The Distribution of Dryopteris caudipinna (Dryopteridaceae), a Sexually Reproducing Counterpart of the Apogamous D. erythrosora, in Japan

2010·8.8 MB·English

by  Yamamoto, Kaoru

TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
ISSN 1346-7565 Acta Phytotax .Geobot, 61 (2 )1:09-114 (2010)
SHORTCOMMUNICATION

The Distribution (DryopteridaceSaeexjua,lly
ofDi:yopteris caudipinna a
Reproducing Counterpart of the Apogamous D. ei:J,throsora, in Japan
KAoRu YAMAMoToi･ ',NORIAKI MURAKAMI' AND ATsusHI EBIHARA2
iMZikino Herbarium, Graduate Scheol ofScien ancdEengineering, 7btvo hetropolitan U}iiversiijL
1-1Minami-osawa,Hlaehicu7'bik,[email protected],ip

(lauthorf coorrrespondencel; iDepartment ofBotat DNlbtiona iMiiseum of?Vbtu arned Science,
4-1- iAmakubo, tsukuba ,Ibarak i305-OO05 ,.ltu)an
The distribut iofoDntlyopteris caudipinna, a sexually reproducing counterpart ofthe apogamous species
D, ecythrosera, was investigat beyd counting the number of spores per sporangium in a larg enumber of
herbarium speeimens. We discovered a number ofnew localitie fsor D, caudipinna, including most ofthe
major island isn the Izu Island arc (Toky Porefeeture) ,and in Chiba ,Aichi, Tottori ,Yamaguchi, Kochi,
Nagasaki, and Fukuoka prefectures ,The habita otfD. caudipinna tends to be island asnd coastal areas in
warm temperate regions under the influenc oefwarm currents.
Key words: apogamy, Dryopteridaceae,Dt:yopt ceaurdiipsinna, Izu Islands,spores
Di:yqpteris (D.C.Eaton)Kuntze,
ei:ythrosora recognized as a sexual counterpart ofD, er:ythro-
and it sallies (Dtlyqpt seecrti. PsoL}tstichod rdiis-) sora, but was thought to be a relative with narrow
versified in the Japanes earchipelago, where there pinnule swhen Nakai (193 1d)escrib eidt S.ubse-
are more than 25 taxa (Iwats u1k9i95) ,including quent studies revealed that D. er:ythrosora is an
apogamous triploids (Takami y1a996). Only two apogamous triploid (Kuri t19a61) ,and that plants
sexual diploi dspecies, which may have been in- morphologically corresponding to D, caudipinna
volved in the formatio nofthe apogamous species are sexual diploid (sHirabaya s1h9i70). Dr:yqpter-
complex, are recegnized in Japan :D, caudipinna is caudipinna tends to have narrower pinnules
Nakai, originarly described from Hachijojima, than D. er:ythrosora, but distinguishi nigt from
and D. koideumiana Tagawa from the Ryukyu Is- the apogamous D. e":ythrosora is not always suc-
lands .Information on their distributi oesnpe,cial- cessfu1 (Yamamot &o Nakaike 1983 )as apoga-
ly of the fbrmer ,has been documented only spo- mous individua lssometimes have a narrow pin-
radically or remains unpublished. The number of nules and appear similar to D, caudipinna. Reli-
spores per sporangium can be an indicato orf the able identificat iofen D, caudipinna, therefore,
reproductive mode ofmost leptosporangi afteerns should be based on counting the number of spores
(Takamiy a1996); the presence of 64 spores usu- per sporangium, counting chromosomes, or de-
ally suggests sexual reproduction, while 32 termining the size ofthe genome, In other words,
spores suggests apogamy. In this study, we up- the taxon is define dby it sreproductive mode, not
dated infbrmati roenlated to the distributi oofn by morphological characters,
these two species mainly by counting the number Previousl yknown localiti efsor Di:yopteris
of spores per sporangium on herbarium speci- caudipinna, confirmed by spore or chromosome
includeTsukuba-shrinein Ibaraki Pre-
mens. counts,
Originall MDr:yopteri scaudipinna was not fecture (Ftijim o19t9o8), Jinmoji in Kanagawa
NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
110 Acta Phytotax .Geobot. Vol. 61
1 2
e
'lv.
.
1oo "m -1OO
FIG. 1. Example of64 spores per sporangium, indicatin sgexual reproduction (S ,7lsuts u1i7349, TNS LVS-57360]]).
FIG. 2. Example of32 spores per sporangium, indicatin agpogamous reproduction (S ,Koba.vash i2993, TNS [VS-573459]).
Prefecture (Yamamot &o Omori 1991) ,Oshima was determined under a stereomicroscope for all
(Nakaik e& Yamamoto 1984) and Hachijojima candidates, unless they were uncountable. For
(Hirabaya s1h9i70, Yamamoto & Nakaike 1983) specimens from the Izu Island se,xcept Oshima
inIbkyo PrefectureK,anmuri IslandinKyoto Hachljojima,for
and which comprehensive spore
Prefectur (eMitsu d1a990) ,the Shimane Penin- counts had alreadybeen made, all ofthe available
sula in Shimane Prefectur e(Miyamo t&o Naka- specimens were examined to determine the gen-
mura 1990) ,and Kirishima in Miyazaki and Ka- eral ratio of sexual to apogamous indiyidua lons
goshima Prefectur e(sMiyamo t&o Nakamura the islands.
1990) A.ccording to Iwatsuk i(1995 t)h,e distribu-
tion of D, caudipinna is confined to the Izu Is- Results
land sand the Miura and Izu peninsulas ,Al-
though both Hirabayashi (1974 a)nd Miyamoto & The spore number per sporangium was deter-
Nakarnura (1990 p)ublishe ddistribut imoapns of mined fbr approximately 500 sheets of the select-
D. caudipinna, i tis unclear whether they exam- ed specimens. Among them, 241 specimens had
ined spores or counted chromosomes, as they did 64 speres per sporangium while about 50 speci-
not state the sources of their data. mens had 32 spores per sporangium. The spores
of the remaining specimens were uncountable.
Materials and Methods Sporangi ataken from the fbllowin gspecimens
contained 64 spores; their morphological charac-
Candidate specimens of D-,opteris caudi- ters matched those of typical D. cattdipinna.
pinna, based on frond shape, were selected from Specimens from previously known localit iaerse
approximately 3,500 sheets ofD. et;vthrosora and excluded from the list
D. caudipinna deposited in the National Museum
of Nature and Science (TNS) and the Makino JAPAN. Honshu. Ibaraki Pref: Nishiibaraki-gun,
Herbarium of Tokyo Metropolita nUniversity I[wVamSa--m9a3ch4i5,2 [3YaM]ma)ank;ea,b e19-8g1-Mulanlk-]a0b8 ,eM-m a}e2hiisu ,sS,an.k a(yToNrSi-
(MAK).The
number of spores per sporangium Shiio .Yakuoin, 1973-12-02, H. Ito s.n, (TNS [VS-
NII-Electronic Library Service
TThheJea paJnaespeSaoncieestyefo rSociety for PPllanatnSyts teSmaytsictsematics
November 2010 YAMAMoTo & AL.-Distributien ofDrlyopteriscaudipinna 111
'
/1-
Ilt//ti.
tll
J
I.:,
1
:F,".:t:=t.::-:Stn!y.r .--r IF-iTF.t-P. V- . :.'- j,..L--1･l{Fl-t1-
,,,.,. e -r- -t･ --. .t.t t ., t ･ t ･ t. ･ tt t t ni.-tpttt/.t. nr-pt;')-･th-t"-
Tms lvt( li:Tt-
a
5
-
''/,'i/"'''
w
g
el
l･1P
tl]i.b,/
FIGS.3-6, Representativ especimens ofDr.t,opteris caudipinna, 3. Ibaraki Prefecture T,sukuba-shi. Tsukuba Shrine, 1987-08-
23, S. ,ldintsumoto s.n.. Iivin gcollection ofthe Tsukuba Botanical Garden 77398, (TNS [VS-9S0825 74]. )Ai.chi Prefec-
ture. Atsumi-gun, Atsumi-cho. Yamada, 1980-11-23 ,K Jnukai 420J {TNS [VS-550286] 5).. Nagasaki Prefecture, Tsu-
shima lsland ,Kamiagata-gun. Kamiagata-cho, Mitake. altitude 80-90 m, 1979-07-19 .S. ftutsui i7349 (TNS [VS-
573601i) .6. Kagoshima Prefecture .Aira-gun, Kamo-cho. Yadome-dake, 1983-07-19, ,e dKiawabata s.n. (TNS [VS-
742741]),
NNIII-IE-leEcltreoncitcronic Library Service
TThheJea paJnaespeSaoncieestye Society ffoorr PPllanatnt SSyystsetmaetmicastics
112 ActaPhytotax.Geobot. Vol,61

tL..'t...'L .,t) . ,.tl.
1eH L . ;･ ,･.:. '.17."/"/'-..rfj t.f'' ' : ''/ . i/. 'LX ' .. ' .. x N .- . . ' " .' ./{.t - ' ,. ." - k, ". .r rL. "crt･' .t
i/be..e'
. x . "!

OshimmlsL
foshtma]s:.
NiiYms lsl.
MiyakeIsi,
.
Aogashimalsl.
t?tI
,e '

tr5
..
L
･ ･i.2i ?' L
FIG, 7. D{stributio nmap ofDs:yopteris caudipinna. Trjangles :previeusly known localitie Csi;rcles :newly recorded tocalities
from this study, Pie charts for Izu island sshow approximate ratio of sexual to apogamous individual s(whit e6:4 spores
per sporaiigium, blaek 3:2 spores per sporangiu m). Count sare based on Nakaike & Yamamoto (1984 f)o rOshima and on
Yamamoto & Nakaike (1983 f)or Haehlj ojima a,nd on our spore observatiolls for the remaining islands,
]038281]) .-Chiba Pref: Chosei-gun, Ichinomiya-ma- oue s,n. (TNS [VS-101749 1M]i)ya.kejima, Oyama, 1932-
chi, Gundari-yama, 2010-03-Og, K. }2imamoto &A, Ebi- 08-17, M .ibtan i5034 (TNS [VS-81570 O8y1am)a;, alt.
hara 100308Gl-8 ,14 ,15 ,I8 ,24, 25 & 27 (MAK). -Tb- 300 m, 1980-IO-1 0H,/ ]Hbtta & S. Mbtsumoto .Iivin cgol-
kyoPret:TbshimaIsl.1:933-07-30Z,.Jbtani8,202(TNS lectioonf7kukuba BotanicalGardenI3490(TNS[VS-

[VS-815690 ]1)9;33-07-31, Y .lotan is.n. (TNS [VS- 9508224, 9508226]); precise Ioca]it yunknown, l936-07-
121073] ;Y: Ybtan i8207 (TNS [VS-815683 ]M) ;Jbtani 11 ,ll ,Hayashi s,n. (TNS [VS-56244 ]1)9;61-04-0 2S,,
8210CTNS[VS-815979l9]5)7;-07-18l, Jbtani26210 Otaki3058(TNS[VS-103846189]6)4;-07-??,
cellector
(TNS [VS-815688 ]Ni)i,jima Isl. H,on-mura, 1981-05-04, unknown s,n. (TNS IVS-815713 ]19)6;4-07-10, collector
H Endo s.n. (TNS [VS-601629 ]1)98;1-05-?? ･I,l Endo unknown s.n. (TNS IVS-81607 781,6078]). Aogashima,
(TNS [VS-931539]K)o,zushima IsL,Tenjo-san, Ikenosawa,1958-07-20,X Jbtani27607 (TNS IVS-
s.n.
1936-07-24,ZJbtani63I2(TNS[VS-81569Z7]Jb)t;ani 815715,815716,815967,815968i)1;977-10-3M1,,Kato&

63B (TNS {VS-815698 ]pr)e;cise localit yunknown, E. A4iki li2 (TNS [VS-369811 T]o)d;aisyojirlj'y a1,988-
1936-07-26, M .Ibtan i6311 (TNS [VS-81569 5pr]e)c;ise 12-21 ,Z ShiJnakaw as.n, (TNS [VS-52220 -1Ka]n)a.ga-
Iocali tuynkllown, 1966-07-2 6to 1966-08-0 3H,. Ohba wa Pre£: [Fajisawa E-nsohshii]ma, 1979-09-1 8M, MI'ne-
30J8 (TNS [VS-173 12422e7,13]). Mikurajima Isl. ,Sato, mura s.n. (TNS [VS-46108 0-A]i)c.hi Pre£: Atsumi-
I934-07-26, Z Jbtani 108i6(TNS [VS-121e76 X] )jb;tani gun, Atsumi-cho, Yamada, 1980-11-23, K, biukai 4201
10818 (TNS [VS-815794 1]9)3;4-07-2 7M, .ibtani 10353 (TNS [VS-550286 ]-)1.[bttor iPre£] [Iwami-gu nFu,-
(TNS [VS-8IS79 O2y]a)ma;, 1934-07-2 6M, Jbtani 108i7 kube-mura ]Mani-san, l931-11-08, N. Hiratsuk as.n,
(TNS [VS-815973 ]pr)e;cise localit yunknown, 1967-07- (TNS [VS-512216 -]S)h.imane Pref. :Masuda-shi, Ta-
29, M .ibtan idi al'. 1279 (TNS [VS-815788 I19)6;7-07-3], kashima, l980-10-10, AC thake s.n. (TNS [VS-449584]).
Z A4titsuokali04(TNS[VS-8157871]9)6;7-08-02,K.bi- OkiIslands,Nishinoshima, Oki-gun,Nishinoshima-cho,

NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
November2010 YAMAMoTo & AL.-Distribution ofDryopteris caudZpinna l13
Mita, Takuhi-yama, 2008-07-Ol, l Kazumi s.n, (TNS Yadome-dake in Kagoshima Prefectur ean,d west
[VS-11053 95-Y]a)m,aguchi Pre£: Yamaguchi-shi, to Soheuksan-do, Korea. In contrast to the wide
N7i4h5o1, 9A2s]a)j1i; 9R9i3v-er0,8 1-91923H,-0M2k-2e0s, aHk,i Adi(nsTaNk isS.[n.V (ST-N7S 4[5V1S9-0]d)i,stributi oofn D, erlythresora, which occurs
s.n.
-Tbyoura-gun. [Tbyota-chToo,y]otanaka-mura, 1947- throughout the warm temperature region of Japan

08-23, K. Oka 1229 (TNS [VS-27610 2To]y)ou;ra-gun, (Kura t&a Nakaike 1985), D,:yopteri scauclipinna
Hohoku-cho, Awano, midstream ofFutanoi River, 1993- is distribu tone disland asnd in coastal areas un-
08-30 ,HL Adbsak is.n. (TNS [VS-745191 ]Ot)s;u-gun, der the influenc oef the warm Kuroshio and Tsu-
Y[uVySa--7c4ho5 ,1S7a7Zk4a5n1e7-8u]e ),1S.9h9i2k-o0k8-u2.2K, oHc.h MibPsrakeiE :s.Snu, k(umToN-S shima currents, although it alse occurs in moun-

shi, Okinoshima Isl. ,1971-07-23, T inobe s.n. (TNS [VS- tainous areas on Kyushu at it ssouthern limi tof
1038280]). Kyushu. Fukuoka Pref: [Tagawa-gund,istributi oTnh.e sexually reproducing D. caudi-
Kawara-machi,]Kawara-dake,1937-08-26A,dlfrie Castanopsisfbrests,
s.n. pinna usually occurs within
(1T00N3S 1[4V]S)--.N5a1g3as2a2k1 i1]9)5P1;r-e0f8-:2T6s, uHs, hOiumcahiIss,ln.. K,(aTmNiSa g[aVtSa-- and in the Kanto distri c(tnorth elrinmiO it is
in The
g2u1n,,S .K7almistustussIuh7ii4m7a9-(chToN,S I[zVuSmi-,5 a7lt3. 64003-5]K0a)m ;mi,a g1a9t7a9--0g7u-n, mdoistsltyri bu twi eoloflnD .p rceausdeirpivnenad ,pre chionwcet vgerro, vdeose, s not

Mine-mura, Kamizato, along Shishinouchi-ka waalt, 60 appear to be sirnply delimit beyd climatic condi-
m, 1979-07-19, S. kutsui 17322 (TNS [VS-573599 K]a)-; tions, because it has not been found on the Izu
rniagata-gun, Kamitsushima-cho, Eko, alt .20-30 m, and Kii peninsula sw,hich probably share similar
1979-07-20S,,tsutsuJi7432(TNS[VS-573602K]a)mi;a-
Ka mi agata- choM,i take , 80-90 1 979-07- meteorolegical cenditions with the localities
g1a9t,aS-,gtusnu, tsulirs4P(TNS [VS-57 a3lt6, 01K]a)m; im,a gata-gun, where i tdoes grow (fbrt Ihzeu Peninsula s,ee Na-

Kamiagata-cho M,itake ,alt .200-400 rn, l968-09-2 8S,. kaike et al. 1984) ,Dryqpteri scaudipinna nor-
foutsui s.n. (TNS [VS-204608 ]Sh)i;moagata-gun, Izau- mally fbrms mixed populations with the apoga-
h(aTrNaS-[ehVo S,A-r2i0a4k3e8-y6a]1m)9a7; ,8,.10926-82-6,09S-.27 3lIs,iu tZsKubi1y5aJm3a1 (2T8N1SP mous D. eilythrosora, although pure populatiQns
[VS- S73596]);T sushima-s h i,Mits ushi ma-cho,Kofa nseexduaaljo i,ndividua l(sD ,caudipinndi exception-
2007-10-09K,.Fleis2e92 (TNS[VS-768795]Ts)u;shima- ally occur on Hachijojima (Yamamot o& Na-

shi, Izuhara-ch oN,ainkawa, 2007-10-06 ,K. Fuse I82 kaike l983) ,Toshima, Niijima ,Kozushima, Mi-
(TNS [VS-768813]-)Ka,goshima Pref: Okuchi-shi, kurajirna, AogashimaIslandsT.he
and ecological
Manegahira ,1959-08-2 2T, famanaka s,n. (TNS [VS- and historic aflactor sthat determin ethe ratio of
734596]);Aira-gun,Kamo-cho, Yadome-dake, 1983-07-
19,MKbw abatas.n. (TNS[VS-7 42738,74274 0,742741]);sexual to apogamous forms are still unclear.
[Izumi-shJoin,o]dan-Shibi-Miyanojo1,957-08-15, S. The relationship between the two sexual dip-

Kurata 882 (TNS [VS-724546]). loi dtaxa, Dcyopteris caudipinna and D. koideu-
SOUTH KOREA. Chollanam-do iSoheuksan-d oIsl., miana, have not been carefully studied. The latter
1949-08-05 ,O, }longsok h8098 (TNS [VS-151180 -]). is usually treate das endemic to the Ryukyu Is-
Cheju-do:Cheju-do IsL,1962-07-28,S,-MOh 26rs(TNS
[VS-10 38337] ). lands, and is characterized by the abruptly nar-
rowed apex ofthe pinnae ,The distributi oofntshe
two species apparently do not overlap, but the dif
Discussion ferenc ein the morphology oftheir frond sis quite
subtle. At present w,e have littl beiologi ceavil-
We confirmed the presenc eof Dnyopteris dence for the two sexually reproducing diploids,
caudipinna in 13 prefectur eofsJapan and on two The dithcul tiyn distinguishi nD.g caudipinna
Korean island sT.he occurrence in seven prefec- from D, eslythrosora may arise from the fac tthat
tures in Japan and in Korea represent new range they share the same alleles. Future analyses of al-
extensions, Except fbr Shikinejim aw,hich lacked leli cdiversi twyith ample sampling should be
specimens with countable spores, we also con- performed in mixed population osf sexuallapoga-
firmed it spresence on all ofthe major Izu Islands. mous individua ransd the two sexual diploids.
Our investigatio nextended the distributio rnange
north to Iwama in Ibarak iPrefectur ee,ast to We thank Mr. Y. Kazumi for providin gvaluable speci-
Gundari-yama in Chiba Prefectur es,outh to mens.
NII-Electronic Library Service
TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
114 ActaPhytotax.Geobot. Vol.61

References tion and morphological characters of sexual and
apogamous individual sin the Kirishima mountains,
Fiijirno St.o ,199g .Observatio onn fiddlehea dasnd other K6y8u.s(hiu.nJ aJ,p Gaenneesreal) Educ,, lbkyo Univ. Agric. 20: 57-
characters of Dtlyopteri scaudipinna at its northern- Na kai, T.1 931.Notulae Japoniae& KoraeXL.
m2o4s.t( idiJnsatprainbeustei )Jo .nN.ippon Fern .Club 3(15-16) 2:2- Bot , Mag. (Tbkyo 4 )a5d: 9p1l-a1n3t7a.s
H i rasbpae yciaessh io, fHD., p,1o9p7t0e.r iCshr o(m4os )Jo..m Jeap. nBuombte.r s45: 1i1n -J1a9p.anese N a kaDkiiylkoye.q ,BpTut. el&rli . NAsea.t tlYyltah.rSmocasoimra.oMtuos ..g r,1o[9u[8pb4 k .i0yn boOsS,heesrrhv.iBamt1ai 0oi:n 5sosn1l -. Tt6,oh2-e,
H Iiw raatDbstaulykaiys,oKhp.it, e1r9H o9i.f5 sJ.ap1Da9rn7y.4o.p HteaCrryiatd oSagcheeooagbero.a ,IpThhb:ikIcyw oat.Sstuukdii,eKs .,oTn. Na ka(iiken, J Ta.p, aA .n eYasmea )moto & M . Maki. 19 84. A s tudy in
distribution,habitat,charactervariationofDrp,opter-
Ku riYJftaaeampr, aSanz.ns a(Ik1,I i9p,6Ip1B ),D , o1C..t2 h,E0rM.-o a1Bm7goo3,u,sf (ofKbo[mrde[da nbn&sukmh b7ayHe,4.ro s: OT3)hb9b k5oayf-os 4o.(m0ee1d. sJ F,alp)oar,na eosfe Ta k ai1ms9i 1yet-al2,yt0 h2rMo,, so( ra1i 9Jn9a6 p,( aIAnnsdeepsxie d)toi aChcre oaomfoe s)tohmee sIzu ofP eJnaipnasnuelsae.
Ku rata, S & T. Na kaike , 1985. Ill ustrati oonf Pterido- Pteridophyta (1910-199 J6a)p,an Pteridologica lSoci-
pkhyyot.es ofJapan Vbl. 4. University of Tokyo Press, Tb- Ya maetmyo, tTook, yAo., & T. Nakaike .1983 .A note on Dryopteris

M i tsfDnueirdssoaet),mr SiKc, at1n )9mK.9yueor, tiAoj niSemwhae kIruseblcuo. rt,Mdsau io zf2uD5rry(uo4-p5st)he: rii 1s(1n- 1eca3w u.dt(iopi i nKnnJiaankpia- Y a macMtaiumuodsoni.pt i,aon nnT,da oAk m.yo or &p,hN oSlaYoke.g yarOi .Bm o (9r :A io1sf, 6p 1s19io-m9de11i,7 aN 1 ao(.cstpieie daJsnaB caeeouopn)luas l.nt, heeNs adetfi)les. rtnSrs ciiibn.u-
M i yaDmio:tyoo,p tEer i& se ,T:y. tNhraoskoraamur gar. ou1p9 9i0n . SJyapsatne m2a. tDii scotsfr itbhue- tYhbek oMsiuurkaa CPietniy Mnussu.la 3, s9o :ut8h5-c-e9nt2r.al (i n JJapaapnaens ,eS)ci .Rep.
RecejvedAprii 2Z 20IO; accepted Adby Z 2010
NII-Electronic Library Service