黒沢高秀 1990年10月22日作成
高等植物の炭酸同化経路には、C3、C4、およびCAMの３っつの型がある。
C3型はリブロース二リン酸 (RuDP) カルボキシラーゼによってホスホグリセリン酸 (PGA) にCO2固定する型で、植物に最も普通に見られる。
C4型はホスホエノールピルビン酸 (PEP) カルボキシラーゼにより、葉肉細胞中でCO2を固定し、アスパラギン酸 (Asp) またはリンゴ酸 (Mal) を初期産物として生じる型で、CO2固定効率が良く、気候抵抗が高い。そのため、強光、乾燥条件に強い特性を持つ。1900年までに、21科2000種以上の植物が確認されたが、すべて種子植物で、その多くがグラミノイド (イネ科、カヤツリグサ科植物) である。双子葉植物ではアカザ科、トウダイグサ科、キク科、ヒユ科などに散見される。
CAM型は、夜間に気孔を開いてCO2を取り入れ、PEPカルボキシラーゼにより炭酸固定を行い、液胞中に有機酸を貯める型である。昼間は気孔を閉じて水の蒸散を防ぐので、極端に乾燥条件に強い。ベンケケイソウ科、サボテン科に一般に見られる他、ツルナ科、トウダイグサ科、キク科などの多肉植物に見られる。また、裸子植物、シダ植物でも知られている。
トウダイグサ科にはこれら３型のすべてが見られる。特にトウダイグサ属 (広義) Euphorbia には１属中に３型が見られる極めて稀な例であり、生活形とともに代謝も大きく変化させて様々な生育地に適応してきたことを伺わせる。トウダイグサ属の中でもニシキソウ亜属 subgen. Chamaesyce はC4植物で、それと近縁なアガロマ亜属 subgen. Agaloma やショウジョウソウ亜属 subgen. Poinsettia はC3植物であることが Moss et al. (1969) によって示された。その後 Webster et al. (1975) によって、トウダイグサ属内のC3、C4、およびCAMの分布が調べられ、C4植物はニシキソウ亜属のみに出現すること、ニシキソウ亜属でもアクタエ亜節 subsect. Acutae の一部はC3植物であることを明らかにした。これをもとに、Websterはニシキソウ亜属の起源がこの亜節にあると推察した。
本研究では、Webster et al. (1975) のリストを原型にして、1990年までのデータを加えてトウダイグサ科全体における３型の炭酸固定経路分布のリストを作成した。その結果、以下のことが明らかになった。
1. トウダイグサ科の５亜科52連のうち、現在までに５亜科17連について調査されたが、C4植物とCAM植物はともにトウダイグサ連 tribe Euphoribieae に限られる。
2. C4植物はニシキソウ亜属のみに見られ、同亜属内ではアクタエ亜節のみにC3植物が見られることが再確認された。
3. CAM植物は多肉のトウダイグサ属植物、モナデニウム属 Monadenium、シナデニウム属 Synadeniumで見られるが、データが少なく、系統との関係ははっきりしない。
REFERENCES
Bender, M. M., I. Rouhani, H. M. Vines & C. C. Black Jr. 1973. 13C/14C ratio
changes in crassulacean acid metabolism plants. Plant Physiol. 52: 427-430.
Brown, W. V. 1974. Another cytological difference among the Kranz subfamilies
of the Gramineae. Bull.Torrey.bot.Club 101: 120-124.
Downton, W. J. S. 1975. The occurrence of C4 photosynthesis among plants.
Photosynthetica 9: 96-105.
Gutierrez, M., V. E. Gracen & G. E. Edwerds 1974. Biochemical and cytological
relationships in C4 plants. Planta 119: 279-300.
Herbst, D. 1972. Ontogeny of foliar venation in Euphorbia forbesii.
Amer.J.Bot. 59: 843-850.
Hnatiuk, R. J. 1980. C4 photosynthesis in the vegetation of Aldabra Atoll.
Oecologia 44:327-334.
Inbanba, S. K. & G. Papa 1979. Distribution of the Kranz type anatomy in some
dicotyledonous families of Kenya. Photosynthetica 13: 315-322.
Krenzer, E. G. Jr., D. N. Moss & R. K. Crookston 1975. Carbon dioxide com-
pensation points of flowering plants. Plant Physiol. 56: 194-206.
Moss, D. N., E. G. Krenzer Jr. & W. A. Brun 1969. Carbon dioxide compensation
points in related plant species. Science 164: 187-188.
McWillams, E. 1970. Comparative rates of dark CO2 uptake and acidification in
the Bromeliaceae, Orchidaceae, and Euphorbiaceae. Bot.Gaz. 131: 285-290.
Nuernbergk, E. L. 1961. Endogener Rhythmus und CO2-Stoffwechsel bei Pflanzen
mit diurnalem Saurerhythmus. Planta 56: 28-70.
Okuda, T. & A. Furukawa 1990. Occurrence and distribution of C4 plants in
Japan. Jpn.J.Ecol. 40: 91-121.
Oudejans, R. C. H. M. 1990. World catalogue of species names published in the
tribe Euphorbieae (Euphorbiaceae) with their geographical distribution.
444pp., Utrecht.
Pearcy, R. W. & J. Troughton 1975. C4 photosynthesis in tree from Euphorbia
species from Hawaiian rainforest sites. Plant Physiol. 55: 1054-1056.
Raghavendra, A. S. & V. S. R. Das 1978. The occurrence of C4-photosynthesis:
A supplementary list of C4 plants reported during late 1974 - mid 1977.
Photosynthetica 12: 200-208.
Sankhla, N., H. Ziegler, O. P. Vyas, W. Stichler & P. Trimborn 1975. Eco-
physiological studies on Indian arid zone plants. V. A. screening of some
species for the C4-pathway of photosynthetic CO2-fixation. Oecologia 21:
123-129.
Schoch, E. & D. Kramer 1971. Korrelation von Merkmalen der C4-Photosynthese
bei Vertretern vershiedener Ordnungen der Angiospermen. Planta 101: 51-66.
Szarek, S. R. & I. P. Ting 1977. The occurrence of crassulacean acid metabo-
lism among plants. Photosynthetica 11: 330-342.
Webster, G. L. 1967. The genera of Euphorbiaceae in the Southeastern United
States. J.Arnold Arb. 48: 303-430.
-------------- 1975. Conspectus of a new classification of the Euphorbiaceae.
Taxon 24: 593-601.
Webster, G. L., W. V. Brown & B. N. Smith 1975. Systematics of photosynthetic
carbon fixation pathways in Euphorbia. Taxon 14: 27-33.
Welkie, G. W. & M. Caldwell 1970. Leaf anatomy of species in some dicotyledon
families as related to the C3 and C4 pathways of carbon fixation.
Canad.J.Bot. 48: 2135-2146.
Ziegler, H., K. H. Batanouny, N. Sankhla, O. P. Vyas & W. Stichler 1981: The
photosynthetic pathway types of some desert plants from India, Saudi Arabia,
Egypt, and Iraq. Oecologia 48: 93-99.
Distribution of photosynthetic syndromes in Euphorbiaceae.
Takahide Kurosawa, 22 Oct. 1990.
Taxon
Circumscriptions and arrangement of the subfamilial and tribal taxa
follow the classifications of Webster(1975), and those of infrageneric taxa
of Euphorbia follow Webster(1967) and Webster et al.(1975). Scientific names
of trib. Euphorbieae follow Oudejans(1990) and synonyms are shown between the
citationmarks.
Status
3...... C3
4...... C4
S...... CAM
?...... unknown.
Methods
A/a...... Kranz/non-Kranz anatomy
C/c...... low/high CO2 compensation point
D/d...... nocturnal acidification exist/not exist
E...... C4 determined by electoron microscopy
Mk/Mc/m...... metabolically C4/CAM/C3 status
N/n...... nocturnal CO2 assimilation exist/not exist
R/r...... high/low delta13C values.
Status and methods
C3...... a c d - m n r nonsuccurent
C4...... A C d E Mk n R nonsuccurent
CAM...... a C D - Mc N R succurent
Reference
<1>, Bender et al.(1971)
<2>, Brown(1974)
<3>, Downton(1975);
<4>, Gutierrez et al.(1974)
<5>, Herbst(1972)
<6>, Hnatiuk(1980)
<7>, Imbamba & Papa(1979)
<8>, Klenzer et al.(1975)
<9>, McWilliams(1970)
<10>, Moss et al.(1969)
<11>, Nuernbergk(1961)
<12>, Okuda & Furukawa(1990)
<13>, Pearcy & Troughton(1975)
<14>, Raghavendra & Das(1978)
<15>, Sankhla et al.(1975)
<16>, Schoch & Kramer(1971)
<17>, Szarek & Ting(1977)
<18,19, 20,21,22,23,24,25,26>, Webster et al.(1975)
<27>, Welkie & Caldwell(1970)
<28>, Ziegler et al.(1981)
Taxon Status; Methods <References>
----------------------------------------------------------------------
EUPHORBIACEAE
Subfam. PHYLLANTHOIDEAE
Trib. DRYPETEAE
Genus Putranjiva
P. roxburghii Roxb. 3; a <27>
Trib. PHYLLANTHEAE
Genus Glochidion
G. harveyanum Domin. 3; a <27>
Genus Margaritaria
M. anomala (Baillon) Fosberg 3; a <7>
Genus Meineckia
M. phyllanthoides Baill. 3; a <7>
Genus Phyllanthus
P. carolinensis Walt. 3; a <27>
P. casticum Soy.-Will. 3; a <6>
P. juglandifolius Willd. 3; nd <9>
P. mandraspatensis L. 3; ar <15>, a <7>
P. maderaspatensis L. 3; a <6>
P. polygonoides Nutt. 3; a <27>
P. rotundifolius Willd. 3; a <7>
Genus Reverchonia
R. arenaria Gray 3; a <27>
Trib. PORANTHERAE
Genus Andrachne
A. cordifolia Muell.Arg. 3; a <27>
A. phyllanthoides (Nutt.) Muell.Arg. 3; a <27>
Genus Breynia
B. cernua Muell.Arg. 3; a <27>
B. nivosa (Bull.) Small 3; nd <9>
Trib. WIELANDIEAE
Genus Astrocasia
A. neurocarpa (Muell.Arg.) Johnst. 3; a <27>
Subfam. OLDFIELDIOIDEAE
Trib. HYAENANCHEAE
Genus Tetracoccus
T. hallii T.S.Brandeg. 3; a <27>
Subfam. ACALYPHOIDEAE
Trib. ACALYPHEAE
Genus Acalypha
A. bipartita Muell.Arg. 3; a <7>
A. claoxyloides Hutch. 3; a <6>
A. gracilens Gray 3; a <27>
A. hederacea Torr. 3; a <27>
A. indica L. 3; a <27>, a <6>
A. lindheimeri Muell.Arg. 3; a <27>
A. ostryaefolia Ridd. 3; a <27>
A. purpurascens 3; a <27>
A. radians Torr. 3; a <27>
A. rhomboidea Raf. 3; a <27>,nd <9>
A. virginica L. 3; a <27>
Genus Mallotus
M. paniculatus Muell.Arg. 3; a <27>
M. philippinensis Muell.Arg. 3; a <27>
M. polyandenus F.M.Fragm. 3; a <27>
Genus Mercurialis
M. annua L. 3; a <27>
M. perennis L. 3; a <27>
Genus Micrococca
M. mercurialis Benth. 3; a <7>
Genus Ricinus
R. communis L. 3; a <27>,c <8>,r <28>, a <6>
Trib. BERNARDIEAE
Genus Bernardia
B. myricaefolia (Scheele) Wats. 3; a <27>
B. obovata Johnst. 3; a <27>
Trib. CHROZOPHOREAE
Genus Chrozophora
C. tinctoria (L.) Adr. 3; a <27>
Genus Ditaxis
D. diversiflora Clokey 3; a <27>
D. humilis (Engelm. et Gray) Pax 3; a <27>
D. lanceolata (Benth.) Pax et Hoffm. 3; a <27>
D. neomexicana (Muell.Arg.) Heller 3; a <27>
D. serrata (Torr.) Heller 3; a <27>
Trib. DALECHAMPIEAE
Genus Dalechampia
D. ipomoeifelia Benth. 3; a <7>
Trib. PLUKENETIEAE
Genus Tragia
T. brevipes Pax 3; a <7>
T. geranifolia Kl. 3; a <27>
T. nepetaefolia Cav. 3; a <27>
T. ramosa Torr. 3; a <27>
T. stylaris Muell.Arg. 3; a <27>
T. urens L. 3; a <27>
T. urticifolia Michx. 3; a <27>
Subfam. CROTONOIDEAE
Trib. ALEURITIDEAE
Genus Garcia
G. nutans Rohr. 3; a <27>
Trib. CROTONEAE
Genus Croton
C. bonplandianus Baill 3; a <27>
C. californicus Muell.Arg. 3; a <27>
C. capitatus Michx. 3; a <27>
C. ciliato-glandulosus Ortega 3; a <27>
C. corymbulosus Engelm. 3; a <27>
C. engelmannii Ferg. 3; a <27>
C. glandulosus L. 3; a <27>
C. humilis S. 3; a <27>
C. insularis Baill 3; a <27>
C. linearis Michx. 3; a <27>
C. lindheimerianus Scheele 3; a <27>
C. monanthogynus Michx. 3; a <27>
C. neomexicana Muell.Arg. 3; a <27>
C. niveus Jacq. 3; a <27>
C. nummulariaefolius A.Rich 3; a <27>
C. torreyanus Muell.Arg. 3; a <27>
Genus Crotonopsis
C. elliptica Willd. 3; a <27>
Genus Ereocarpus
E. setigerus (Hook.) Benth. 3; a <27>
Trib. JOANNESIEAE
Genus Jatropha
J. angustidens (Torr.) Muell.Arg. 3; a <27>
J. dioicea Sesse 3; a <27>
J. macrorhiza Benth. 3; a <27>
J. spicata Pax 3; a <7>
J. stimulosa Michx. 3; a <27>
J. spathulata Muell. 3; nd <9>
Genus Mozinna(=Jatropha)
M. spathulata Ort. 3; a <27>
Trib. MANIHOTEAE
Genus Cnidoscolus
C. rotundifolius (Muell.Arg.) McVaugh 3; a <27>
C. texanus (Muell.Arg.) Small 3; a <27>
Trib. MICRANDREAE
Genus Havea
H. benthamiana Muell.Arg. 3; a <27>
H. brasiliensis (Willd. et Andr.Jass.) Muell.Arg. 3; a <27>
H. guianensis Aubl. 3; a <27>
H. nitada Muell.Arg. 3; a <27>
H. spruceana (Benth.) Muell.Arg. 3; a <27>
Genus Micrandra
M. lopezii R.E.Schult. 3; a <27>
M. minor Benth. 3; a <27>
M. siphonioides Benth. 3; a <27>
M. spruceana (Baill) R.E.Schult. 3; a <27>
M. sprucei (Muell.Arg.) R.E.Schult. 3; a <27>
Subfam. EUPHORBIOIDEAE
Trib. HIPPOMANEAE
Genus Homalanthus (=Carumbium)
H. populius Grah. 3; a <27>
Genus Maprounea
M. brasiliensis St.Hil. 3; a <27>
Genus Sapium
S. biloculare (Wats.) Pax 3; a <27>
Genus Sebastiania
S. pavoniana Muell.Arg. 3; a <27>
Genus Stillingia
S. dentata (Torr.) Britt. et Rushby 3; a <27>
S. linearifolia Wats. 3; a <27>
S. salicifolia (Torr.) Raf. 3; a <27>
S. sanguinolenta Muell.Arg. 3; a <27>
S. sylvatica L. 3; a <27>
S. texana Jhonst. 3; a <27>
Trib. EUPHORBIEAE
Genus Euphorbia
Subgen. ESULA
Sect. BALSAMIS
E. balsamifera Ait. 3; a <25>
E. calyculata Kunth 3;r <18>
E. dendroides L. 3; a <27>
Sect. LATHYRIS
E. lathyris L. 3; a <19>, a <27>
Sect. MYRSINITEAE
E. biglandulosa Desfontaines 3; a <25>
E. broteri Daveau 3; a <21>
Sect. ESULA
E. commutata Engelm. 3; a <27>
E. cyparissias L. 3; a <27>,c <8>
E. esula L. 3; a <27>,c <8>
E. peplus L. 3; a <27>
Sect. TITHYMALUS
E. helioscopia L. 3; a <27>
E. portulacoides L. 3;r <18>
E. purpurea (Raf.) Fern.
'E. darlingtonii Gray' 3; a <27>
E. spathulata Lam. 3; a <27>
'E. arkansana Engelm.' 3; a <27>
'E. dictyosperma Fisch. et Mey.' 3; a <27>
'E. missouriensis (Norton) Small' 3; a <27>
(sect. unknown)
E. arguta Banks et Sol. 3;r <28>
E. crenulata Engelm. 3; a <27>
E. palmeri Watson 3; a <27>
E. segetalis L. 3; a <27>
E. terracina L. 3; a <27>
Subgen. EUPHORBIA
Sect. TREISIA
E. bubalina Boiss. S;R <18>
Sect. EUPHORBIA
E. caducifolia Haines S;Mc <24>,D <17>, a <15>
E. grandidens Haworth S; n <11>
E. ingens Boiss. 3;m <23>
E. nivulia Hamilton S;R <18>
E. submammillaris (Berger) Berger S; nD <9>
E. trigona Miller S;R <18>
E. xylophylloides Lem. S; nD <9>
Sect. ELAEOPHORBIA
E. drupifera Schumacher S;R <18>
Sect. GONIOSTEMA
E. didierioides Leandri 3;r <18>
E. milii Moulins 3; n <11>,nd <9>
'E. splendens Boj.' 3; a <8>,n <11>
Sect. APHYLLIS
E. tirucalli L. S;R <1>
(sect. unknown)
E. caducifolia Haines S;R <28>
E. lactea Haw. 3;c <8>
E. lurida Engelm. ?; a <27>
E. submammillaris BRG. S; nD <9>
Subgen. POINSETTIA
E. cyathophora Murr. 3;r <18>
E. dentata Michx. 3; a <27>,c <10>,c <8>
E. eriantha Benth. 3; a <27>
E. heterophylla L. 3; a <27>,c <10>,c <8>
'E. geniculata Orteg' 3; a <7>
E. pulcherrima Klotzsch 3;r <1>,c <8>,n <11>
Subgen. AGALOMA
Sect. ZYGOPHILLIDIUM
E. henricksonii Johnston 3; a <26>
E. hexagona Nutt. 3; a <27>,c <10>,c <8>
Sect. CYTTAROSPERMUM
E. macropus (Klotzsch) Boiss. 3;r <18>
E. macropodoides Robinson et Greenman 3;r <18>
Sect. TITHYMALOPSIS
E. corollata L. 3; a <27>,c <10>,c <8>,r <1>,nd <9>
E. innocua Wheeler 3;r <18>
E. ipecacuanhae L. 3; a <27>
Sect. PETALOMA
E. bicolor Engerlm. 3; a <27>
E. marginata Pursh. 3; a <27>,c <10>,c <8>
Sect. TRICHEROSTIGMA
E. misera Benth. 3; a <27>
Sect. NUMMULARIOPSIS
E. peperomioides Boiss. 3; ar <18>
Subgen. CHAMAESYCE
Sect. SCLEROPHYLLAE
Subsect. GYMNADENIAE
E. clusiifolia Hook. et Arn. 4; aR <13>
'E. forbesii Sherff' 4; a <5>, aR <13>,C <8>
E. rockii Forbes 4; aR <13>
Subsect. SCLEROPHYLLAE
E. articulata Burman 4;R <18>, aR <14>
E. mesembryanthemifolia Jacquin 4;
'E. buxifolia Lam.' A <27>
'Cham. buxifolia (Hitchc.) Small' ACMk <16>
(subsect. unknown)
E. arnottiana Endlicher 4; aR <13>
'E. hookerii Steud.' AR <14>
E. atoto Forst. 4; a(124), aR <13>
E. atrococca Heller 4; aR <13>
E. celastroides Boiss. 4; aR <13>
'E. skottsbergii Sherff' AR <13>
E. degeneri Sherff 4; aR <13>
E. hillebrandii Leveille 4; aR <13>
E. multiformis Hook. 4; aR <13>
E. olowaluana Sherff 4; aR <13>
E. remyi Boiss. 4; aR <13>
Sect. CHAMAESYCE
Subsect. ACUTAE
E. georgei Oudejans
'E. acuta Engelm.' 3; a <27>,r <18>
E. angusta Engelm. 3;r <18>
E. lata Engelm. 4; a <27>,R <18>
Subsect. HYPERICIFOLIAE
E. hirta L. 4; a <27>,C <20>, aR <15>, a <6>
E. hypericifolia L. 4;C <20>
'E. glomerifera (Millspaugh) Wheeler' A <12>
E. nutans Lagasca 4; a <25>, a <27>
'E. maculata L.' AMk <4>,C <19>, a <27>,C <8>
Subsect. CHAMAESYCEAE
E. carunculata Waterfall 4;R <18>,#a <27>
E. chamaesyce L. 4; a <27>
E. drummondii Boiss. 4; a <19>
E. glyptosperma Engelm. 4; a <27>,C <19>,C <10>,C <8>
E. granulata Forsk. 4; a <25>, aR <15>, a <7>
E. maculata L. 4;E <22>
'E. supina Raf.' AMk <4>,C <19>,C <8>,C <10>, a <27>
E. missurica Raf. 4; a <27>,C <10>,C <19>,C <8>
'E. petaloidea (Engelm.) Small' A <27>
'E. zygophylloides Boiss.' A <27>
E. peplis L. 4; a <21>, a <25>
E. polygonifolia L. 4; a <27>
E. prostrata Ait. 4; a <7>
'Cham. prostrata (Ait.) Small' A <2>
E. thymifolia L. 4; a <3>
E. wheeleri Baillon 4; a <19>
Subsect. PLEIADENIAE
E. caecorum Boiss. 4;R <18>
E. viscoides Boiss. 4;R <18>
(subsect. unknown)
E. albomarginata Torr. et Gray 4; a <27>
E. arizonica Engelm. 4; a <27>
E. fendleri Torr. et Gray 4; a <27>
E. geyeri Engelm. 4; a <27>,C <8>
E. hooveri Wheeler 4; a <27>
E. humistrata Engelm. 4; a <27>
E. melanadenia Torr. 4; a <27>
E. micromera Engelm. 4; a <27>
E. parryi Engelm. 4; a <27>
E. pediculifera Engelm. 4; a <27>
E. serpens Kunth 4; a <27>
E. serpyllifolia Pers. 4;C <19>,C <10>, a <27>,C <8>
'E. hirtula Engelm.' A <27>
'E. neomexicana (K. et G.) Wheeler' A <27>
E. setiloba Engelm. 4; a <27>
E. stictospora Engelm. 4; a <27>
E. vermiculata Raf. 4; a <27>
'E. hirsuta (Torrey) Wiegand' 4; a <27>
(sect. unknown)
E. australis Boiss. 4; aR <3>
E. capitellata Engelm. 4; a <27>
E. cinerascens Engelm. 4; a <27>
E. densiflora (Klotzsch) Klotzsch 4; a <12>
E. florida Engelm. 4; a <27>
E. forskalii G.Jay 4;R <28>
E. heineana Sprengel
'E. microphylla Roth' 4; aR <15>
E. hyssopifolia L. 4; a <27>
E. inaequilatera Sonder 4; a <7>
E. revoluta Engelm. 4; a <27>
E. scordifolia Jacq. 4;R <28>
E. serrula Engelm. 4; a <27>
E. stodartii Fosberg 4; a <6>
E. taitensis Boiss. 4; aR <13>
E. tomentulosa Wats. 4; a <27>
E. villifera Scheele 4; a <27>
(subgen. unknown)
E. brachycera Engelm. ?; a <27>
E. chamaesula Boiss. ?; a <27>
E. crotonoides Boiss. ?; a <7>
E. hildebrandtii Baill. 4; a <27>
E. parishii Greene ?; a <27>
E. philora (Cockrell) Tidestrom
'E. montana Engelm.' ?; a <27>
E. platyphyllos L. ?; a <27>
E. pyrifolia Lam. ?; a <6>
E. robusta (Engelm.) Small ?; a <27>
E. schizoloba Engelm. ?; a <27>
'E. incisa Engelm.' ?; a <27>
E. schimperiana Scheele ?; a <7>
E. squamigera Lois.
'E. rupicola Boiss.' ?; a <27>
E. tetrapora Engelm. ?; a <27>
E. waldsteinii(Sojak) Radcliff-Smith
'E. virgata Waldst. et Kit.' ?; a <27>
Genus Monadenium
M. lugardiae N.E.Br. S; nD <9>
Genus Pedilanthus
P. tithymaloides (L.) Poit. ?; a <6>,Dn <9>,R <1>
Genus Synadenium
S. cupulare (Boiss.) A.White S; nD <9>
S. grantii Hook.f. S; nD <9>