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後期更新世から完新世の渦鞭毛藻シストのシ 原著論文 I ーゴニオラ ー
NOM 11 号(1 983 年 9 月 ) 1-32 原著論文 後期更新世から完新世の渦鞭毛藻シストのシ ノニムリスト I ーゴニオラ ックスグル ー プ List of Synonyms o fl a t e Pleistocene-t o -Holocene dinoflagellate c y s t s . 1Gonyaulax group. 松岡数充* Kazumi MATSUOKA \VI LSO l'i, 「 化 石渦鞭毛藻 i ま ,休眠期 の藻体 (シ スト z よ っ て提示された 。 これに続く Wall ( 1968 他) and Oale の現生シス卜の培養実験(.:1: , i s 1 α byr inth e α これに関係して上の例でもわかるように , のレ ベ ルでも同じ 状況 が存在する 。 E vi t t and Oavidson (1 964 ) に Ev itt (l 96 1 ) , Nem α tosphaerops ( OSTENFELO ) , T ec t α t odi nium pel litum WALL )。 cys t , 休眠 l生接合子= h y pn ozy got e ) が堆積物中 に取り込まれたものである」という見解は, t epikiense Ritecl α todinium ( RO SS IGNO L ) , はじめに Evitt 属 つまり iÕf 泳 体の Go ny α ul αx 属に 対 応するシストの Spini. (erites 属 , At α れ odinium 属 • Bit e ct α todin 山間 属, Tect α todinium属 Nematosph αeγ ops IS 属 , の見解を支持する 資料を提供する とともに両 である 。 鞭毛藻の分 類 についても 新たな問題を提起する れに対応する i方針永 体 を指摘 するのは困 難 であり , こ とに な っ た 3 この経緯については,すでに紹 さらにはシストの 研究者 介し た 1980 ) と i新制本の研究者 ( planktologist ) との 聞 に種の { 松岡 . が, その要点をここに述 1) 渦鞭毛主義とシストの形 態は, 互いに著し く巽ることが多 L ミ 。 そのために • ifj'i 泳 f本との対 応、がつかなか っ たシストに関する初期の研究で は, シストの形 態に基づく独自 の命名と分 類が ( f列 え{ま ROSSIG N OL , 1964 ; もN ALL , 1967 など )。 えて REID (l 974 ) は,シ又トの研究についてはシ ス卜独自の命 名分類を行う ( 命名規約は国際植 物命名規約に基 づく)の もやむを得ないとした 。 この観有、に立って , REID ( 1974 , 1977) , BRADFORO (1 975 , 1977) , HAR LA l'i D (1 977 ) は後期更 新世 から 完 新世 のシストを記 載・命 名 ・ 分類を行 っ た 。 2) このようにして得られたシストの分妻貢 f本 系ドでは競泳 体 とシスト の 種 が 必ず し も 一 対 ーの 対応 に な っ でおらず , 複数の シ ス卜慢が単 一 の続泳 体積に対応 することがある t永fキ: ( 主に pal y nologist ) 認識が異る ということである 。 以上 の点を 踏ま べておく 。 f子 わ れてきた 以上の事実は シ ス卜の形態のみからそ Gο ny α ul α x LA C HMA NN ) とシ ス卜 Spini(e γ it e s ( EHRENB ERG ) , S pin. e lon g α lus m e mb γ an α ceu s ( 例えば瀞 spini(er α ( CLAPAREDE ( ROS SIGNOL) and rαmosus REID , S pin. , Spin.miγα b il is 一方 ìõ'f?永{本の分類もこれまでは主に形態に基 づいてあ、り . 種や属の認識については複数の見 角干のあるも のもある ( f列え l ま G ony α ulax (er αco mpl ex とし寸表記 , sp in ト Py γ ophacus 属に おける s t e inii とむ a nc α mpo α e との関係 , Pγ oto. peridinium 属 における o c 印刷 cum と oblongum との 関係や Pγ otoce γα tlu m γ e ticul αtum ( CLA' PAREDE and LA C H~IANN ) カ、 Gony α ul α x gγ indl eyi * 長崎大学教養部地学教室 -1- N O~l ]1 号 REI NEC1 1~ (1 983 年 ) か)乙 と え!1 1主され て :.,..\ る Gi 叩旬 }' /.J C ド ta Þ・'f~. A rpolig e r α また表層 i'Ði- h史上t~ 械物中 に はまっ 16 0 のンス 卜型 197f ))í が 矢 11 られ -(' t., 、る ( DA l. r-" j馬 . {本 ぴHfl;ム、 :t ゎ r か 20 主主干重 コ\. T ,,' ) み 11)jら , ueridium i,fi,; 土 e pi c. yst al から!な-~ / )J f; ;l込 l こ のような f \;J状 から ,現世 の ( /I~ 鞭毛必 シス| m'(,を 111 ,、る グ7 仁.;. '; 仁を i与な L 、と 号 えっ る樋は, ノ1 占 i tî環 ¥_ ~: ! . .r~t 1/,n 二 子 11 (11, 1_ , またとし d禛J < ; : ¥1/;変 fじ j ( 1 ' . )i こ J担保 L て認 識 される ' え さ C、 あ るー 会 iS た j , a l' cheopyle 主~ ;},るじi造り :J II主主引がJ ~.... L とがつ ぐ Gony ωla~... 二 こは 論 る 点 n ;l l'注文であ り そ の 系統 を I.i n eage と する ,) /(~寸 し l li e ag t.~ ぷ pin l/e l' l/ e s 3 ¥ J4 ! . ー の同名任 L "),.'l か 7i Ac hÜ 11l u sph ue l α ,\'ρ 1T! O/ ο sphu C I υ/!S 1 . < も〉 R oll/'ne s/i α Hilecl CL! ocl 川 IU 1rl. l幻JI 一 ク の') ,上 う な 在観}見t.'バlり仏以 点 ゐ ~~ Ij) リ ノ :J ト % に L 限 会 iifi いて いる カ 純物 L l' [ 践 り ~H い 仁 J r; 〆 LL. ご , , .λ Lin gll l ndi η : !l m lá 近の五 If 究 ヤ, J 域 。 pe rc !l l υ dinlum ゾ ト が ん!i i幸夫耳東を 通 Tecl uludiniunl して 日立 Jd さ ~ l ' Cl. 心 こ 、りよう な らメ| 、 仁 柑 j J ,, \1α x iod i 'l1 ilU71 f.UJJ る二 と に した 1) メト í'iト裂 の [派 ' こ参 !~1~. し Im p α gi r1 inill l1l た 文献 の') t ) , iÜ: llld974 , HnADFo 院 υ i ( 197 7 ) , Homotryblio id L ,neage H 山 λ州 1977; , 1978ji : .uミ iよされとい',~!も Polysph α crldìll l7l T uh e I' c u ludι" のは除外 し ,そ Jれ: ~に:〈G 令'j: れた 文幻l献拡 :に子 つ いて ジのJ ノみ}. ぶ / jド ' 9 二 <'ι-白:に e Cel'atioi d ~\J () n についてのコメント ノ ヴノ Jt Hよ じりうf'f:fiH, ;y,、 ' ''',-.' ょ(,Ë ;f,、 グ〕 gonyaulacoid")し 7 i ニ it し ( Or er .J I ¥1 ) j '. ノ:;1 , J1I ィJド ノ J 、! 1t{ l ,) ) と T f, \Lo [( ( 1 98 0 ): 上... _..ンノ;; , t~ i _ _ _ , '~ .J t 川二 f可 ま れる手千 人,: J J,瓜 先Ii)j ノJ ~い な L 、 ゾつJ γ ,ノ Jル 1"' 一ソ〆/ :に みμノト i'Ar~ υJ け川7η1 .'1 α 川 (.J汁1J. μ 1ι uJ."<:て í'yrodinium , }'yruph u 仁 u s ( '' (,. ul iU11l し て L 、.;. ト 手あ ;,.:) ヵー し l'yrop;'ll C ll$ J,-.t'!, t!J ]'u!;er c u. !I, d けllU In aI'c h eop 、 l p 五 十 f , 、 1 :t .C },.J~~ :!.: rU l l . i が i i庁!ぶ j ,j) ノ/、 hypl)C 、 s ta! ,. { [t 1U mI~'f! , 1 > シ l' " Ii 刀、 J> r οfο cer οIυ{um η , 1 勺 υ I ο 百 υ nyilU ! α:r , Gυ ぷ n l J 土 1 ~Sμrド {キ: (/・ l I X:) iÆ:, A..lと .十 11 、 九 . ~ぺ.: i/) ケ l Conya ll Îa c 品 les ' 予 払l !I l 1 v)う > 恕i (平i JJ で . 辻 i n t ' age i nI T la rine こ の グ j レーノ( : . -1ナ i t l ld l .' "eagρ Tub"川、 1I {ο (i i 1 li un l 九 t コニオラ y クスグループ ( Gon :v aul a x group) こ こ! , f a う! f也の 的傾 (.:つ f投 υ 二小 /,) 然であ る 」こ と 有;: rt~. !i依 しご ル くてとが lj! 汲1:- ご あ 人、 ト;ユ {作 ノ 1乍:予~~ (δ ち ) ~l : Linf'age } gr り up ;J fv まれる ン スト泌を次~.: Conyaul acoi d ' H ¥C r j , ij { i~; L 、 でも れïl fX Jj Jr~ を紙: tHl!IJI~! の;/,',')革佐て~ ;~~ と いえ Jf L ーニ ジ) 上手 す ト にあ る わ け で l よ ない ♂ :~ ~え もし、 i よそれを jよ本に し た引) G()n) 'HuluC'oid .L' : 十 ( 枯十liO) 人才 d決 り や 与れに f 'jój I円殖機ょに くり変 化, 1 GOllya μlα ( Hl, や f勺" ofll c f' ratium Jr'.ft の ン スト:之 preci ngu l α 命れ規約にかヵ ー わる 問題ーが 残 るも ス ト 19 \ ' liJ )',j'j の 0> , ar('目 h eopy le をも つ こ と会とから .こ れ九 (! ))f~ i 土 宮 on:, Dllla co i d 7う 'i 二 な ってい つに j ー さ .~い, に関 し て :土, arche o py l e をも お , 主らに Pyr 什 d 川 ium J,of1, vり /ストである f" oly ぉ p h ー も IJ るかわら -r . ..rli 祭実験 い よ!~ -j く それらと巧干泳 シ ス卜 の づ苦労ー: Cani η gl α J~ : 正 apical J、 .一》 c 、出 t s ご p'{~ ri{. されど . 、る ン ストと ( - ノi 、 す TA゚LE Taxcnomi 仁 c o rrel a t io ns of tllccute anJ cy 只!.. 10 正 m 只 nf t l ¥ ( ! Urde1" C o nY ;J \ll .-]l 臼 1 0 :; Thec< : It e f orl!l n : Cy 川[ り l J l .AC ACEAE IGONYA¥ C ;O NY!, ULM:O I I IL INl lt ヒ!巳 re l1 c 八 CF I " . [ d (197 ん) . ] ) "1 0 ( 1'J l 川1 Gonyaulax spinifcri1 (Clapar de and LH~hman l1 ) c O Ol p l 回転 ご一一一一一一ー に pinifc rit es c l o Jl g c1 tu 只 H. cid 、忘忘ぶ一二ご 寸 l山川】 1げfc肝n日t 叩 川 111川 t日川川 山川川 il川 1111、 、ふふ\\ \ 、よとど 山川 tcs 川 d >i 1i 只川… 1) \ \ > \ \ S !)jrliferlt es r~n IUS1 I S (Ro s 引 gn ol) ¥¥¥ ¥ \\\、 Np lTl <l lospha p. r op s "j 京 lilbyríntllc ユ X… 日山 w Gonyaulax cf. spinifera (Clapar~ Je and L.:lchm c1 lln) ー一一 nl 1d 1 " ' 1 " (1 %8) 日、 110 ( J976 ) D,de (19i ら) 弘!n ( 川 8) ) o n (1968) ム 1 nNU Qノ 'b ) ( 1 J01 0 り 〆 は 'ど 111 ,, "d e 1 1 i ) d e l l w ‘ ( 札口 d 1)<, 1e 9 nd ] e l A < i - l ー -、 7 f ) 3 t t s ρし LK o t i t l r 』 c r 1i n n u ( f e t U 、, 3 1 r c じ U t E L ", 山 ( 1%8) し I ドじ,\( :1: 一一一一一一一一一一一 ilnly 叩 hiJ e r i. dj um 忽 山 山ツ i 一=ー 111he r c l! \山 1 ( K .ossignul) é\ 川 l 1I al 1J l Jnle ( J .968) A - o - ) l o h - GU RM [ o t } u r (HO!-ì 只 l 民 11 01) < t l dL . : l l :l J 1 l ¥n n) ruHERCL; LODI~I ()ID Pyrophacus stelni . i (Schi . llC l ") - L i L n Pyruphacus van 亡<l mpotle M バ 11 lいdω 川 h 川山 日州凡川州 ,1 ;-)1 川、 iハl (Defl .:.i Ildreθnd Cυo k son) i 只 l- nc l.i 礼!1 U l1l (t川 ! … M 山 iυ jい … 川一 川川 1刊l川 川 1l… 1此 川 Iω 山川 e ( < ' I i! CJ ト I O '[[{II I .lI. I(1 1.1J I 'YR OPHACACEAE e, (1 ')7 品) nl l m m:lch 川~rophoγ um (Defl andrc L 1n d Cookson) 1,:<l1 l 。 pe γ 口 l\ od i. n iu Jn Pyrodinium b. 1h amense PL1te- n ' o t t O l f > t (Clupulー を de Il ulloideU5 s e i e L .i nι l 1.Ì odi { Prutoceratium rcticulatum r Stcin 』 lyed γa [ 3 Gonyau1ax po i digltalis (' lo l l chp .t ) t - 5 じonyaulax (c r it e 日 11 ( ¥ < 3 1 1 ) 一一一一一一一日 pinif 己 r1 じ 旧日 ahr\Ilus - 令 一一一 一一一日 p jni Gonyaulax scrippsae Ko[oi (0 日 touf c1 d) 山 m t町叩叩山》は以州 ωk[ i比 山 L recrato<d ini υ m peJli: tum 、 L凶イ心,, (J リ 1 6) 1 ' 0 1 e \,:;111 and Dal t., Re. i \\人 /\laxlodinlu H1じ hoan U1 ll D i l l e (\<~1 1 ) \, ,,)1 anJ l .a1e (1 リ 70 ) I. INI:A じ l' in i:lH V i Jl l(川 poac (Rüssi 邑 nd) ¥ . : 0 1J . and ~la J), lle ( J .9J J . ) tS¥lo k ; 1 (1~} 82) :'¥OM 11 号(1 983 年 ) Note on c1assification system Many incubation experiments of 1iving dinof1age11ate cysts carried out by Wa11 and Da1e (1968 and other papers) proved that fossi1 dinof1age11ates inc1uding hystrichospheres in sensu strict are hypnozygotes (=cysts). On the other hand , their resu1ts brought us important and comp1ex taxonomic ・ prob1ems of the modern d 工 noflage1 1 a te c1assification. At the present , we ho1d two c1assification systems concerning the modern dinof1age11ares. of the vegetative and by phyco1ogists One of them is main1y based on the morpho1ogy mot 工 1e and ce11s , and this system is wide1y adopted p1ankto1og ユ S 仁 S 酬 depends on the cyst morph 0 10gy , Another system essentia11y and the uti1ity of this system is stressed by pa1y ロ ologists and micropa1eont010gists. But the 1atter system can not cover the who1e of the modern dinof1 age 11 a tes , hecause on1y a few parts of the modern dinof1age11ates produce the cy sts , which are preserved in surface sed iments , in the 1ife cyc1e. It is evident that a sing1e species shou1d keep a sing1e bio1oュ gica1 name. But actua11y we use two names for a sing1e dinof1age11ate spe c ies ,' for examp1e Gon ya u1M 己主語 ra St e 工 n , 1883 for the moti1e form and Lingu10dinium machaerophorum (Def1andre and Cüoksü n 、 1955) Wa11 , 1967 for the cyst form. Of co urse , according t the ru1e of the priority in the Internationa1 Code of the Botanica1 Nome n cle atur e , the varid name is Gonyau1ax po1yedra Stein , 1883 in this case. Rut in other cases , a sing1e thecate and moti1e species is correspond with severa1 cyst specie s Lachmann , 1859) , for examp1e Gonyau1ax spinifera (C1aparede and Di esing , 1866 for the thecate and m ti1e spiniferites e10ngatus Reid , 1974 , Spin. rnembr: 1naceus Sarjeant , 1970 , Spin. r nirabilis 王旦旦旦三旦s (Ehrenberg , choanum Reid , spheropsis 1964) ( Ro ss ign o 1 、 Sarjeant. 1970 、 a nd 19 64) S pini . 1838) Loeb1ich and Loeb1ich , 1 96 6 , Ataxiodinium 1974 , Bitectatodinium 1abyr 工 nthea pe11itum Wa1 1 , (Rossigno1 、 form 、 tepikiense \,: i1son 司 1973 . Nemato- (Ostenfe1d‘ 1903) Reid , 1974 and Tectatodinium 1967 for the cyst f rm. In the case of Gonyau1ax - 4- spinifera (C1aparede and Lachmann) , overc1assified in the cyst forms are probab1y the pa1eonto1ogic-based system (Da1e , 1978) , especia11y in the genus Spiniferites. On the contrary , as known in presentation of "Gonyau1ax spinifera comp1ex" (Dobe11 and Tay1or , 1981) , the thecate and moti1e forms are probab1y underc1assified. Therefore , at our present 1eve1 of know1edge of cyst-theca re1tions , we can not take a sing1e and idea1 c1assification system of the modern dinof1age11ates. Under these circumstances , stressed that in the interim , Da1e (1978) the cyst nomenc1eature shou1d ref1ect known pa1eonto1ogic and bio1ogic informations. Probab1y according to Da1e' . sconcept , Har1and (1982a) made a critica1 review of Recent and Quaternary dinof1age11ate cysts of the genus Protoperidinium. Har1and (1982a) started from the point that the taxonomic informations concerning the modern Protoperidinium are more comp1ete than those of cysts , and rearrenged and reattributed these cysts to q sing1e 1iving species , ProtoperidiniulT1. And he conc1uded that the different 1eve1 of the genus between thecate and cyst forms indicatξs the hiera1chica1 difference. On the other hand , Matsuoka (in press) does not direct1y accept the hiera1chia1 concept in the case of the heterosporous prob1em observed in such species as Gonyau1ax spinifera Lachmann) , (C1apar る de and and adopted the 1ineage system fundamenta11y proposed by Wa11 and Da1e (1968) and deve10ped by Wi11iams (1977). On the bases of this system , though the names of higher ranks -ーー Order and Fami1y in the thecate and moti1e form and Lineage in the cyst form --- are sti11 different between cyst and thecate-moti1e forms , the members consisting of these higher ranks are most1y common in both. This report dea1s with the P1eistocene and Ho1ocene dinof1age11ate cysts be10nging to the Order Gonyau1aca1es (Gonyau1ax group). References in the synonym 1ist pub1ished before 1978 are recorded in Bradford (1978) , Har1and (1977) and Reid (1974 , 1977). - 5 一 NOM 11 号(1 983 年 ) Genus Achomosphaera Evitt 1963 Type species: Achornosphaera rarnu1ifera (Deflandre 1937) Evitt 1963 Hystrich 心 sphaera rarnu1ifera Def1andre 1937 Achomosphaera anda10udsiense Chene 1977 1977 Achornosphaera anda1usiense Chene , p. 112 , p1. 1 , figs. 1-4. 1977 Spiniferites septentriona1is Har1and , p. 103-104 , p1. 1 ‘ figs. 12-18 , text-fig. 4. Genus Ataxiodiniurn Reid 1974 Type species: Ataxiodiniurn choanurn Reid 1974 Ataxiodiniurn choanurn Reid 1974 1966 Mernbranilarnax ? sp. Morzadec-Kerfourn , p. 141 , p1. 4 , figs. 1, 2 . 1974 Ataxiodiniurn choanurn Reid , p. 588-589 , p1. 1 , figs. 1 , 2 . 1976 P1aninosphaeridiurn rnernbranaceurn Eisenack 1965; Da1e , p1. 1 , f i g . 11. Genus Bitectatodiniurn Wi1son 1973 Synonyrn: Genus Ca1edonidiniurn Reid 1974 Type species: Bitectatodiτliurn tepikiense W 工 lson 1973 Ca1edonidiniurn verrnicu1aturn Reid 1974 1973 Bitectatodiniurn tepikiense Wi1son , p. 351-353 , fig. 2 , nos. 1-12. . 236 , 1974 Gen. et sp. nov.; Gonyau1ax sp. indet. ; Har1and , p p1. 1T , figs. 1 - 3 , 5 , 6 . . 589 - 590 , p1. 1 , 1974 Ca1edonidiniurn verrnicu1aturn Reid , p figs. 3-5. 1976 Bitectatodinium tepikiense Wi1son; Jux , p1. 1 , fig. 1 , pl. 2 , figs. 1-3 , pl. 3 , figs. 1-6. 1976 Bitectatodinium tepikiense Wi1son; Da1e , p1. 1 , fig. 12. 1977 Bitectatodinium tepikiense Wi1son; Har1and , p. 93-94 , pl. 1 , fig. 20 , pl. 4 , fig. 8 . 1977 Bitectatodinium tepikiense Wi1son; Reid and Har1and , p1. 1 , fig. 4 . Genus Impagidinium Stover and Evitt 1978 Type species: Impagidinium dispertitum (Cookson and Eisenack 1965) Stover and Evitt 1978 Leptodinium dispertitum Cookson and Eisenack 1965 Impagidinium acu1eatum (Wa11 1967) 1967 Leptodinium acu1eatum Wa11 , p. 104 , p1. 14 , figs. 18 , 19 , text-figs. 3c , d. 1979 Leptodinium acu1eatum Wa11; Har1and , p1. 1 , figs. 17-18. - ハヲ - QU s C上 ., ob • 今ム 1ム , ny 『J' QU s C上 -工 ob 1i 1上 p • , ュ - 1981 Impagidinium acu1eatum (Wa11) Stover and Evitt; Matsuoka , Impagidinium churchi11ii (Har1and 1968) 1968 Leptodinium churchi11ii Har1and , p. 548-551 , figs. 12 , l3, 22-24. 。 This species may be a corroded form of Spinifeγites bentori. Impagidinium mu1tip1exum (Wa11 and Da1e 1968) 1961 Anhystrichosphaera mu1tip1ex West , p. 452-453 (nom.nud.). 1968 Leptodinium mu1tip1exum Wa11 and Da1e , p. 318-319 , p1. 1 , nos. 1-7 , fig. 1 . Impagidinium paradoxum (Wa11 1967) Stover and Evitt 1978 1967 Leptodinium paradoxum Wa11 , p. 166-107 , p1. 15 , figs. 5-8 , -7- NOM 11 号 ( 1983 年 ) text-figs. 2 , 3A.B. 1978 Impagidinium paradoxum (Wa11) Stover and Evitt , p. 166. 1979 Leptodinium paradoxum Wa11; Har1and , p1. 1 , figs. 19-20. 1981 Impagidinium paradoxum (Wa11) Stover and Evitt; Matsuoka , p1. 1 , figs. 1-4 , p1. 2 , figs. 1-2. Impagidinium patu1um (Wa11 1967) Stover and Evitt 1978 1967 Leptodinium patu1um Wa11 , p. 105-106 , p1. 14 , fig. 20 , p1. 15 , figs. 1-4 , text-fig. 4. 1978 Impagidinium patu1um (Wa11) stover and Evitt , p. 166. 1979 Leptodinium patu1um Wa11; Har1and , p1. 1 , figs. 5-8. 1981 Impagidinium cf. patu1um (Wa11) stover and Evitt; Matsuoka , p1. 2 , fig. 10. Impagidinium sphaericum (Wa11 1967) 1967 Leptodinium sphaericum Wa11 , p. 108 , p1. 15 , figs. 11-15 , text-figs. 2a-c. 1976 Leptodinium shpaericum Wa11; Da1e , p1. 1 , fig. 10. Impagidinium stria1atum (Wa11 1967) Stover and Evitt 1978 1967 Leptodinium stria1atum Wa11 , p. 107-108 , p1. 15 , figs. 9-10 , text-fig. 5. 1978 Impagidinium stria1atum (Wa11) Stover and Evitt , p. 166. 1981 Impagidinium stria1atum (Wa11) stover and Evitt; Matsuoka , pl. 1 , figs. 5-6 , pl. 2 , figs. 3-4. Genus Lingu10dinium Wa11 1967 emend. Wa11 and Da1e 1973 Type species: Lingu10dinium machaerophorum (Def1andre and Cookson) Wa11 Hystrichosphaera machaerophorum Def1andre and Cookson 1955 Lingu10dinium machaerophorum (Def1andre and Cookson) Wa11 1967 - 8 1951 Resting spore of Gonyau1ax po1yedra Stein; Nord1i , p. 207-208 , figs. 1a-1f. 1954 cf. Gonyau1ax po1yerda Stein; Erdtman , fig. 2B. 1955 Hystrichosphaerdium machaerophorum Def1andre and Cookson , p. 274 , pl . 9 , figs. 4 , 8. 1959 Cyst of Goniau1ax , Mckee et a1. , p1. 1 , fig. 9 . 1961 Hystrichosphaeridium ashdodense Rossigno1 , p. 322 , p1. 1 , fig. 9 . nom. nud. 1962 Hystrichosphaeridium ashdodense Rossigno1; Rossigno1 , p. 132 , p1. 2 , fig. 2 , nom. nud. 1964 Ba1tisphaeridium machaerophorum (Def1andre and Cookson); Ro s signo1 , p. 90-91 , p1. 2 , fig. 14 , p1. 3 , figs. 20 , 21. 1964 Cyst of Gonyau1ax po1yedra Stein; Evitt and Davidson , p. 4-5 , p1. 1 , figs. 6-8 , 13. ?1966 Hystrichosphaeridium isoca1amus Def1andre and Ccokson; Morzadec-Kerfourn , p. 139 , p1. 2 , figs. 1 , 2 . ?1966 Hystrichosphaeridium redonensis Morzadec-Kerfourn , p. 139 , 140 , p1. 2 , fig. 3 . ?1966 Hystrichosphaeridium zoharyi Rossigno1; Morzadec-Kerfourn , p. 140 , p1. 2 , fig. 4 (not p1. 4 , fig. 3). 1966 Ba1tisphaeridiummachaerophorumDef1andre and Cookson; Morzadec-Ker fo urn , p. 140 , p1. 3 , figs. 1 , 2 . 1966 C1eistosphaeridium machaerophorum (Def1andre and Cookson) Davey et a t. , p. 170. 1967 Cyst of Gonyau1ax po1yedra Stein; Evitt , p. 43 , p1. 6 , figs. 13-19 , text-figs. 13-14. 1967 Lingu10dinium machaerophorum (Def1andre and Cookson) Wa11 , p. 109-110 , p1. 15 , figs. 16 , 17 , text-fig. 6 . 1967 Cyst of Gonyau1ax po1yedra Stein; Wa11 et a t. , fig. 1. 1967 Cyst of Gonyau1ax po1yedra Stein; Wa11 and Da1e , p1. 1 , fig. N. 1968 Lingu10dinium cf. machaerophorum(Def1andre and Cookson) Wa11; Har1and , p. 548 , figs. 8 , 8a , 27 , 28. - 9- N O M 11 号 (1 983 年 ) 1968 Resting spore of Gonyau1ax po1yedra Stein;Wa11 and Da1 e , p. 271-272 , p1 . 1 , fig. 18 , p1 . 3 , figs. 4-n . 1969 Ba1tisphaeridium machaerophorum (Def1andre and Cookson); Rossigno1 , figs. 24 (16) , p1. IV , fig. 1. 1969 Lingu10dinium machaerophorum (Def1andre and Cookson) Wa11; Downie and Singh , figs. 7 , 8 , 1 0 , 11 . 1971 Ba1tisphaeridium sp. Takahashi , p. 20- 21 , p1. 3 , figs. 2 , 3 , 5. 1971 Cyst of Gonyau1ax po1yedra Stein; Wa11 , p1. 2 , figs. 7 , 8 . 1971 Lingu10diniummachaerophorum(Def1andre and Cookson) Wa11; Jux , p. 168 , 1 7 1-172 , p1. 28 , figs. 23-29. 1973 Lingu10dinium machaerophorum (Cookson and Eisenack); Wa1 1 , Da1e and Harada p. 24 , p1. 2 , figs. 5 , 6 , p1. 3 , f 工 gs . 11 , 12. 1974 Lingu10diniummachaerophorum(Deflandre and Cokson) Wa11; Wa11 and Da 1e , fig. 1P. 1974 Lingu10dinium machaerophorum(Def1andre and Cookson) Wa11; Ried , p. 591-592 , p1. 1 , figs. 6 , 7. 1977 Lingu10dinium machaerophorum(Deflandre and Cookson) va11 ; Har1and , p . 94-96 , p1. 2 , figs. 22 - 24 , p1. 4 , figs. 11 , 17 、 text-fig. 2. 1977 Lingu10dinium machaerophorum (Def1andre and Cookson) Wa11; Reid and Har1and , p1. 1 , fig. 2. 1981 Cyst form of Gonyau1ax po1yedra Stein; Kobayashi , Matsuoka and Iizuka , p. 53 - 55 , p1. 1 , figs. A , C , D. Genus Nematosphaeropsis Def1andre and Cookson 1955 emend. Wi11iams and Downie 1966. Type species: Nematosphaeropsis 1abyrinthea (Ostenfe1d 1903 ) Reid 1974 Ne matosphaeropsis ba1combiana Def1a nd 了 e and Cookson 1955. Nematosphaeropsis 1abyrinthea (Ostenfe1d) Reid 1974 10 ー Da1e , p. 270-271 , p1. 1 , figs. 14 , 15. ?1970 Spiniferites ramosus (Ehrenberg); Wa11 and Da1e , p. 49-50 , p1. 1 , figs. 1-15. ?1971 Cyst of Gonyau1ax sp. aff. G. spinifera (C1apar鐡e and Lachemann); Wa11 , p1. 2 , fig. 4. 1970 Spiniferites bu110ideus (Def1andre and Cookson) Sarjeant , p. 75. 1981 Spiniferites cf. bu110ideus (Ehrenberg); Ma tsuoka , p1. 2 , fig. 7 . Spiniferites ramosus (Ehrenberg) Mante11 1854 ?1959 cf. Hystrichosphaera furcata (Ehrenberg); McKee et a1. , p1 . 1 , fig. 7 . ?1961 Hystrichosphaera furcata (Ehrenberg); Rossigno1 , p1. 1 , fig. 6 . ?1962 Hystrichosphaera furcata (Ehrenberg);Rossigno1 , p. 132. ?1963 Hystrichosphaera cf. furcata (Ehrenberg); Rossigno1 , p1. 1 , figs. 11 , 12. 1964 Hystrichosphaera furcata (Ehrenberg); Rossigno1 , p. 85-86 , p1 . 1 , fig. 11 , p1 . 3 , fig. 9 . 1966 Hystrichosphaera bu110idea (Def1andre and Cookson); Morzadec-Kerfourn , p. 138 , p1. 1 , fig. 4. 1967 Hystrichosphaera furcata (Ehrenberg); Wa11 , p. 99-100 , p1. 14 , figs. 1 , 2 , text-fig. 2 . 1968 Hystrichosphaera ramosa var. ramosa (Ehrenberg); Har1and , p. 540-541 , fig. 21 . 1968 Hystrichosphaera bu110idea Def1andre and Cookson; Har1and , p. 544 , fig. 20. 1969 Hystrichosphaera ramosa (Ehrenberg); Har1and and Downie , p1 . 7 , figs. 3 , 7 . 1971 Spiniferites ramosus (Ehrenberg); Takahashi , p. 16-17 , p1 . 2 , figs. 1-3. 1974 Spinifarites cf. ramosus (Ehrenberg); Har1and , p. 231233 , pl. 1 , figs. 9 , 10 , pl. IV , fig. 6 . 2 1 NOM 11 号(1 983 年 ) 1974 Spiniferites bu110ideus (Def1andre and Cookson); Reid , P. 600-601 , p1. 2 , figs. 1-3. ?1975 Spiniferites ramosus ramosus (Ehrenberg); Davey and Rogers , p1. 1 , fig. 5 . 1976 Spiniferites bu110ideus ( Def1andre and Cookson); Da1e , pl. 1 , fig. 4. 1977 Spiniferites ramosus (Ehrenberg); Har1and , p. 102-10 3 , p1. 1 , figs. 5 , 6 . 1977 Spiniferites ramosus (Ehrenberg); Reid and Har1and , p1. 1 , fig. 5. Spiniferites hyperacanthus (Def1andre and Cookson) Reid 1974 1955 Hystrichosphaera hyperacantha Def1andre and Coo kson , p. 264 , pl. 6 , fig. 7 . 1967 Hystrichosphaera hyperacantha Def1andre and Cookson; Wa 11 , p. 100 , p1. 14 , fig. 3. 1974 Spiniferites hyperacanthus (Def1andre and Cookson) Reid , p. 603-604 , pl. 4 , fig. 35. Spiniferites pachydermus (Rossigno1) Reid 1974 1964 Hystrichosphaera furcata var. pachyderma Rossigno1 , p . 86 , pl. 1 , figs. 1 , 2 , pl. 3 , fig. 6 . 1968 Hystrichosphaera ramosa var . reticu1ata Davey and Wi11 工 ams Har1an d , p. 541. 1969 Hystrichosphaera furcata var. pachyderma Rossignol; Rossig no1 , p. 98 , text-fig. 24 , fig. 5. 1969 Hystrichosph a ♂ r a ramosa var. pachyderma sensu Har1and and Downie , p. 232 , p1. 7 , fig.9. 1974 Spiniferites pachyde r r nus (Rossigno1) Reid , p . 607 - 608 , p1. 4 , figs. 36-38. Spiniferites frigidus Har1and and Reid 1980 . 2 , fig. 5. ?1977 Cyst of Gonyau1ax sp. A Reid and Har1and , p1 , 内ノ』 内ノ】 1980 Spiniferites frigidus Har1and and Reid in Har1and et a1. , p. 213-214 , Figs. 2A-J , 3 . 1982 Spiniferites frigidus Har1and and Reid; Har1and , p1.2 , figs. 5-16. Spiniferites e10ngatus Reid 1974 1968 Resting spore of Gonyau1ax sp. 1 Wa11 and Da1e , p. 271 , pl . 1 , fig. 16. 1969 cf. Hystrichosphaera sp. a Har1and and Downie , p. 232 , pl . 7 , fig. 4 . 1974 Spiniferites e10ngatus Reid , p. 602-603 , p1. 3 , figs. 23 , 24. 1976 Spiniferites e10ngatus Reid; Da1e , p1. 1 , fig. 8 . 1982 Spiniferites e10ngatus Reid; Har1and , p1. 1 , figs. 9-10 , text-fig. 2 . 1982 Spiniferites e10ngatus Reid; Ma tsuoka , p1. 2 , figs. 1-2. Spiniferites nanus Matsuoka 1976 1976 Spiniferites nanus Matsuoka , p. 111 , p1. 28 , figs. 1-3. 。Th is species is a synonym of S . bu11oideus. Spiniferites curciformis Wa11 and Da1e 1973 1973 Spiniferites cursiformis Wa11 and Da1e in Wa11 , Da1e and Harada , p. 21-22 , p1. 1 , figs. 1-6; p1. 2 , figs. 1-4. Spiniferites inaequa1is Wa11 and Da1e 1973 1973 Spiniferites inaequa1is Wa11 and Da1e , in Wa11 , Da1e and Harada , p. 22 , p1. 1 , figs. 7 , 8 . Spiniferites pseudofurcatus (K1umpp 1953) subsp. ob1iquus - 23- N O M 11 号(1 983 年 ) (Wa11 1967) Lentin and Wi11iams 1973. 1967 Hystrichosphaera tertiaria var. ob1iqua Wa11 , p . 103 , p1 . 14 , fig ・ 16 , text-fig. 2. 1973 Spiniferites pseudofurcatus subsp. ob1iquus (Wa11 1967) Lentin and Wi11iams , p. 129. Spiniferites rubinus (Rossigno1 1964) Sarjeant 1970 1962 Hystrichosphaeridium rubinum Rossigno1 , p. 134 , nom. nud. 1964 Hystrichosphaera rubina Rossigno1 , p. 87 - 88 , p1. 1 , figs. 12-13. 1970 Spiniferites rubinus (Rossigno1) Sarjeant , p . 76. 1979 Spiniferites rubinus (Rossigno1); Har1and , p. 537 , p1. 2 , figs. 4-11 . Spiniferites scabratus (Wa11 1967) Sarjeant 1970 1967 Hystrichosphaera scabrata Wa11 , p.102 , p1. 4 , figs. 10-14 , text-fig. 2. 1970 Spiniferites scabratus (Wa11) Sarjeant , p. 76. . 1976 Spiniferites scabratus (Wa11); Da1e , p1. 1 , fig. 6 Hystrichosphaera nodosa Wa11 1967 1967 Hystrichosphaera nodosa Wa11 , p. 101-102 , p1. 14 , figs. 7-9 , text-fig. 2 . 。 This species is a synomym of Spiniferites bentori. Spiniferites ramu1iferus (Def1andre 1937) Reid 1974 1935 Hystrichosphaera cf. ramosa (Ehrenberg 1838) Def1andre , p1. 5 , fig. 11. 1937 Hystrichosphaeridium ramu1iferum Daf1endre , p. 74 , p1. 14 , figs. 5-6 , p1. 17 , fig. 10. 1963 Ba1tisphaeridium ramu1iferum (Def1andre) Downie and Sarjeant , p . 92. -2 4 1963 Achomosphaera ramu1ifera -(Def1andre) Evitt , p. 163 , Fig. 2 . 1974 Spiniferites ramuriferus (Def1andre) Reid , p. 608 , p1 . 4 , figs. 39 , 40. 1981 Spiniferites sp. 1 --- Gonyau1ax sp. 1 Dobe11 and Tay1or , p. 100 , p1 . 1 , fig. 4. 。 The thecate form of this species resemb1es Gonyau1ax a1askensis Kofoid 1911 , and the cyst has an opercu1um in two pieces (2P). 1981 Spiniferites sp. 2 --- Gonyau1ax sp. 2 Dobe11 and Tay1or , p. 102 , p 1. 1 , figs. 9-13 , 16 , 19 , 20. 。 The thecate form of this species may be conspecific with G . sp. 1 Dobe11 and Tay1or , and this cyst has an opercu1um in two pieces. 1981 Other 2P cysts Dobe11 and Tay1or , p. 102 , p1. 1 , figs. 14 , 15 , 17 , 18. Genus Tectatodinium Wa11 1967 Type species: Tectatodinium pe11itum Wa11 1967 Tectatodinium pe11itum Wa11 1967 1963 Leiosphaeridium scrobicu1ata (Def1andre and Cookson 1955); Rossigno1 , p1. 2 , fig. 22 , nom. nud. 1964 Leiosphaeridia scrobicu1ata (Def1andre and Cookson); Rossigno1 , p. 92 , p1. 1 , fig. 19. 1967 Tectatodinium pe11itum Wa11 , p. 113 , p1. 16 , figs. 11 , 12. 1968 Resting spore of Gonyau1ax spinifera (C1aparede and Lachmmann) ;Wa11 and Da1e , p. 269 , p1. 1 , fig. 12. iu ph qL NOM 1]号(1 983 年 ) 1968 Tectatodinium pe11itum Wa11; Wa11 and Da1e , p1. 1 , no. 8. 1977 Tectatodinium pe11itumWa11; Reid and Har1and , p1. 1 , fig. 3 . 1979 Tectatodinium pe11itum Wa11; Har1and , p1. 1 , fig. 9 , 10. Tectatodinium psi1atum Wa11 and Da1e 1973 1973 Tectatodinium psi1atum Wa11 and Da1e in Wa11 , Da1e and Harada , p. 27 - 33 , p1. 2 , figs. 9-15; p1. 3 , figs. 1-6. Tubercu1odinioid Lineage Genus Tubercu10dinium Wa11 1967 emend. Wa11 and Da1e 1971 Type species: Tubercu10dinium vancampoae (Rossigno1 1962) Wa11 1967 Pteropspermopsis ? van campoae Rossigno1 1962 Tubercu10dinium vancampoae (Rossigno1 1962) Wa11 1967 1962 Pterospermopsis ? van campoae Rossigno1 , p. 134 , p1. 2 , f i g . 1. 1964 Pterospermopsis ? vancampoae Rossigno1; Rossigno1 , p. 90 , p1 . 2 , figs. 17 , 18 , p1 . 3 , fig. 15. 1967 Tubercu10dinium vancampoae (Rossigno1 1962) Wa11 , p. 114 , p1. 16 , figs. 15 , 16. 1971 Cyst of Pyrophacus vancampoae (Rossigno1) Wa11 and Da1e , p. 234 - 235 , Figs. 10-25. 1976 Tubercu10dinium vancampoae (Rossigno1) Wa11; Matsuðka , p . 114 , p1 . 29 , figs. 6 , 7. 1982 Tubercu10dinium vancampoae (Rossigno1) Wa11; Matsuoka. p1 . 1 , figs. 10 , 11 . 1979 Tubercu10dinium sp. Har1and , p1. 3 , fig. 8. -2 6- GONYAULACALES CYST without cyst ・name Genus Protogonyau1ax Tay10r 1979 1980 Cyst of Protogonyau1ax catene11a (Whedon and Kofoid) Tay10r 1979; Fukuyo , p. 51. 1981 Hypnozygote of Protogonyau1ax catene11a (Whedon and Kofoid) Tay1or; Yoshimatsu , p. 136 , p1. 11 , figs. 9-12. 1945 Cyst of Goniau1ax Tamarensis Lebour var. excavata . 10-12 , p1. 11 , Figs. n , o. Braarud 1945 , p 1978 Cyst of Gonyau1ax tamarensis Lebour; Turpin , Dobe11 and Tay1or , p. 236-237 , Figs. 2-6 , 9 . 1980 Cyst of Protogonyau1ax tamarensis (Lebour) Tay10r 1979; Fukuyo , p. 51 , Fig. 1 (8-2). 1977 Cyst of Gonyau1ax excavata (Braarud 1945) Ba1ech 1971; Da1e , p. 30 , Figs. A-C. 1981 Cyst of Protogonyau1ax sp. (H1R) Fukuyo , p. ーーーーーーーーーーーー 1982 Cyst of Protogonyau1ax sp. Matsuoka , p. 200 , p1. 2 , fig. 12. 1979 Cyst of Gonyau1ax moni1ata Howe11; Wa1ker and Steidinger , p. 313-314 , Fig. 21. Genus Fyrodinium P1ate 1906 1978 Cyst of Pyrodinium bahamense var. compressa (Bδhm) Steidinger , Tester and Tay1or , p . 331. Genus He1go1andinium Stosch 1969 1969 Cyst of He1go1andinium subglobosum Stosch , p. 576-577 - 27- , N O M 11 号(1 98 3 年 ) figs. 1-6. Genus Pyrophacus Stein 1883 1982 Cyst of Pyrophacus steinii (Schi11er) Wa11 and Da1e; Matsuoka , p. 199 , p1. 1 , figs. 10 , 11. 1971 Resting spore of Pyrophacus horo1ogiurn Stein 1883; Wa11 and Da1e , p. 230-231 , Figs. 34-36. QU ? “ References Bradford , M.R. (1975) New dinof1age11ate cyst genera from the Recent sediments of the Persian Gu1f. Can. Jour. Bot. 53. 3064-3074. ー一一ーーーー ー.-. (1977) New species attributab1e to the dinof1age11ate . Leieunii ! . .Ger1ach ,工 961 emend. Lentin and Wi11iams , 1975. cyst genusGrana , 16 , 45-59. ーーーーーー一一, ー.ー. (1978) An annoted bib1iographic and geographic review of P1eistocene and Quaternary dinof1age11ate cysts and acritarchs. Amer. Asso. Strati. Pa1yno1. , Contr. 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Sci. , ';01, iii+300pp. Wa11 , D. (1967) Fossi1 microp1ankton in deep-sea cores from the Caribbean Sea. Pa1aeonto1ogy , 10(1) ーーーー, ー. , 95-123. and Da1e , B. (1968) Modern dinof1age11ate cysts and evo1utior . of the Peridinia1es. Micropa1eonto1ogy , 14(3) ーーーー, ー. and -一一ー, ー. , 265-304. (1971) A reconsideration of 1iving and fossi1 Pyrophacus Stein , 1883 (Dinophyceae). Jour. Phyco1ogy , 7 , 221-235. - 31- NOM 11 号(1 983 年 ) Listofsynon ~' ms ofl a t e Pleistocene-to-Holocenedinoflagellate cysts Gonyaulax Group Ka z umi 九1ATSUOKA Dep a r t m en tofGeology , Fa cu lt yofL i b e r alArts, Nagas a k iL n ive r si tト Bunk yo- じ h o ‘ '\;agasaki 852 、 lapan ABSTRACT e i st oceneandHol o cened i n o t l a g e l l a t ec y st swered e s c r i b e d S i n c el as ttwen t yyears , manyPl andi l l l l s t r a t e d. M05t0' 1t h e s ec ys t swer ecl as s f ie c li nt hep a l e o n t o l o g i c b a s e ds y stemwhich i s i n d e p e n d e n t l ydev e loped from t he b i o logi cb as e ds y st em. Under t he s e circllmstances , we In ders t a n d i ngoft h er el a t i o ns h i pbetweent h et he c a t eandcys tforms s hol l l dhavet h es t a n dofl Ourp r e s e n tknowledgcoft h ecl a s s f ic at i ons y st em betweent hev e g e t a t i v ec e l landc y s ti 5 revicw ed 、 a nd t h et e n tat i vcc y s t c l as si f cai tons ystemi sp r o p o s e d . Ther epor tal 50includ 巴 s t hc i l s tofsy nony 口l S ofl at eP lei s t o c e n ea n c lHo l o c ened i n ot l a ge lat ec ys t sa s si g n a b l et ot heOrder Gonyal l l a c a le s(GonyaulaxGroup) 引用文献リスト中の注 ( 牢 ) 本 1 福代康夫(1 980 ) ;:~底 j尼 iゃからの Go ny α ul α x Stein のンストの発見, 有毒 プラ /クト ン : 赤潮 に 関 する近年の知見と研究の問題点ー赤 潮研究会・編集委員会編, p p .46- 5 3 . t : 2i 凶i 数充(1 983 ) *4ti~ r出!主主 7だ 渦鞭毛藻シス 2, ( 1980 ) , 1,ゐ 革便 モ ;災ンス卜 そのう〉 vol . no.4 , pp.319. .326 牢 5 松間数先 ( 1982 ) 大村湾表!長 堆 F貨物 '.1' の;f,',\ 鞭毛藻 シスト ・ 海洋環境 特 性と赤潮発生 p p .389-40 3 ,↑訂正止社厚生 :二 関 する 基礎研究,環境科 学 研究報告書, 時~j 本 3 小林 vo1 .28 , 類学 1: の 意義 について-- Jj刊地球, ト ・ 沿トj'!J!!X 保全のための海の五環 境科学. 平野敏行編 , Flプ幸1;1, n o .1 , pp.53- 5 7 . け本水産資源保設協会 . 本 2 福代康 夫 . polyedr α B148-R 14-8 ‘ 聡 ・ 松岡数先-飯塚 H~L~ (1 981 ) 大村 - 32 一 pp. 197- 204.