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Venuloid Idioblastsin Pteris Systematicimplications Jr.*

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Venuloid Idioblastsin Pteris Systematicimplications Jr.* The JapaneseSocietyJapanese Society for Plant Systematics May, 1978 Acta Phytotax. Geobot. 33 Venuloid Idioblasts in Pteris and Their Systematic Implications W. H. WAGNER, Jr.* w. H. v fft- : i / Je s vvKoeelkse. ng$mHa In many ways anatomical characters relied upon more have been in pteridophytes than in spermatophytes in working out taxonomic relationships, especially during the early development of classificatien at the levels of genus and family. Nevertheless, we are still engaged in seeking new characters that can be exploited in solving controversial problems of relationships and this research continues. In this article I wish to discuss "false the peculiar veins'' that are found in the brake ferns, Pteris. Such structures have "spicular ``recurrent [Cfalse "Scheinnerven," been referred to as idioblasts," veins," veins," "sclerites," "venzaloid and other terms. As a name fbr this phenomenon I propose idio- blasts." This would cover all structures of various fbrms and homologies which produce the illusion of tiny veinlets scattered in the interveinal, submarginal, or sinus regiens of the lamina. They are not attached to the true veins. Ordinarily the venuloid idio- blasts can be seen with a high-powered hand lens or dissecting microscope with no difllculty, appearing as narrow Iines or streaks. In dried herbarium specimens they are often slightly raised. Such veinlet-like structures are widelv . scattered in the various groups of ferns, but are most familiar in such families and subfamilies as Marattiaceae in Marattiales and Hymenophylloideae and Vittarioideae in the Polypodiales. In some genera (e.g., 7Vichomanes, Angioptere's) they have been used fbr purposes of species discrimination. The presence of venuloid idioblasts has also been used as a source of evidence for relationships at the higher levels of genus and family. As part of a general study ef venuloid idioblasts in fern leaves, I wish to describe in this paper some of my observations in the genus Pteris. This is especially appropriate on this occasien because Professor Motozi TAGAwA probably dealt more than any other individual with the species of Pten-s that display these structures. Indeed he described several of these taxa, I wish here not only to expand the existing descriptions of the venuloid idioblasts of Pteris, but also to discuss some of their possible systematic im- plications. * Department of Botany, University of Michigan, Ann Arbor, MI 48le9, U. S. A. Research supported by NSF DEB75-03550. Taxonemic Applications ofFoliar Characters. I wish to acknowledge Florence S, NVAGNER for her constant assistance in this prQject. NII-Electronic Library Service The JapaneseSocietyJapanese Society for Plant Systematics 34 Acta Phytotax. Geobot. Vol. XXIX, Nos. 1-5 Generally speaking thc presence of venuloid idiob]asts in the brake ferns has gone unnoticed by the majority ef authors. BowER (1928) did not mention them in Pteris, nor did CopELAND or OGuRA In connection with Pterz's WALL･, (1947) (1972). .arewilleana "between HoLTTuM (1954) noticed that the veins on both surfaces are irregular short ralsed lines, commonly two rows between each pair of veins." In his synopsis of the genus Pteris in Japan, Ryukyu, and Taiwan, Wang-Cheung SmEH (1966) used what he termed [[false-veins ``Subsect. in mesophyll" te key out 2. Cadieri." He also adopted the "many ``few contrast false-veins" vs. faIse-veins'i as the primary division in his key to the seven taxa in this section, name]y P. -ptanzatensis (TAGAwA) TAGAwA, P. angustipinna muttijitla KuRATA, P. WALL., TAGAwA, P. tyukruensisTAGAwA, P. PoiR.,P. kidoi .arevilteana and P. cadieri CHRisT, A modification of this treatment was also used by SHiEH (1975) in the Flora of Taiw, an. , Description. The fbllowing descriptions of the venuloid idioblasts of Pterzs are based primarily on the species that represent the two extreme types of frond structure, namely P. multifdd with linear, entire major segments, and P. grevilleana with pectinate or deeply lobed major segments. The specimens used are in the University of Michigan Herbarium*. Frond scgments were cleared in sodium hydroxide aqueeus solution, and stained with tannic acid and ferric chloride. evident early this study that the idiohlasts actually do not occur in the It became in mesophyll, as has been reported; instead they are scattered in the upPer and lower epi- dcrmis, They are, however, somewhat overlapped by adjacent epidermal cells, as shown in Figure 1. The cross-sectional diameters of the epidermal cells and the idioblasts are sma}ler by one-halfto one-fifth than the true mesophyll cells. There are usually two to four layers of inesophyll parenchyma cells visible in section, and they are rounded and lack projections. ' Fig. 1. Cross-sectien of a leaf segrnent of Pteris gretiUeana showing position of venuloid idioblasts in the upper and lower epidermises, * Volcher specimEns'for anatomical preparations. (In University ofMichigan Herbarium unless otherwise indicated). Pterisgrevilteana-CmNA: Tonkin, Balansa 1978; western Tonkin, Bonin May 1B87. BoRNEo: Sarawak, Penrisson Mt., Brooks 14 in May 191O. P. cadien'-AssAM: Maknon Forest, Mann s.n., CopELA)m Herb. 13777; CHiNA: Kwantung, Tsing Wan Shan ["P. Iauii CHiNG sp. nov.'i], Lau 21S4; Hainan, Chim Mt., Lau 5469; Fokien Prov., M55, CopELAND Herb. 7575; FoRMesA, Chilung, Simoaawa 558. P. Fung multijida-A?AN: in-eno 1713; Hondo: YamamotQ in Settsu, 71)gasi709; KoREA: Cheju-Do, C7itLng Man Chi Shan, 71sang Shantung, Huang Shan, CHiNA: Kwantung, Fan Shiu Shan, Lau 2758; 264I4; UNiTED STATEs oF AMERicA: AIabama, Mobile, Dttkes in 1904 Fern Sec. Herb.). (]eiao2894; ' (Amer. P. mnttijidu × semipinnata?-OKiNAwA: Nuha-oyama, Amano 6738. P, wdouensis: RyuKyu, Amami-isl.: Naze-city, Slerizawa 11765. NII-Electronic Library Service The JapaneseSocietyJapanese Society for Plant Systematics May, 1978 Acta Phytotax. Geobot. 35 ' ' -t -t ttttt- - t The idioblasts are narrow, approximately 20-25 microns in diameter, and the secondary walls are extremely thick, the lumen being only 6-9 microns wide. Their lengths vary from O.! to 1.2 (rarely more) mm., and most of them are between O.5 and 1.0 mm. Thc longest idioblasts tend to be those located approximately midway between the costa and the margin of the segment, and the shortest are fbund near the cQsta, and especially along the margin. With few exceptions, they taper at both ends to pointed Fig, 2. View ofcleared segment of P. gmoilleana under Nomarski interference contrast lighting showing the similar refraction of true veins and false veins. ' tips. Fertile segments show the idioblasts running along the adaxial surface and the rolled margin above the sorus, but not in the false indusium itsel£ The idioblasts have a slightly scalloped appearance as seen under the dissecting rnicroscope and in clearings duq to the greater er lesser overlap of the numerous adjacent epidermal cells, The refractive characteristics of the false veins under Nomarski interference contrast illumi- nation are similar to those ef the true veins (Fig. 2). are nearly straight or only slightly curved. curved Usualiythe idioblasts Strongly or angular ones are most often fbund close tQ the costa in the angle of vein departure. Also, along the segment margins many of the idioblasts curve upward toward the apex of the segment. Although there are occasional individuals with more or Iess knobbed tips, only rarely do the idioblasts branch. On the abaxial surfaces one sometimes finds examplcs of forking where the tip comes into contact with the auxiliary cells of the stomates. Pteris grevilteana (Fig. 2) displays the most striking development of spicular cells, and these are immediately evident to the eye, even with a low powered hand lens. They are very numerous and crowded, and a typical interveinal area including both laminar surfaces totals roughly between 20 and 40 idioblasts (Fig. 2). [I]hey occur throughout the interveinal areas and there is no special tendency to cluster along the true veins. Numerous examples ofjoined pairs or triplets of idioblasts can be found. They are NII-Electronic Library Service The JapaneseSocletyJapanese Society forforPlantSystematlcs Plant Systematics 36 Acta Phytotax. Geobot. Vol. XXIX, Nos. 1-5 Fig. 3. Distribution of venuloid idioblasts in Pteris multti7de showing their tendeney to d'u'hte'r drotu'iui the truc vcins. joined in various ways. Although they may sometimes be contiguous for nearly or quite their fu11 length, they are commonly attached near tlteir tips. Carefu1 examination often is needed to estab]ish that what appears to be a single unusually long idioblast is in reality two or three connected along their tips. Pteris multijda (Fig. 3, 4) differs in having only relatively few idioblasts, usually O-5, rarely more, per interveinal area. These tcnd to cluster along the true veins, and this may make it possible to overlook them in a casual examination. Joined idiob]asts are rare in P. muttifda, except along the midrib, and curved examples are also not so commen as in I'. grewilleana. However, short individuals, as in the latter, do tend to occur mainly along the costa and along the margin. P. ryulpuensis (Fig. 5) has the fewest idioblasts of any J of the taxa I studied. wM-v#i/･,/////T,g/.i･.･ix,'g,f' ;･" " ,e/g･/itwh,i,s,ny-i,,/g'i"zee･g.ma,l,/sew.. ..gts/a･-#'ggli//r,\･isii:.'/･"'ps ' ci ww th: ' ' ' -' ca･srm.t,. -g' - de -i,//S,/s // .k.if/T,,/::gif\ nc . gs/i･g i/i'#./g:x:-g･eelk/4'},g･i sigS･,Y･･ist/j?{,//t-/r.lg' ss･/:ii//Y.l･g 'ik' grf,IillS･/l/r,//'//1,g･i･',,rr･il",l:11ii-l':111?l･,i/11/l,lti･l･Li,l'tlSl'･li#,I,//lilllli-,i･,-i,ii,.,E,11e..'ijll'l.,$',//tsi.l//f-,fl/Tit"Tl,g,kll,lw" mekg:msznys-&im' -g･-i -t/tde'-'i,･: ,k g'l'i s/k ti･ i･ i- iLe,l ;, L, l.
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