A Taxonomic Study of Asarum sect. Asiasarum (Aristolochiaceae) in Japan

2007·0.84 MB·English

by  Hiroki Yamaji

植物研究雑誌
J.Jpn.Bot.
82:79–105(2007)
A Taxonomic Study of Asarum sect. Asiasarum (Aristolochiaceae) in Japan
Hiroki YAMAJIa,d, Teruko NAKAMURAb, Jun YOKOYAMAc, Kenji KONDOd,
Takashi MOROTAd, Shuichi TAKEDAd, Hiroshi SASAKId and Masayuki MAKIc
aBiologicalInstitute,GraduateSchoolofScience,TohokuUniversity,
Aoba,Sendai,980-8578JAPAN;
bFacultyofPharmaceuticalSciences,TokyoUniversityofScience,
2641Yamazaki,Noda,278-8510JAPAN;
cDivisionofEcologyandEvolutionaryBiology,GraduateSchoolofLifeSciences,TohokuUniversity,
Aoba,Sendai,980-8578JAPAN;
dResearchDivision,Tsumura&Co.,3586,Yoshiwara,Ami,Ibaraki,300-1192JAPAN
(ReceivedonJune2,2006)
ThreespeciesandonevarietyofAsarumsect.Asiasarumhavehithertobeenknown
in Japan. They are reclassified into seven species including three new species: A.
maruyamae,A.tohokuense,andA.mikuniense.Mostofthetaxonomicstudiesonthesec-
tiongaveweighttothecharactersofleaves,calyxlobes,andthenumberofstamensand
pistils.However,thisstudyisbaseonthefollowingcharacterstodistinguishthespecies:
color pattern in the adaxial surface of calyx tube, the shape of calyx tubes, the width of
calyx tube throat, the shape of calyx lobe, and the number and height of ridges on inner
surface of calyx tubes. The distribution range of the seven species are geographically
segregated. Asarum sieboldii is distributed in central and western Honshu, as well as
Korea and central to southern China; A. misandrum in Kyushu, the Aso Mountains as
well as southern Korea; A. heterotropoides var. heterotropoides in northern Honshu,
Hokkaido,southernSakhalinandKuriles;A.tohokuenseincentralandnorthernHonshu;
A. mikuniense in central Honshu; A. maruyamae in western Honshu; A. dimidiatum in
western Honshu, Shikoku and Kyushu.
Key words: Asarum sect. Asiasarum, distribution, Japan, new species.
Asarum L. consists of about 80 species of and sect. Heterotropa (Morr. & Decne) A.
rhizomatous herbs distributed in eastern Braun (Kelly 1998).
Asia, Europe and North America (Kelly Asarum sect. Asiasarum comprises six
2001). Based on cladistic analyses of mor- species and three varieties and occurs in
phological data and internal transcribed northeastern Asia; Japan, China, Korea and
spacer (ITS) sequences of nuclear ribosomal eastern Russia (Maekawa 1936b, Cheng and
DNA (Kelly 1997, 1998), Kelly (1998) Yang 1983, Yamaki et al. 1996, Watanabe
proposed an infrageneric classification sys- 1996, Oh et al. 1997, Lee and Lee 2000). In
tem of Asarum. The genus is divided into Japan, totally four taxa (three species and
two subgenera, subgenus Asarum and subge- one variety) have been recorded so far:
nus Heterotropa (Morr. & Decne.) O. C. Asarum sieboldii Miq., A. heterotropoides F.
Schmidt, and subgenus Heterotropa into two Schmidt var. heterotropoides and var.
sections: sect. Asiasarum (F. Maek.) Araki mandshuricum (Maxim.) Kitag., and A.
—79—
80 植物研究雑誌 第82巻 第2号 平成19年4月
dimidiatum F. Maek. (Maekawa 1956, Ohwi lobes; form L3 has larger, pentagonoid calyx
1965, Satake and Momiyama 1982, lobes. These forms were also supposed to be
Hatusima 1993). Most of the precedent taxo- different in pollen, and chemical component
nomic studies of the section gave weight to characters (Nakamura 1986, Nakamura and
the characters of leaves, calyx lobes, and Nagasawa 1987, Nakamura et al. 1979,
number of stamens and pistils. 1982, 1987).
Yamaji et al. (2007) revealed that eight Theaimofthisstudyistoclarifythetaxo-
distinct forms were recognized based on nomic position of the eight forms. For this
morphological examinations. In this study, purpose, we examined specimens deposited
each form was recognized based on a multi- in herbaria, and compared with the taxa hith-
variate analysis of various floral characters erto described in Japan and the other areas.
and examination of new floral characters Consequently we revised the taxonomy of
together with those used in the previous Asarum sect. Asiasarum of Japan.
works. First, two color patterns in the inner
surface of calyx tube were found (form D Identification of herbarium specimens
and L). Form D is dark purple, on the other About 500 fertile specimens in the
hand, form L is dark purple proximally, herbaria: KYO, MAK, TI, THS, and TUS
ivory white, yellowish green, light purple, or including about 100 specimens collected by
their intermediate in the middle part, dark the authors were also examined (see each
purple or rarely ivory white at the throat. description of the following taxonomic treat-
Form D is further divided by the cell number ment).Throughthecomparisonamongfresh,
of trichomes on the inner surface of calyx fixed,andpressedanddriedflowersfromthe
tube and adaxial surface of calyx lobes; form populations examined in Yamaji et al.
D1 has single-cell trichomes; forms D2 and (2007), we found that pressed flowers in
D4 have three-cell trichomes; form D3 has herbarium specimens were enough to iden-
seven to ten-cell trichomes. Forms D2 and tify each form recognized in this study
D4 are further divided by the number of sta- except for difficulties of discrimination be-
mens and pistils; form D2 has the regular tweenformsD1andD2(Fig.1).Aconsider-
number of them in the sect. Asiasarum: 12 able number of specimens keep their floral
stamens and six pistils while form D4 has color pattern as well as the number of ridges
half of them: six and three, respectively. of the inner surface of the calyx tube, calyx
Form L is further divided by the shape of the lobe shape/condition, and length/inclination
calyx tube. Form L4 has a round, elliptical, of pistil protuberances. Moreover, from the
urceolatecalyxtube;itslengthisaboutahalf total size/shape of flower, and basal width of
of the external diameter. On the other hand, calyx lobe, the size/shape of the calyx tube
forms L1, L2, and L3 have round urceolate including calyx tube throat width were able
calyx tube; its length is more than a half of to be anticipated.
the external diameter. Forms L1, L2, and L3 However, we found specimens with unsta-
are identified by the width of calyx tube ble number of stamens and pistils, 6–12 and
throat. Forms L1 and L3 have narrow throat, 3–6, respectively (e. g., H. Yamaji 8006,
less than a half in diameter to external TUS 284589, THS 72985). These specimens
diamete, on the other hand, form L2 have belong to forms D1, D2, and D4. For such
wide throat, more than a half in diameter to plants, Nakamura et al. (1982) showed that
external diameter. Forms L1 and L3 are the number of stamens and pistils were un-
identified by the size and shape of calyx stable,andvaried6–12and3–6,respectively
lobes; form L1 has smaller, triangular calyx in a restricted area of central Kyushu. As the
April2007 JournalofJapaneseBotanyVol.82No.2 81
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April2007 JournalofJapaneseBotanyVol.82No.2 83
area is surrounded by the distribution of The other specimens from Sakhalin were
form D4, and as Nakamura et al. (1982) also identified as form L1.
reported that such plants also reduced the Type specimen of A. dimidiatum was col-
number of stamens and pistils under culti- lected in Nagasaki, Kyushu and deposited in
vated condition, we concluded that these the University of Tokyo (TI). Though its re-
plants should be included in form D4. duced stamens and pistils were not recon-
The number of specimens identified into firmed in this specimen, Maekawa (1936a)
each form was as follows: form D1(cid:1)7, D2 recorded that its number of stamens and pis-
(cid:1)9,D3(cid:1)2,D4(cid:1)35,L4(cid:1)2,L1(cid:1)138,L2(cid:1) tilswere6and3,respectively.Therefore,we
220, and L3(cid:1)6. In addition to these, 118 confirmed that A. dimidiatum corresponds to
specimens could not be identified into one form D4.
form but be narrowed down to forms D1 and In Japan, A. heterotropoides var. mand-
D2. Because the most distinct discriminant shuricumisonlyreportedtobedistributedin
character between forms D1 and D2, the the restricted area of central Kyushu
trichome condition of flowers, was espe- (Hatusima 1993, Azegami 1996). This taxon
cially fragile, it was difficult to discriminate is also distributed in Far East Russia,
them among the herbarium specimens. Northeastern China (Maximowicz 1872,
Highly discolored or collapsed specimens Cheng and Yang 1983), and Korea (Yamaki
could not be identified to any forms. et al. 1996). Its type specimens were
collected in Far East Russia, and are
Comparison with taxa hitherto recognized deposited at V. L. Komarov Botanical
in Japan Institute (LE). The distribution of this taxon
We compared the eight forms with taxa in Japan and its characteristics reported by
belonging to sect. Asiasarum hitherto known Hatusima(1993)completelycorrespondedto
in Japan. that of form L4. However, the typical form
Asarum sieboldii was obviously hitherto of the taxon on the continent, including the
recognized as a species including forms D1, types is clearly distinct from form L4 in
D2, L2, and L3 by its distribution and the comparatively large, pentangular, gently
diagnostic characters described previously recurved or patent calyx lobes, compara-
(Maekawa 1936b, Ohwi 1965, Satake and tively dark purple calyx, and pilose petioles
Momiyama 1982). Examination of the type andtheabaxialsurfaceoflaminae.Therefore
specimen of A. sieboldii in the National the attribution of form L4 is examined in the
Herbarium Nederland, Leiden University following chapter.
Branch (L) showed that the type is identified Forms D3, L2, and L3 did not corre-
as forms D1 or D2 by its calyx tube: com- sponded to any taxa which have been hith-
paratively shallow, with wide throat, and erto recorded in Japan.
dark colored inner surface. In addition, this
specimen is supposed to have been collected Comparison with taxa and populations
in the west of central Honshu (Nakamura outside Japan
and Ômori 1990). We compared the eight forms with the
The type specimen of A. heterotropoides taxa and populations outside Japan. About
var. heterotropoides was collected in south- 110 specimens from outside of Japan were
ern Sakhalin, and deposited at V. L. also examined (Appendix).
Komarov Botanical Institute (LE). It was As mentioned above, Asarum hetero-
identified as form L1 by its reflexed, small tropoides var. mandshuricum did not corre-
calyx lobes, and round, urceolate calyx tube. spond to the form L4. On the other hand, A.
84 植物研究雑誌 第82巻 第2号 平成19年4月
misandrum B. Oh & J. Kim, which was than those of forms L1 and L3, 1–3 cm and
described from southern Korea, is similar to 1–2 cm, respectively, though those of form
form L4 in having strongly recurved and tri- L1 rarely had peduncles up to 6 cm.
angular calyx lobes and glabrous petioles Consequently, we decided forms L1 and L3
(Oh et al. 1997) and yellowish green pistil as distinct from A. versicolor in this study.
protuberances.WeconcludedthatformL4is Asarum patens (Yamaki) Y. Lee is unique
identified as A. misandrum. in having long, protruded pistil protuber-
Asarum sieboldii f. seoulense (Nakai) C. ances, comparatively large and patent calyx
Y. Cheng & C. S. Yang, which is also dis- lobes, and densely pilose leaves in adaxial
tributed in Northeastern China and Korea surface. No individuals corresponding to this
(Cheng and Yang 1983, Yamaki et al. 1996). species were found in Japan.
Thistaxonisuniqueinhavingdenselypilose
petioles and the abaxial surface of laminae, Conclusion
which was not detected in sect. Asiasarum Among the eight forms, three forms have
plants in Japan. Therefore we concluded that not been described previously.
no Japanese plants correspond to this taxon. The precedent systems of Asarum sect.
Individuals recognized as A. sieboldii in Asiasarum (Maekawa 1936b) applied hierar-
Korea were almost similar to form D1 in this chical classification including intraspecific
study in the floral color pattern, the number taxa, and weighted each character with
of stamens/pistils and the trichomes of flow- somewhatsubjectivestandardtoestimatethe
ers. In addition, they completely agreed with relationship of the taxa. However, such
the form D1 in chemical and pollen charac- hierarchical classifications may force to
ters (Yamaji et al. unpubl. data). Individuals inferencesofphylogeneticrelationshipseven
recognized as A. sieboldii in southern China if they are not yet supported by DNA evi-
corresponded to forms D1 or D2. dence. Progress of molecular phylogenetics
Asarumversicolor(Yamaki)Y.Leeisdis- in the last decade promoted systematic study
tributed in central Korea, and is character- to infer the phylogeny with molecular infor-
ized by variegated leaves with some mation.
exceptions, smaller flowers than A. sieboldii In this study, we intended to recognize
(Yamakietal.1996,LeeandLee2000).The morphologically distinct forms of this group,
shape and size of flowers of this species is and to regard the recognized forms as hypo-
similar to forms L1 and L3, especially in the thetical taxonomic units for the following
calyx tube with a narrow throat. Calyx lobes study, e. g., a molecular phylogenetic study.
of this species are patent or oblique, undu- Therefore, we do not discuss about phylo-
late, acuminate at apex, and are similar to a genetic relationship between the forms
part of form L1. Floral color pattern of this recognized in this study. We equally treated
species is also similar to forms L1 and L3: the forms as distinct species.
ivory to light purple, olive green to purple in Exceptionally, we decided to include
outer surface, dark purple at the throat, ivory forms D1 and D2 into the same species.
white to ivory, sometimes light purple at the Though our analysis indicated that these two
middle part, dark purple proximally, ivory forms are different in floral characters by the
white at the base. However, the number of CVAs, and they are supposed to have almost
cells of each trichome of flowers is almost disjunct distributions from each other, dis-
three in A. versicolor though those of form crimination of these forms in herbarium
L1 and L3 are 3–7 and 5–7, respectively. specimens is almost impossible. In the same
Peduncles of this species are longer, 3–6 cm, way, the type specimen of Asarum sieboldii
April2007 JournalofJapaneseBotanyVol.82No.2 85
Fig.2. Distributionsofeightforms(sevenspecies)ofAsarumsect.AsiasaruminJapan.A:FormDspecies.
B:FormLspecies.*Specimensthatcouldnotbeidentifiedintooneformbutbenarroweddowntoform
D1andD2.
is impossible to place into either form. distributed in the west of central Honshu,
Hence we suspended the taxonomic treat- Sadogashima Is. and Tsushima Is. Asarum
ment of two forms and treat them as a single maruyamae is distributed in the restricted
taxon. area of western Honshu. Asarum dimidiatum
Consequently, we recognized the follow- is distributed in Shikoku, Kyushu and two
ing seven species in Asarum sect. Asiasarum restricted areas of western Honshu. Asarum
in Japan; forms D1 and D2(cid:1)A. sieboldii, misandrum is only recognized in the
form D4(cid:1)A. dimidiatum, form L1(cid:1)A. restricted area of central Kyushu. Asarum
heterotropoides var. heterotropoides, form heterotropoides var. heterotropoides is dis-
L4(cid:1)A. misandrum; three new species de- tributed in Hokkaido, northern Honshu,
scribed here are as follows: form D3(cid:1)A. Sakhalin, and the southern Kuriles. Asarum
maruyamae, form L2(cid:1)A. tohokuense, and tohokuense was distributed in northern and
form L3(cid:1)A. mikuniense. central Honshu and Sadogashima Is. Asarum
The distribution of each form in Japan is mikuniense is distributed in the restricted
shown in Fig. 2. Asarum sieboldii is area of central Honshu.
86 植物研究雑誌 第82巻 第2号 平成19年4月
d
e n
A.misandrum(FormL4) straightorzigzag salmonpinktopalepurpleivorywhitetoivory,withdarkpurplestrip11.5–165–87.5–11 1.0–1.519–27 deltoid acutestronglyrecurved4.0–7.59.5–11 3–470–100 1.0–1.7closedyushu(Kumamoto)asouthernKorea
K
A.mikuniense(FormL3) straight olivegreen–purple vorywhite–palepurple,withdarkpurplestripe11–146.5–105–8 0.6–1.217–21 pentangular acuminatepatent–oblique7–118–11 5–7120–180 0.6–1.5closed–detachedCentralHonshu
i
ciesinJapan A.tohokuense(FormL2) straight ivory–palepurple ivorywhite–palepurple,withdarkpurplestripe10–156.5–10.55.5–11 0.4–1.015–21 transverseovatusdeltoidacuminateoblique5.0–9.57.0–12.0 3–460–120 0.4–1.0detachednorthofcentralHonshu
e
sect.Asiasarumsp A.heterotropoides(FormL1) straight pink–darkpurple ivorywhite–palepurple,withdarkpurplestripe10–155–104–7.5 0.6–1.715–27 deltoid acute–acuminaterecurved–patent3.5–9.05.0–10.5 3–780–150 0.5–1.6closedHokkaido,northeastHonshu,SakhalinandsouthernKurileIslands
parisonofAsarum A.dimidiatum(FormD4) straight olivegreenwithsmallpurplespotsdarkpurple 8.5–125–85.5–9.0 0.7–1.315–24 transverseovatusdeltoid–pentangularacuminatepatent–oblique5.5–96.5–10.5 3–680–140 0.8–1.6detachedWesternHonshu(Nara,Hiroshima),ShikokuandKyushu
m
o all
Table1.C A.maruyamae(FormD3) straight egreenwithsmpurplespotsdarkpurple 10–147.5–105–7 0.7–1.015–17 pentangular acuminatepatent–oblique5.5–96–9 7–10220–240 0.8–1.1closedWesternHonshu:ShimanePref.
v
oli
A.sieboldii(FormsD1,D2) straight olivegreenwithsmallpurplespotsdarkpurple 9.5–155–9.56–11.5 0.7–1.717–26 transverseovatusdeltoid–pentangularacute–acuminaterecurved–oblique4.5–13.57.0–14.5 1–520–140 0.7–1.9detachedCentralandWesternHonshu,KoreaandsouthofcentralChina
e
c
a
urf
RhizomeCalyxtubeColorpatternOutersurface InnersurfaceSize(mm)DiameterLengthThroatwidthLongitudinalridgesHeight(mm)NumberCalyxlobeShape ApexDirectionLengh(mm)Width(mm)TrichomesinadaxialsNumberofcellsLength(µm)StyleprotuberanesLength(mm)TipconditionDistribution
April2007 JournalofJapaneseBotanyVol.82No.2 87
Comparison of the recognized species in solitary, actinomorphic. Peduncle erect or
this study is shown in Table 1. For taxo- oblique,glabrous.Sepals3,connate,forming
nomic key to the taxa, in addition to the calyxtube;externalsurfaceglabrous,striated
qualitative characters, useful quantitative longitudinally; inner surface glabrous or pu-
characters recognized in the univariate bescent with trichomes, 15–27 ridged. Calyx
analyses (Yamaji et al. 2007) were also lobes recurved, patent, or obliquely erect;
adopted. adaxial surface glabrous or pubescent with
trichomes; abaxial surface glabrous. Stamens
Taxonomic treatment 6 or12, or variable 6–12 in two whorls; fila-
Asarum L. ment greenish ivory or light purple, anther
Subgen. Heterotropa (Morr. & Decne.) extrorse, pale yellow. Ovary half inferior or
O. C. Schmidt superior, conic, slightly ribbed, 3–6-locular.
Sect. Asiasarum (F. Maek.) Araki in Acta Styles 3 or 6, or variable 3–6, free, with
Phytotax. Geobot. 6: 125 (1937) – L. Kelly horn-like stylar protuberances at apex; stylar
in Amer. J. Bot. 85: 1467 (1998) – protuberances bifulcated, greenish ivory or
Asiasarum F. Maek. in Nakai, Fl. Syl. Kor. dark purple, 0.4–3.0 mm long, close or
21: 17(1936). TYPE: Asarum sieboldii Miq. detached each other, ovules 4–10 in each
Perennial herb. Rhizome about 5 mm in locule.
diameter, glabrous, zigzag or straight, slen-
der, with leaf and flower trace, sometimes Key to the Species of
branched. Roots usually about 1 mm in di- Asarum Sect. Asiasarum in Japan
ameter, fleshy, slender, distributed evenly 1. Inner surface of calyx tube entirely dark
along rhizomes. Hibernaculum at the axil of purple
leaves and cataphylls, composed of cata- 2. Calyx tube roundish urceolate, 0.7–0.9
phylls,preformedleavesandflower,unequal times longer than the external diameter;
ovate or elliptic, obtuse, acute or acuminate throat about half in diameter to external
at apex, olive green, purplish green, or pur- diameter;trichomesonadaxialsurfaceof
plish ivory, with purple veins, 10–16 mm calyx lobe and tube 220–240 µm long,
long, 5–10 mm wide in hibernaculum with composed of 7–10 cells; pistil
floral bud. Cataphylls 3, caducous, glabrous, protuberances generally greenish ivory
ciliate, triangular ovate, obtuse, acute or (Shimane/Honshu) .... 1. A. maruyamae
acuminate at apex; proximal cataphyll: 1–3 2. Calyx tube cylindroid or urceolate, 0.5–
mm long, 2–6 mm wide, median cataphyll: 0.7 times longer than the external diame-
4–14 mm long, 6–12 mm wide; distal ter; throat more than half in diameter to
cataphyll: 8–18 mm long, 7–20 mm wide. external diameter; trichomes on adaxial
Leaves deciduous, solitary in shoot without surface of calyx lobe and tube 20–140
flower, two, alternate in flowering shoot, µm long, composed of 1–5 cells; pistil
petioles 3–20 cm long, adaxial ridges protuberance purple at apex
sparsely pilose, the other part pilose or 3. Stamens and pistils 12 and 6 (west of
glabrous; lower lamina ovate cordate–broad- central Honshu, Tsushima, Korea, and
ly cordate; upper lamina pentangular ovate southeast China) ........... 2. A. sieboldii
cordate; membranous, palmately veined; 3. Stamens and pistils usually 6 and 3,
abaxial surface glaucous, glabrous or pilose; sometimes 6–12 and 2–6 (Honshu
adaxial surface green, variegated or non- (Hiroshima, Nara), Shikoku, Kyushu)...
variegated, sparsely pilose; margin ciliate, ................................... 3. A. dimidiatum
the cilia incurved toward the apex. Flower 1. Inner surface of calyx tube dark purple or
88 植物研究雑誌 第82巻 第2号 平成19年4月
Fig.3. Comparison of form D taxa: A: Asarum maruyamae (H. Yamaji 7002). B: A. sieboldii (H.
Yamaji 6051). C: A. dimidiatum (H. Yamaji 8002). ov: Overlook view. up: Upper view. ls:
Longitudinalsection.p&o:Pistilandovary.ts:Transversesectionofcalyxlobe.Bar(cid:1)5mm.
VoucherspecimensaredepositedatTUS.
rarely ivory white at throat, ivory white, wide, pentagonoid, patent or oblique,
ivory, or pale purple at middle, dark pur- acuminate at apex (central Honshu) ....
ple proximally ................................... 6. A. mikuniense
4. Throat narrow, less than half diameter of 4. Throat wide, more than half diameter of
external diameter external diameter
5. Calyx lobe 4–8 mm long, 5–9 mm 6. Calyx tube round urceolate, more than
wide, triangular, reflexed in most flow- half as long as external diameter; calyx
ers,sometimespatentoroblique(north- lobes patent or oblique, acuminate at
ern Honshu, Hokkaido, Sakhalin and apex (northern and central Honshu) ....
the Kuriles) ....... 4. A. heterotropoides .................................... 5. A. tohokuense
5. Calyx lobe 7–10 mm long, 8–10 mm 6. Calyx tube round elliptical urceolate,