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Of Hortonia (Monimiaceae) Tree (Philipson The JapaneseSocietyJapanese Society forforPlant Plant Systematics ISSN eOel-6799 Acta Phytotax. Geobot. 47 (1): 53-59 (1996) Wood and Bark Anatomy of Hortonia (Monimiaceae) KWEON HEO Department of Ptant Resources, College ofAgriculturat Lije Science, Kangwon National Universit.v, Chuncheon 200-701, Kbrea Abstract. Wood and bark anatomy of Hortonia, which is generally considered the rnost primitive genus in Monimiaceae, was investigated to reconfirm early information and to fi11 up a gap in our knowledge, Remarkable features include: vesseL eiements very long and nar- row with exclusive]y scalariform perforation pEates; intei'vascular pittings in transitional state; fiber-tracheids non-septate and non-stratified with bordered pits; axial parenchyma apot- racheal; rays heterogeneous and multiseriate; phloem fibers present; phloem rays not dilated. Comparisons with other genera suggest that Hortonia is not the only genus retaining prkni- tive wood anatomical features in the family. Key words: bark anatomy, Hortonia, Monimiaceae, wood anatomy Received March 16, I996; accepted May 8, 1996 Hortonia Wight, comprising two or three species endemic to Sri Lanka, is a large shrub or small tree (Philipson, 1987, 1993; Mabberley, 1987) and is usually placed in its own subfamily Hortonioideae of Moni- miaceae, Hortonia has a few least specialized features such as alternate phyllotaxy, bisexual fiowers, shailow floral cup, tetrasporangiate anthers, and anther dehiscence by longitudinal slits. The genus has a distinct acol- pate pollen with striate scuipturing, which is similar to pollen of Dahtgre- nodendron of Lauraceae (Sampson, 1993). Recent authors thus consider Hortonia as the most primitive genus, linking Monimiaceae with Lauraceae (e.g., Endress, 1980; Takhtajan, 1980; Sampson, 1993). In a study of cladistic analyses of Lauraceae which we are now attempting, 13 wood and bark characters of Hbrtonia are included in a data matrix as a basis for comparison between Monimiaceae and genera of Lauraceae. However, four of the 13 characters are described without any illustration (see Money et al., 1950; Metcalfe and Chalk, 1950), and the nine remaining chara ¢ ters have not been investigated. The of this to purpose paper is provide full descriptions for the 13 characters of the wood and bark anatomy of Hortonia, and thereby to sup- plement a data matrix of the genus for cladistic analyses of Lauraceae, This will allow us to reduce the number of trees and to reach a better re- solution for phylogenetic relationships in Lauraceae. An evolutionary posi- tion of Hbrtonia in Monimiaceae will also be briefly discussed on the basis of anatomical characteristics of the wood and bark. NII-Electronic Library Service The JapaneseSocietyJapanese Society forforPlant Plant Systematics 54 Acta Phytotax. Geobot. Vol. 47 Materials and Methods Four wood specimens from Hortonia angustijblia and H. floribunda were examined in this study. Their collection data are presented in Table 1. Woods and barks were softened in boiling water for 24 hours. Trans- verse, radial, and tangential sections were cut at 15-20ym in thickness with sliding microtome, and stained by a combination of 1% safranin and fastgreen FCF or by safranin alone. After dehydration, sections were mounted with Entellan. Pit structure was observed by scanning electron microscopy. In order to measure sizes of individual xylem elements, weod pieces were macerated by Jeffrey's fluid (Johansen, 1940). Observations The wood and bark anatomy are very similar in Hortonia angustijbtia and H. floribunda. The following descriptions are common to the two spe- cies unless otherwise indicated. The terminology basically follows Radford et al. (1974). Wood rings are not is diffuse 1). Growth distinct(Fig.1).Wood porous (Fig. Porcs are angular, arranged mostly solitarily and very rarely in a multiple of cells. Pores are 40-60ptm in tangential diameter and 40-90ptm in radial i" thickness) (Fig･2)･ pore frequency ig.a.Mg,e,tefr,6.Wi'IB tt.hisno iYgl,1,,(f2,.-.4,:flM Vessel elements are very long (12eO-1500ptm long). End walls are TABLli 1. Hortonia species studied and their collection data Taxa Collection data fio;tonll;lngustijb-iiaTrimen Sri Lanka. Conservatory Forests. SJRw 29767 (MAD); wood collection #25939 (A); rree size unknown and sample about1 cm in diam. H, floribunda Wight Sri Lanka. Kandy distr. KunckLes. Nooteboom 3057 (L), Tree 4 m tall; sample about 1 cm in diam. Sri Lanktt. Kandy distr, Mirisketiya. Nooteboom 3406 (L), Tree 8 m tall; sample about 1 cm in diam. Sri Lanka. Censervatory FoTests. SJRw 29788 (MAD); wood collection #2594e (A); tree size unknown; sample about 1 cm in diam. NII-Electronic Library Service The JapaneseSocietyJapanese Society forforPlant Plant Systematics June 1996 HEO: Wood and bark anatomy of Hortenia 55 steeply inclined. Perforation plates are exclusively scalariform (Fig. 3) as already described (Money et al., 1950; Metcalfe and Chalk, 1950). The number of bars varies from 18 to 41, and they are rarely branched. Inter- vascular pits are more or less scalariform to opposite, and 5-15ptrn long in horizontal diameter (Fig. 4). Imperforated tracheal elements are fiber-tracheids, Fibers-tracheids are angular to polygonal in shape in transverse section (Fig. 2), long (1000-1200#m long) and thick-walled (8-10ptm thick) (Fig. 2). The fiber- tracheids constituting a ground mass of wood are non-septate as already described (Money et al., 1950; Metcalfe and Chalk, 1950) and not strati- fied, and the pits are small circular with slit-like apertures in radial section (Fig. 4). Spiral thickenings are not observed. Neither tyloses nor gum de- posits are observed in vessels, Axial parenchyma strands are apotracheal diffuse, polygonal in trans- verse section (Fig. 2), Banded parenchyma is not observed (Figs. 1, 2), Rays are fusiform in tangential view (Fig. 5), occupying much amount of wood, and multiseriate as already described (Money et al., 1950; Met- calfe and Chalk, 1950); they are heterogeneous, comprising square and procumbent cells in radial section (Fig. 6). The rays are more or less high (1000-1300stm long) and narrow (4-5 cells wide) in Hortonia angustijblia, whereas in H. floribunda they are lower (800-1000ym long) and wider (6-8 cells wide), Silica deposits are not formed but hexagonal crystals are present in the rays (Fig. 7). No oil cells were observed in and around the rays. Bark Phloem fibersare Phloem present. rays are mostly 3-4 seriate in tangential section, and dilated rays are absent. No aggregates of scierotic cells are present (Fig. 8). Disrupted islands of rays are present, bur they are not comparable with aggregates of sclereids in some Lauraceae (see Richter, 1981). No rhomboidal crystals are found in sclerotic cells of rays. Lignified parenchyma cells are not observed. Discussion Wood and bark anatomi'cal features of Hbrtonia can be summarized as follows. Features relevant to the 13 characters that were known without illustration or not known are indicated with numerals in bracket. Wood diffuse porous; vessel elements very long and narrow; (1) per- foration plates exclusively scalariform; intervascular pitting more o'r less scalariform to opposite, representing transitional state; fiber-tracheids (2) non-septate and (3) non-stratified (4) with bordered pits; (5) tyloses absent (as £ar as the materials observed are concerned); axial parenchyma apot- racheal no ,but (6) banded parenchyma present; rays heterogeneous and (7) multiseriate; (8) silica absent in rays, but crystals present in rays; no oil cells present in and around rays. In bark, (9) phloem fibers present; phloem rays mostly 3-4 seriate but NII-Electronic Library Service TheTheJapaneseSocletyfor Japanese Society for Plant Systematics 56 Acta Phytotax. Geobot. Vol. 47 FIGs. 1-4. Wood anatomy of HortonTafloribunda. 1. Transverse section (TS) showing diffusc porous wood without distinet growth rings, 2, TS shewing apotracheal-diffuse parenchyma and thick- walled fiber-tracheds. 3. Scanning electron micrograph showing vessels with scalariform perforation platcs, 4. Radial section tshowing scalariform te opposite arrangement of intervascular pits, Scale bars equal 100"m in Figs. 2, 3, and 4; 500ym m Fig. t. NII-Electronic Library Service The JapaneseSocletyJapanese Society forforPlantSystematlcs Plant Systematics June1996 HEOWood and barkanatomy of Hortoma 57 i/ l{,i#,fil; $mmislli-l,xi,31Iklli,,I/l/ }I l ll}1// @ "l XL "IIk eqgww,,.tw k ...g}ig/I-..},kl}l{ . i} "rede.-- lli FIas 5-8 Wood and bark anatorny of Hortonia flonb"nda 5 Tdngential seLtion of wood showing multisenate fusiform idys 6 Rddial section showing a heterogeneous iay 7 Rddidl section show ing crystdls in ray cells 8 Trangverse gect]on of bdrk Scale barb cqual 100Am m Figs 6 and 7, SOOym ln Figs 5 and 8 NII-Electronic Library Service The JapaneseSocietyJapanese Society forforPlant Plant Systematics 58 Acta Phytotax, Geobot, Vol. 47 (10) not dilated; (11) no aggregates of sclerotic cells found; (12) no rhom- boidal crystals present in sclerotic cells; (13) no lignified parenchyma pre- sent. As regards other Monimiaceae, the wood is studied to a varying de- gree for 20 of 34 genera (i.e., Atherosperma, Dqphnandra, Decai:ydendron, Doryphera, Dryadodaphne, Glossocalyx, Hedycailya, Hennecartia, Kibara, Laurelia, Levieria, Matthaea, Mollinedia, Monimia, Nl7muaron, Palmeria, Pe"mus, Siparuna, 7-timbourissa, and .Xlymatos), and the bark for six genera (i.e., Atherosperma, Hedycarlya, Laurelia, Monimia, Peumus, and Siparuna) (see Garratt, 1934; Money et al., 1950; Metcalfe and Chalk, 1950; Patel, 1973). In comparison with those of the other genera of Moni- miaceae based on information available, the wood of Hortonia has been considered primitive. Money et al, (1950, p. 400) stated that vessels and imperforate tracheal elements are of primitive form, and this statement is reiterated by later authors (e.g., Schodde, 1970; Philipson, 1993). It is cer- tain in the light of general evolutionary trends of the secondary wood that the long and narrow vessels with exclusively scalariform perforation, the intervascular pitting in transitional state, and the non-septate fibers, which are observed in Hortonia, all represent primitive states (Tippo, 1946).
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