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Biosystematic Studies on the Genus Ixeris − （Compositae Lactuceae）1． Fruit Wali Anatomy and Its Taxonomic Implications

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Biosystematic Studies on the Genus Ixeris − （Compositae Lactuceae）1． Fruit Wali Anatomy and Its Taxonomic Implications The Japanese SocietySooiety for Plant SystematicsSystematios − − ISSN OOO1 6799 Acta Phytotax． Geobot．41 ： 43 60 （1990） Biosystematic Studies on the Genus Ixeris − （Compositae Lactuceae）1． Fruit WalI Anatomy and Its Taxonomic Implications − JaeHong PAKI and Shoichi KAwANo2 ・ 一 朴 宰 弘 河 野 昭 ： ニ ガ ナ属 （キ ク科 ・タ ン ポ ポ 連 ） の 分 類 学 的 研 究 1． 果皮 の 解 剖 学 的 特 徴 と そ の 分 類 学 的 意 義 Abstract The mature fruit wall structures were invcstigated fer l80f about 20 傭 癇 spec 三es − （Compos 童tae Lactucea ¢ 〉． Fourteen species had 10 costae ， nearly the same size in transverse sections and fbur species had − lO− − ， 5 70r 13 costae ， Fruit waU structures were divided intotwo ． maimypes ： wi nged type subdiv 蓋ded into Type I and Type ILA B G and ribbed type Type （ ， ， ）， （ III ．Type I has thick 量ntercosta L repens Type II−A B and G have weU −developed costae ） （ ’）； ， 孟 ‘ePhala ， 1． debilis五 5 ‘oJo η γα L longirestrata 孟 ‘海 η 8 π 廊 and L tamagatvaensi ． The ribbed （ Poly ， 静 ， ， の type （Type III）has the least conspicuous costae （1． dentata and 工Oother species of sect ． Ixeridium）． In the winged type ， costae are f（）rmed by cell divisiens in the i nitial stage of costa1 tissue develop− ment was ， but its development less conspicuous in the intercostal tissuc of ∬． rep 砌 50f Type L In the ribbed type cQstae are of rather ， fbrmed by the disintcgration intercostaltissue ， than by − − cell divisions in costal tissue throughout pre and post fとrtil 重zation stages ． Based on the different fruit−wall types and ontogeneticprocesses as well ， as on karyological data， we suggest that l l species ng sect w 量 r 量 belong ｛ to the ．Ixeridi ， th bbed achenes ， and largechromosomes with a basic number of κ 富 7， can be distinguished from the rema ｛ning spec 互es ， which have wingcd achencs and s 皿 all chromosomes and a bas量cnumber of κ 磊8。 New evidence conceming 丘u 董twall structures obtained ’ in the study does not supPort earlier scctional dassifications except fbr sect ． ChoTisls to present ， ， which a single species L γ 8π∫ 三s referred ． resuhs suggest species referred sect ， ψ ， Our that the to ， Ixeriditrmwith ribbed ， achcnes alld large chromosomes with a basic number of x ＝7 represent ， adistinct and thus should phylogenetic group ， ， be placed in a separatc genus f｝om 勧 ガ∬ ．str ． Key words ： Gompositae−Lactuceae IxeriS Ixeridium wall anatomy ， ， ，fruit ． and species The generic delimitation relat 量onships of 、rxeris and allied genera Compositae ； tribe Lactuceae ， all of which are fbund f｝om north to south −eastern Asia （ ） ， have been controvers 三al f‘）r many ． The Ixeris years genus ， as originally describedin 1826by CAssINI GAsslNI l 8263 KING and DAwsoN l on （ 975）based Ixerispoipcephala， c 圭rcumscribed only several species （DE CANDoLLE 1838）． The gcnus has also been lDepartment of Botany ， Faculty of science Kyoto university Kyoto 606 ． ， ， ， Japan Present address ： Departmeat of Biology Gollcge of Natural Science Kyung −Pook National ， ， Unlversity， − Taegu 702 701， Korea． 2Depart 皿 t of Botany Faculty of Sc 量ence Kyoto Univcrsity Kyoto ， ， ， 606，Japan， and Instituteof Genetic E 〔ology Tohoku Univers 三ty Sendai 980 ． ， ， ， Japan 一 NII-ElectronicN 工 工 Eleotronio Library Service The JapaneseSocietyJapanese Society for Plant Systematics " Acta Phytotax. Geobot. Vol. XLI, Nos. 1-3 included in the broadly defined Lactuca (BENTHAM 1872). In more recent classifications, however, freris has been conEidered to consist either of approximately 20 species (NAKAi 1920; KiTAMuRA 1956, 1981; see also GRAy 1859) or ofabout 50 species, including some species of &opidiastrum, Paraixeris (NAKAi 1920) and even Lactuca (SfrEBBiNs 1937; ToMB 1977). In these classificatory systems, achene morphology, the shapes ofthe involucre and coTolla, and the numbers ofvascular bundles in the ovary wall (STEBBiNs l937) have been used as key characters for defining these genera. According to KiTAMuRA (1956, l981), the species of ixen's are characterized by fusiform achenes with a mQre or less elongated beak, ten ribs or wings, and also a persistent bristled pappus. A related genus, ewopidiastram, also possesses fusiform achenes, but lacks a conspicuous beak. KiTAMuRA (1956) suggested, based on the Iife form, growth habits, leafmorphology, euter involucre size, head number, and pappus color, that llverds is divided into five sections: Chorisc's, Rsende-chon'sis, indb-ixeris, Sobotixeris and .1beeridezam. In these earlier studies, however, little attention was paid to the anatomical features efachenes. The only exception is a recent contribution by BELiAEvA and BoyKo (1980), in which achene morphology and anatomy of 13 species ofLactasa from the eastern region of the Soviet Union, including three 1]teris species (i.e., 1}veris ropens (=C`thorisds ropens"), L chinensis (--"freridium gramineum"), and L dentata (=:"1iveridium dentatum"), were critically investigated. They suggested that the achene wall structures are important and usefu1 fbr the delimitation of genera and for assessing species relationships. Thus, a more extensive study on fruit wall anatomy covering all the species in 11ven's and related genera became necessary in order to elucidate the exact status of these taxa. The purpose ofthe present study is twofbld: first, to describe the fruit wal1 structures of 18 llven's species, representing all five sections (KiTAMuRA 1956), that occur in Japan, Korea, Taiwan, Malaysia and New Guinea; second, to compare and discuss aMnities of the species based on these results and other gross morphological as well as karyological characters. For further understanding of the character states in fruit wall anatomy, the ontogeny of the fruit wall was also critically examined in some representative species. Material and methods The eighteen ofspecies of1beeris (KiTAMuRA 1956, 1981) are listed in Table 1. Most of the fruits were obtained from herbarium specimens in the Herbarium of Kyoto University (KYO)s and some were collected from wild habitats or cultivated plants. Fruits were always soaked in or' fixed with FAA (formalin : glacial acetic acid : 50% ethanol=1 : 1 : l8), and then dehydrated through a t-buty1 alcohol series and'embedded in Paraplast with MP 57-58 C for microtoming. Transverse and longitudinal sections cut at 8-12 psm were stained with safranin and fast green FCF, and mounted with Entellan. NII-Electronic Library Service The JapaneseSocietyJapanese Society for Plant Systematics Septernber, 1990 Acta Phytotax. Geobot. 45 Table 1. Species of ixeris examined and cellection sitesi). Sectionfspecies2} Voucher specimens Sect. C7lorisis (DC.) A. GRAy I. ropens (L.)A. GRAy Japan.Chiba: Tateyama,Hiroe17221. Hyogot Isl. Awaji, Kbnde 2J99. Hoklcaido: Kitami,IVbrtthashil13e. Sect, lhab-ixeris KiTAM･ I. Palycophala CAss. Japan. Osaka: Takai,S;,to 3735. Kanagawa: Kawasaki,vazteshima16417. Sect, Psetedb-cherisis KrTAM･ J. debilisA, GRAy Japan, Aichi: Shinoiima, 1lakahashi2145. Mie: Mt. Suzuka, 24 May 1938, Okamoto s.n.. Korea, Chollanam-do: Dalri-do, Pak 780. L stolontfara A. GRAy Japan. Kyoto: Ooeyama, Mltrata 45066. Nagano; Mt, Shirouma, furuse 30. Ishikawa: Tatsunoguchi,Sugde 1022, Sect. Sbbelixeris NAI AI L longirostrata NAicAi (HAyATA) Japan. Isls.Bonin: Hahajima, Kbbayashi et al. 8gt1･ Sect. 1hreridium A. GRAy I. tamagatvaensis KiTAMJ '8178. CMAKiNo) Japan. Nagano: Kami-ina, Mlatrata Matsumoto, Pak 920, Shizueka: Ohkawachi 17 Oct. 193!, Klatrosawa s.n., L chinensis (THuNB.) NAKAi Korea. Taegu: Mt, Palgong, Pde 679-2, Republicef ahina (Taiwan). Hua]ien, Wenshan, S)Vimilu 12639. L dentata (THuNB.) NAKAr Japan. Nara: Yoshino, 7:sugaruet al. ee98. Nagano: Shiga-kogen,Pak900. L YAHARA Parva(KITAM.) Japan. Kagoshima: Isl. Yakushima, 1toatstLkietaL 59. L vakuinsularis YAHARA3) Japan, Kagoshirna: Isl, Yakushima, JteatstLki 329e. L transnekoensis KiTAM. (SAsAKi) Republic of China CI"aiwan). Nantou: Mt. Nehg-kao, Ttimura et al. 23348, Takao: Mt, Kanzan, Okamoto 9. L makinoana KiTAM. (M-ciNo) Japan, Hyogo: Akou,Pak 993. Mie: Igaueno,6June I937, Okayama s･n.J PeeplesRepublicofChina, Yunnan: Kumming, Kayama et al, 1413. L grctcitis (DCI.) STEBBINS Bhutan,EasternHimalaya: GonChungnang-Punakha, Hlzra et aL 15990 CI[-I). L laevigata paLuME) SaHuLTZ-Bip. ex Japan. Kageshima: Tashiro, 24July 1919, 7lashire, MAxTM. s. n. Papua New Guinea. Western Highlands: Wabag Subdist. Hbagland et at. 6665 (L). L nen;aea (ZOLL. et MoRITZI) STEBBINs3) Malaysia. Sabah: Mt. Kinabalu, Sinith 557 (L). Papua New Guinea. Star Mts, W, Sepik, Vletcteamp 6577 (L), Lptesitla (MATr,) SviBBiNs3) PapuaNewGuinea. WesternHighlandsDist.: Mt. Kinkain, Vink l60sc (L). L sttbasaulis (KosiER, J) PA,K et KAwANo3) ,4) Papua New Guinea, East. Eastern Highlands Dist.: Mt. Wihelm, Shzith ANU li 129 (holotype: L!). L siamensis <KERR) STEBBINsS) Thai]and. Chiang Mai: Omkoi Distr., Kll7ana et at. r32257. i) All colloctions deposited at KYO unless otherwise noted. 2) Sectional classification fo1]ows KiTAMuRA (1956), 3) Species assignment in sections by the present authors, 4) LactLeca subaaaulis KosrER is transfbrred to the genus lbeeris as L szibticautis by the present authors tentatively. NII-Electronic Library Service The JapaneseSocletyJapanese Society forforPlant Plant Systematics ca Acta Phytotax. Geobot. VoL XLI, Nos. 1-3 observations were made on mQre than mature collection, in Critical five fruitsper which the endosperm was entirely absorbed and the embryo occupied the entire embryo sac. The only exception was 1hreris parva, in which only sornewhat immature fruits were pu bk by sc sl lb 1mec exc 1 ' 'glt't',･Wl･ - , /.., ..t, , ,zr.", vl'l ;:f,,l'i'1' ..;,;t-, Wim#fienFva'' -- 4C-lb t-,k,rw･--.ttin,ttt - ,;.// . ... ttt ,.tnytb., .tt ttt tt t tt ttttt t..t.t.'t'e. .. lt ,.,, .tt. Figs. 1-4. Fruit morpholegy and anatomy ofixeris, Fig. I: Scanning electron micrographs ofmature ofL debilis, Fig. 2: Diagram il]ustrating transverse sectionofL debilis.Fig. ffuit a 3: Diagram illustrating the structure of a single costa of I, debilis. Fig. 4: Longitudinal section of costa ef L ropens. cos, costa; em, embryo; exc, exocarp; lb, iibriform fiber cells (4C) ; inc, intercosta; mec, mesocarp; sl, fiber-selereid cells (4A, B); sc, seed coat; vb, vascular bundle.
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