O lea，D odonaea and others
勾
3-5-parted; plants with caudices and tap- in evergreen bushes，al t. 920-1200 m.
roots (sect. Rigida) ・... ・.....・ ・..21. A. rigida Specimens examined: SOMALIA. In midst belt of
H H
N facing limestone，e sca叩mentwith considerable win-
1. Anemone somaliensis Hepper in Kew ter rainfall from NE Monsoon. Evergreen，b ushland
with Acokanthera，B uxus，D odonaea，O lea africana.
Bull. 26: 57 (1971). Type: N. Africa.
N. of Galgallo，11 00 l'N，49 002'E，13 00 m，7. 12.1969.
SOMALIA. South of Al Hillas，st ony ground
Lavranos 7300 (K); B i，Escarment，S of Bunder
訂
in shade，3 000 f ，.t10.11.1929，C. Barrington Murraya，B uraha Dhasi，1 1038-39 'N，5 0029-32 'E，
Brown in Herb. Collenette 413 (holotype- 1050 m，1 6-17.11.1986，T hulin & Warfa s. n. (UPS-
K!). K).
Note: The collector of the type specimen
Rhizomes tuberous，s tout and irregular，
ca.15 x1 2 mm，n on-branched. Basal leaves Barrington Brown believed this plant was A.
blanda，b ut Hepper (1971) after his re-
2-3; petioles 5-15 cm long，s ubglabrous or
examination determined this plant as an ew
sp selypubescent; blades monomorphic，
紅
species (A. somαliensis) which he regarded
palmately 3-parted，3 -7 x5 -10 cm，wi th ses-
as taxonomically close to A. hortensis but
sile crenate-dentate primaηsegments and
200 植物研究雑誌第81巻第4号 平成18年8月
b
a
t
小
W
H
a
na
u
ug
z

d e
Fig. 1. Flowers of Anemone species from the Southem Hemisphere. a. A.
decapetala (Argentina. Tucuman Tabi，1 0.5.1950，R ocha 2902，L E). b.
A. triternatα，(Bolivia. Meguel Bang Lectae ex Herb. Collegii Columbix，
1891，B ritton & Rusby 1041，L E). c. A. mult折dα(Argentina.Estancia
Harberton，B rown Chico，7. 1.1967，G oodale 468，M HA). d. A. thomsonii
(Sudan. Mts. Matong，1 2.2.1936，J ihnston 1512，K ). e. A. capensis
(South Africa. Cape Distr.，7 .1890，G amble 22096，K ). 1: Petal. 2:
Stamen. 3: Carpel. Scale indicates 1m m.
differing from it by much larger involucral regarded it as at axon close to A. hortensis
leaves (similar to basal ones) and smaller but differing by its stout rhizomes，s parsely
perianth. Thulin (1993)，w ho included A. pubescent or subglabrous basaI leaf petioles，
somαliensis in“Flora of Somaliaぺalso tall stems (to 20 cm long) appressed-
August 2006 Joumal of Japanese Botany Vol. 81 No. 4 201
pubescent distally，3 -4 involucral leaves， A. bilobata Phi ，.lCat. Plants Vasc. Chile 5
partially united basally and sp selypubes- (1881).
紅
cent，a nd pedicels 2-4 cm long，d ensely pu- A. polypetala Laanagain Escritos 2: 178
町
bescent in the upper part，f lowers with 10- (1922).
18 blue or mauve tepals 10-15 mml ong，ob - Rhizomes tuberous，ov oid-cylindroid，1. 7-
tuselyemginateor variously divided at the 2.5 x1 .2-1.8c m. Basalleaves 3-5，p etioles
訂
apex，a nd pilose carpels with erect styles. 2-10 cm long，s carcely pubescent，w ith
According to the results of our re- gradually dilated scale-like bases (6-8 mm
examination of the type specimens of A. wide); blades dimorphic，2 -ternate，1 -2 x1 -
somaliensis in K (Barrington Brown 413)， 7m m，sc arcely pubescent. Early basalleaves
we confirmed most characters noted by with 10-15 broadly ovate obtuse ultimate di-
Thulin but we regard that the characters in visions; primary segments subsessile or on
common with A. hortensis are few (few petiolules 2-3 mml ong; later basal leaves
tepals and elongate achene heads) but the with 40-60 linear-oblong wide-obtuse ulti-
characters in common with A. tsche.rηjaewii mate lobules; primary segments on petiolules
Regel and A. serawschanica Kom. from the 5-20 mml ong (Fig. 2a). Fruiting plants
flora of Central Asia are more numerous sometimes lacking basalleaves. Scapes 5-35
(tuber shape，b asal leaf petioles basally not cm long，f ew-flowered，s carcely puberulent.
dilated，n ot connate basal part of involucral lnvolucral leaves 3，o n petioles 2-5 x3 -5
leaf petioles similar to those in basal leaves， mm; blades 1-temate，d issimilar to the basal
and tepals having solitary anastomosing ones，2 -10 cm long，s carcely puberulent;
veins). The most significant phenomenon is blades 1-ternate; primary segments with 20-
the tricolpate pollen grains of A. somaliensis 30 linear-oblong long-acute ultimate lobules.
which we studied firstly for this species Lateral flowers with two small linear
(仕omThulin & W紅白， UPS-K) characteris- bracteoles. Pedicels 5-25 cm long，s c紅cely
tic for subsect. Carolinianae and absent in puberulent. Tepals 10-12，d eciduous，li near-
both subsect. Anemone and Biflora There- oblong，b lue or whitish-pink，m onomorphic，
fore，w e note A. somaliensis stands apart 8-20 x2 -5 mm，w ith (3-)5-7 basal veins
from the both Mediterranean subsect. and 1-2 anastomosing veins，d ensely pubes
聞
Anemone and Central Asian subsect. Biflora cent. Stamens 3-5 mml ong，w ith linear fila-
and we sep ateit into the monotypic sub- ments， globose anthers and naow
紅 町
section Somaliense，s ubsect. nov. connectives. Carpels subglobose， slightly
compressed (marginal ribs ca. 0.2 mmw ide)，
2. Anemone decapetala Arduino， 1-2 mml ong，d ensely covered with hairs
Animadv. Bot. Spec. Alt. 2: 27 (1764). 0.7-1 mml ong; styles curved，0 .5-0.7 mm
Type: N.710-21 (lectotype-LINN!). De long; stigmas linear (Fig. la). Fruiting heads
semillas procedentes de Brasil (paratype- elongate-cylindroid， 2.0-2.5 x1 .5-2.0 cm.
P!). Achenes subglobose，w ith ribs 0.3-0.6 mm
A. trilobata Juss. in Ann. Mus. Hist. Nat. wide，1 .5-2.5 x1 .3-1.8 mm; densely pube-
Paris 3: 248 (1804). scent (hairs 4.5-5.7 mm long); styles
A. sphenophylla Poepp.，F ragm. Syn. Pl.: substright or subulate，0 .7-1.2 mml ong，
27 (1833). stigmas linear (Fig. 2b).
A. chilensis Spreng. ex Pritz. in Linnaea Chromosome number: n= 8 ，1 2 (Joseph
15: 626 (1841)，n om. nud. and Heimburger 1966，R othfels et al. 1966，
A. macrorrhiza Domb.，F l. Bras. 13: 151 Baumberger 1970).
(1864). Pollen grains: pantocolpate (Huynh 1970).
202 植物研究雑誌第81巻第4号 平成18年8月
Distribution and habitat: mainly eastern ter unique within the tuberous species of
part of extratropical South America Anemone in the New World) ， and
(Argentina，B razil，C hile，P eru，U ruguay); on subglobose achenes with mainly subulate
rocky slopes， gravelly soil and shaded styles densely covered with hairs 4.5-5.7
places，al t. 100-3000 m. mml ong.
Specimens examined: ARGENTINA. San Martin Meanwhile，w e have to note that the data
de los Andes，3 000 m，3 .11.1926，C omber 725 (K); for our comparative-mo叩hological study
Tucuman: Estancia Santa Maria，3 0.8.1949，P ederson were rather limited and its results are not suf-
412 (K); San Javier 10.2.1950， Rocha (WU);
ficient base for elaboration of the
Empedrado: Corientes，22 .8.1971，Pe derson s. n. (GH);
interspecific structure of the polymorphic
Sierra de la Venlana，C erro Ventruz，2 8.9.1981，R oig
taxon A. decapetala.
47037，( K); Magellan，C ape Negro，C ape Darwin，
without date，H emslow s. n. (K). CHILE. Conception，
乱1acrae，10.1825，B ridges s. n. (K); Valparaiso，1 831， 3. Anemone triternata Vahl in Symb.
Cuming s. n. (K); Valparaiso，1 832，Br idges (K); 1856， Bot. 3: 74 (1794). Type: URUGUAY. Circa
H vey(K); Santiago， 2.1856 (KW); Valparaiso，
訂 Monte-Video ad ostium fluminis Plata，
Durren Ebenen， 5.8.1895， Buchtien s. n. (E);
Commerson，n o date nor herbarium locality.
Valparaiso，Q uintero，9 .1923，W enderman s. n. (E);
San Martin de los Andes，3 000 m，3. 11.1926，C omber A. tridentata Vahl in Symb. Bot. 3: 75
725 (K); Colchagua: San Femando，C erro Nicunlanta， (1794).
9.1928，M ontero 732 (K); Prov. Coquimbo，D ept. A.βmariefolia Juss. in Ann. Mus. Hist.
，(，
Illapel，L a Vega Escondida，3 450 m，2 0.12.1938，
Nat. Paris 3: 247 (1804).
Morrison s. n. (K); Alto del Puerto， 18.8.1940，
A. decapetala Arduino var. foliolosa H.
Santesson s. n. (K). URUGUAY . Montevideo: Cerro
Cassabo，9. 1926，He rter s. n. (GH). Eichler，F l. Brasil. 13: 151 (1864).
Note: This species was described from the A. cicutゆlia1. M. Johnst. in J. Am.
flora of Chile in 1764 by Arduino，a nd it was Arbor. 19: 248 (1938).
inc1uded in early works (e. g.，L innaeus Rhizomes tuberous， elongatecylindric，
町
1753，C andolle 1817，P ritzel 1841)，a s well 1.5-2.5 x0 .7-1.0c m. Basalleaves 2-5，pe ti-
as more recent papers (e. g.，B ritton 1892 oles 3-13 cm long; expanded basally，
and Lourteig 1951). In Linnaeus' Species glabrous; blades 1-2-temate，m onomorphic，
Plantarum of 1753，th is species was still ab- 2-5 x2 -5 cm; primary segments with long，
sent，b ut we have examined in LINN the narrow-linear acute ultimate lobules，
lectotype specimen which was the basis for scarcely pubescent; petiolules 10-25 mm
the Arduino description. long. Scapes 10-20 cm long，l -flowered，
It is the rather polymorphic species within sc lypubescent. Involucral leaves similar
紅白
which several n rowor segregate species to basal ones，1 -2-temate，b lades 5-8 cm
紅
were described (viz.， A. trilobαta， A. long，w ith petiole-like bases; ultimate lob-
polypetala，et c.). However，w e note白atall ules linear，a cute，s c訂celypubescent (Fig.
specimens referred to this taxon which we 1b). Pedicels 5-25 cm long， densely
examined in K，B M，G Ha nd other herbaria pubescent. Tepals 10-15，p ersistent，m ono-
have several common differential characters: morphic， lanceolate， apically acuminate，
ovoid-cylindroid tubers， 1-2tematebasal white to pink，1 0-15 x2 -3 mm，wi th 3b asal
四
and l-temate involucral leaves (latter ones veins and without anastomosing veins，
dissimilar to former leaves)，pr esence of two densely pubescent only basally. Stamens 5-
small bracteoles)，f ew-flowered scapes，1 0- 6m ml ong，wi th filiform filaments，e l1ipsoid
12 deciduous，m onomo中hic，linear-oblong， anthers and na町owconnectives. Carpels
blue or whitish-pink tepals having 5-9 basal subglobose，s lightly compressed (ribs 0.2-
veins with 1-2 anastomosing veins (charac- 0.3 mmw ide)，ca . 1m ml ong，d ensely cov-