IAWA Journal, VoL 14 (4), 1993: 391-412

WOOD ANATOMY OF TREES AND SIffiUBS FROM CHINA. VI. MAGNOLIACEAE

by

Chen Bao Liangt 1, Pieter Baas2, Elisabeth A. Wheeler3 and Wu Shuming4

Summary The wood anatomy offive genera of Mag­ ces; oil cells in rays mostly occur in the taxa noliaceae (59 native species, 2 introduced from tropical and subtropical provenances. species) of China is described. Although the Simple perforation plates are mostly present wood anatomy of this family is rather homo­ in the temperate taxa. Counter to trends for geneous, it is possible to identify most speci­ the dicotyledons at large, helical thickenings mens to genus. Magnoliaceae wood from are more common in tropical species than in China is characterised by diffuse-porosity, temperate species, and, when present, are scalariform to opposite vessel wall pitting, usually not distinct in deciduous species. scalariform perforations with few bars or in Key words: Magnoliaceae, Kmeria, Lirio­ some Magnolia species simple perforations, dendron, Magnolia, Mangiietia, Michelia, ground tissue fibres with distinctly to minute­ China, systematic wood anatomy, ecologi­ ly bordered pits, marginal parenchyma and cal wood anatomy, wood identification. heterocellular rays mostly with one marginal row of square/upright cells. Intervessel and vessel-parenchyma pits are almost exclusive­ Introduction ly opposite in the Liriodendroideae; they are Magnoliaceae are shrubs, small trees, or almost exclusively scalariform in the Magno­ trees, and range from tropical and subtropical lioideae, except for Magnolia section Rhyti­ to temperate areas. In China, most Magnolia­ dospermum in which pits are predominantly ceae occur in tropical to subtropical forests, a opposite. Although the wood anatomical char­ few species occur in temperate areas. The acters more or less overlap between Magnolia family Magnoliaceae can be divided into two and Manglietia, these genera are wood ana­ subfamilies: Liriodendroideae and Magnolioi­ tomically distinguishable. Wood anatomy is deae. There is uncertainty and controversy similar in the evergreen species of Magnolia about the delimitation of the genera of Magno­ and Michelia. Kmeria is the only genus in lioideae because of considerable overlap in which crystals were observed. Taxa from the characters (Dandy 1927; Keng 1978; Law tropics to subtropics tend to have longer and 1984; Nooteboom 1985; Qiu et al. in press). wider vessel elements, and a lower vessel fre­ For this study, we basically adopt Noote­ quency than those from temperate provenan- boom's 1985 concept of genera (Nooteboom t) The late Dr. Chen Bao Liang was a staff member of the Department of Biology, Zhongshan University, PRC, where he carried out a Ph.D. study on the taxonomy and wood anatomy of selected genera of the Magnoliaceae in China. This wood anatomical study was largely car­ ried out at the Rijksherbarium/Hortus Botanicus in 1990 and 1991. He died a few months after his return to China. The three co-authors wish to dedicate their contributions to this paper to his memory.

1) Department of Biology, Zhongshan University, Guangzhou, People's Republic of China. 2) Rijksherbarium/Hortus Botanicus, P.O. Box 9514, 2300 RA Leiden, The Netherlands. 3) Department of Wood & Paper Science, North Carolina State University, Box 8005, Raleigh, N.C. 27695-8005, U.S.A. 4) Department of Biology, Nankai University, Tianjing, People's Republic of China.

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Table 1. Genera of Magnoliaceae with numbers of species occurring in China, and studied wood anatomically.

Subfamily I Genus World China Number examined

Liriodendroideae Liriodendron 2

Magnolioideae Elmerrillia 5 0 0 Kmeria 2 1 Magnolia 110 32 20 (+ 1 U.S.A. native) Manglietia 24 18 12 Michelia 46 36 25 Pachylarnax 2 0 0

Total 190 87 59

1985), except that Manglietiastrum is merged tive to China (Oleaceae, Baas & Zhang 1986; into Manglietia instead of Magnolia (Chen & Theaceae, Deng & Baas 1990; Rosaceae, Nooteboom in press). Zhang & Baas 1992; Ulmaceae, Zhong et a1. Some species of Magnoliaceae provide 1992; Anacardiaceae, Dong & Baas 1993). useful timber for the local people; many spe­ cies are used as ornamentals. Table 1 gives Material and Methods the number of species for China (cf. Chen & Wood samples were collected by the first Nooteboom in press) and the whole world and fourth author, as well as obtained from (Dandy 1964). General accounts of Magno­ various institutional wood collections and liaceae wood anatomy include those by universities in China. Some of the vouchers McLaughlin (1933), Canright (1955), and were identified by Chen Baoliang. Wood Metcalfe (1987). Fossil wood with character­ samples were collected mostly from mature istics of Magnoliaceae occurs in the Upper stems, with a few samples from young stems. Cretaceous of North America (Page 1970). Some of the slides used in this study are per­ By the middle Eocene, there were woods with manently housed in China. For those samples characteristics of the genera Liriodendron and with slides at Leiden, the age of the stem is Magnolia (reported from North America: Scott estimated and this information is given in the & Wheeler 1982; Cevallos-Ferriz & Stockey generic tables: * = juvenile wood samples. 1990). Wood of Michelia is reported from the In this paper, nomenclature follows the Oligocene of Japan (Suzuki 1976). Publica­ Revision of Magnoliaceae from China (Chen tions specifically about wood anatomy of & Nooteboom in press). Sections and mace­ Chinese Magnoliaceae include those by Ho rations for light microscopic studies were pre­ (1949), Tang (1973), Chang (1974, 1982, pared in the usual manner (cf. Baas & Zhang 1984), Liu et aI. (1987), Wu et aI. (1993), 1986). Samples for scanning electron micro­ Wu & Li (1988, 1989) and Chen (1989). scopy were prepared according to Exley et aI. The present paper deals with the systematic (1977). Conventions for descriptions and for and ecological wood anatomy and micro­ determination of quantitative values are the scopic wood identification of Magnoliaceae same as those explained by Baas and Zhang from China and is part of a by-family series (1986) or were slightly modified to bring on the wood anatomy of trees and shrubs na-

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them in line with recommendations of an ing vessels are typically in short radial multi­ IAWA Committee (1989). Vessel element and ples, with some oblique or tangential pairs fibre lengths are not given for those samples and some clusters. available only as sections. Frequency and element size - Vessel fre­ quency ranges from 12-158/mm2. Average Survey of wood anatomical character tangential vessel diameter varies from 30-130 states in the Magnoliaceae from China 11m, and average vessel element length from 570-1480 11m. Generally, within a species, Growth rings (Figs. 1-6) vessel element length and vessel diameter in­ Growth rings in the Magnoliaceae from crease with stem diameter. Vessel frequency China are distinct to faint and usually marked is very variable within individual genera and by marginal parenchyma bands and changes also within species of Magnolia and Michelia. in the radial dimensions of the ray parenchy­ This variation is related to stem diameter, en­ ma cells. Sometimes, particularly in tempe­ vironment, and habit. rate species, the last formed latewood fibres Perforations (Figs. 13-16) - Perfora­ are also radially flattened. tions are predominantly scalariform. Forty­ five of the 59 species (76%) have exclusively Vessels scalariform perforations. The highest bar Distribution and grouping (Figs. 1-6)­ numbers (up to 28) were found in Kmeria Magnoliaceae are diffuse-porous. The per­ septentrionalis. There usually are fewer than centage of solitary vessels is extremely vari­ able and ranges from 12-80%, the remain- (text continued on page 398)

Legends of Figures 1-24:

Figs. 1-6. TS, x 50. - 1: Magnolia coco (shrub), wood diffuse-porous with indistinct growth ring boundaries and narrow vessels. - 2: M. sieboldii, wood diffuse-porous with distinct growth ring boundary, most vessels in radial multiples. - 3: Kmeria septentrionalis, wood diffuse-porous. - 4: Liriodendron chinense, wood diffuse-porous, distinct growth ring boundary, vessels soli­ tary and in radial or oblique multiples. - 5: Michelia martinii, wood diffuse-porous, vessels solitary and in radial multiples and clusters, parenchyma marginal and also in zonate bands. - 6: Manglietia grandis, vessels solitary and in radial multiples.

Figs. 7 & 8. RLS, x 510. - 7: Manglietia grandis, vessel-ray pits scalariform and unilaterally com­ pound with reduced borders. - 8: Liriodendron chinense, vessel-ray pits opposite and unilater­ ally compound. - Figs. 9 & 10. TLS, x 125. - 9: Magnolia coco, rays 1-4 cells wide, oil cell (arrow). - 10: Manglietia insignis, rays 1-2 cells wide, scalariform intervessel pits. - Fig. 11. Liriodendron chinense, ray with oil cell (arrow), TLS, x 510. - Fig. 12. Manglietia grandis, rays heterocellular, with one row of square to upright marginal cells (bottom), RLS, x 125.

Figs. 13-16. RLS, all SEM. - 13: Manglietia megaphylla, perforation plate scalariform with seven bars; x 370. -14: Michelia martinii, perforation plate scalariform with two bars; x 390. - 15: Magnolia hypoleuca, perforation plate simple with irregular outline; x 690. - 16: Michelia hypolampra, perforation plate simple; x 475. - 17: Manglietia dandyi, intervessel pits scalari­ form; x 1200. -18: Magnolia sieboldii, intervessel pits opposite; x 1900.

Figs. 19-24. All SEM. - 19: Michelia martinii, vessel wall with coarse helical thickenings; x 1700. - 20: Manglietia sinica, vessel wall with discontinuous helical thickenings; x 1700. - 21: M. insignis, septate fibres; x 550. - 22: M. insignis, tylose is in fibre; x 2000. - 23: M. insignis, tylose is in vessel; x 435. - 24: Magnolia championii, oil cell in ray; x 370.

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10 bars, and the bars are widely spaced. also occur on the vessel walls in Kmeria and Mixed simple and scalarifonn plates occur in two species of Manglietia. seven species: Magnolia amoena, M. hepta­ Tyloses and vessel contents (Figs. 22,23) peta, M. sargentiana and M. zenii (all of sect. - Tyloses are of common occurrence in ves­ Yulania); Manglietia pachyclada; Michelia ve­ sels, and are occasional in fibres. They are lutina and M. hypolampra. Irregularly forked usually thin-walled and do not fill the vessels, bars rarely occur in some species. Exclusive­ but appear similar to cross-walls. Gums or ly simple perforations only occur in some other deposits were not found. deciduous species of Magnolia. Usually, the end walls of vessel elements are oblique with Fibres (Figs. 21, 22) short or long tails. With the exception of some fibres in Mang­ Wall pitting (Figs. 17, 18) -Lirioden­ lietia insignis, fibres are nonseptate, with bor­ dron has exclusively opposite intervessel pits, dered pits arranged in a single row in the some Magnolia species have predominantly radial walls, the pits occur occasionally in the opposite pits (M. officinalis and M. hypo­ tangential walls. Average pit chamber diam­ leuca of sect. Rhytidospermum, and M. sie­ eter ranges from 4-7 I!m (total range 2-11 boldii of sect. Oyama). In the other Magno­ I!m). Fibre wall thickness varies from 1.5-8 liaceae intervessel pits are exclusively to pre­ I!m, on average from 3-5 I!m and is of very dominantly scalariform, sometimes with a little diagnostic value. Average fibre length few opposite pits. Pits are always non-ves­ ranges from 1010-2030 I!m (total range tured. Opposite pits are rounded to slightly 530-2980 I!m). The ratio of fibre to vessel oval, with an average horizontal pit chamber element length (F/V ratio) varies from 1.28 diameter ranging from 3-21 I!m (Fig. 18). to 2.30. The ranges of both fibre length and The apertures are generally slit-like. In scala­ F IV ratio overlap considerably within and riform pits the average horizontal pit chamber between genera, so that they cannot be used diameter ranges from 9-57 I!m (total range as taxonomic characters. Tyloses occur in 5-100 I!ffi) (Fig. 17). Vessel-ray pits are com­ Manglietia insignis, M. pachyclada and M. monly large and simple or half-bordered with sinica. reduced borders, typically scalariform, some­ times scalariform-opposite, often unilaterally Axial parenchyma (Figs. 1-6, 10) compound with a row of rounded to suboval Axial parenchyma typically is banded; in parenchyma pits subtended by a single elon­ woods with distinct growth rings marked by gate vessel pit. In the woods with predomi­ radial flattening of the fibres, these bands are nantly opposite pits, the vessel-ray paren­ obviously marginal. In other woods the bands chyma pits are also opposite and with very likely are marginal, but are not associated reduced borders. Vessel-axial parenchyma with any changes in fibre dimensions or, at pits are more or less similar to vessel-ray pits. times, with changes in ray cell radial dimen­ Wall thickness and sculpturing (Figs. 19, sions (as viewed in cross section, see Fig. 6). 20) - Vessel walls are thin (1.5-3I!m). The Thus, it is difficult to determine whether these presence or absence of helical thickenings on bands are correlated with growth cycles, and the vessel walls is a valuable diagnostic char­ they might better be termed zonate. Band acter for Chinese Magnoliaceae. They vary width ranges from quite narrow (1-2 cells from distinct to quite faint, being only visible wide in some species) to broad (greater than with SEM or phase contrast. Liriodendron 4 cells, up to 6). In a few species there rarely lacks helical thickenings; conversely, most occurs some scanty paratracheal and diffuse Michelia species have helical thickenings. In apotracheal parenchyma. Strands are usually Magnolia the occurrence of helical thickenings 3-7 cells long (total range 2-10). seems related with phenology: all evergreen species examined show various degrees of Rays (Figs. 9-12) helical thickenings; however, only 3 of the Most rays are heterocellular. The body ray 14 deciduous species examined have helical cells of multi seriate rays are almost always thickenings. Very faint helical thickenings procumbent, but in young samples of Kmeria

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they may be mixed with square and upright al to scalariform, 3-21 11m in horizontal cells. In mature wood samples most rays have diameter, with slit-like apertures. Vessel-ray but a single marginal row, but they are ac­ and vessel-parenchyma pits opposite, with companied by some rays with 2-4 marginal reduced borders to almost simple, or rarely rows of square or upright cells. In young unilaterally compound. Helical thickenings stems rays may have up to 14 rows of square absent. or upright marginal cells. Ray frequency Fibres 1660 (1320-2240) 11m long, thin­ ranges from 4-12/mm and is of very limited to thick-walled, with small bordered pits, 5-8 diagnostic value. Ray width ranges from 1-3 11m in diameter, mainly confined to the radial (6) cells. Ray height varies from 6 up to 48 walls. cells or 320 to 3400 11m. Uniseriate rays are Parenchyma in terminal bands of 2-6 cells infrequent in mature wood samples, and when wide, in strands of 6-8 cells. present quite low (fewer than 4 cells high), Rays 5-6/mm, 1-3(-4) cells wide, 6- but are common and composed exclusively 48 cells or up to 1160 11m high, body ray cells of upright cells in young stems. procumbent, with 1 row of upright and/or square marginal cells. Crystals Oil cells occasionally present in ray mar­ Crystals are very rare. Minute acicular gins ofFHOw 11640 only. crystals have been noted only in some ray Crystals absent. cells of Kmeria. Notes: 1. This is, to our knowledge, the first record of oil cells in Liriodendron; such Oil cells (Figs. 11,24) cells are absent from L. tulipijera of eastern No oil cells were observed in Kmeria or North America. Manglietia, but oil cells were observed in the 2. Canright (1955) suggested that wood of rays of some species of all other genera. Oil L. chinense might be distinguished from the cells are reported for the first time in Lirio­ closely related L. tulipijera by vessel diam­ dendron chinense, where they are of very eter and ray size (smaller in L. chinense), but rare occurrence. They also occur in the axial noted that "it is entirely possible that insuffi­ parenchyma of some species of Michelia. cient sampling makes these differences ... more apparent than real." In this study, the mean vessel tangential diameter for L. chi­ Generic wood anatomical descriptions nense was 55 11m which overlaps with the values Stark (1954) reported for L. tulipijera Subfamily Liriodendroideae (53-66 11m). Canright reported that rays in L. chinense rarely were over 20 cells high, Liriodendron L. (Figs. 4, 8, 11) while rays in L. tulipijera often reached Material examined: L. chinense (Hemsley) heights of 25-35 cells. We found rays as Sarg.: S. China, Fujian s.n., FHOw 11640. high as 48 cells in L. chinense. Differences in Deciduous trees, from temperate or sub­ vessel diameter and ray size do not seem to tropical regions. be reliable means of distinguishing the two Growth rings distinct, marked by terminal species. parenchyma. Wood diffuse-porous. Vessels 68-103/mm2, 31-65% solitary, remainder Subfamily Magnoloideae in radial multiples of 2-4(-7), occasionally in oblique to tangential multiples of 2-4, and Kmeria (Pierre) Dandy (Fig. 3) clusters, oval to round or slightly angular, tan­ gential diameter 50-65 (35-80) 11m, radial Material examined: K. septentrionalis Dan­ diameter up to 30-115 11m, walls 2-3 11m dy: SE. Yunnan, B.L. Chen 87F169. thick. Vessel element length 870 (610-1130) Evergreen trees, in tropical to subtropical 11m. Perforations scalariform in oblique end forests, on limestone, at about 500 m altitude. walls, with 4-6 (1-8) bars. Intervessel pits Growth rings distinct, marked by margi­ nonvestured, opposite, sometimes transition- nal parenchyma. Wood diffuse-porous. Ves-

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Table 2. Variation in selected wood anatomical characters of Magnolia.

Ph VF %sol TD B VEL FL RW HOC

Subgenus Magnolia

Section Gwillimia, tropical except for M. delavayi

M. albosericea evg 28 46 50 6 940 1610 2 (3) + + Chun & C.R. Tsoong NK 205, Hainan M. championii evg B.L. Chen S560, Hongkong, 1900 m* 35 75 45 10 680 1090 2 (3-4) + - 87F163, Yunnan, 1800 m 20 28 35 6 670 1460 2-4 + + NK 130, Hainan 56 24 53 5 703 1574 2-3 + + SCCFC 2720 25 40 45 4 790 1630 2-3 + + M. coco evg B.L. Chen 86S561, Guangzhou* 21 56 30 9 570 1074 2-4 + + B.L. Chen 21, Guangzhou 23 36 32 9 570 1230 3-4 (5) + - NK 174, Hunan 84 18 32 8 577 1038 2-6 + - M. delavayi evg SWFC s.n., Yunnan 70 12 50 4 680 1390 4 (5) + - NK 113, Yunnan 60 20 60 5 637 2-4 (5) + - M. henry, NK 190, Yunnan evg 19 47 78 7 882 1770 2-3 + - M. phanerophlebia evg B.L. Chen 87TOOl, Yunnan, 725 m* 22 50 66 8 659 1220 2-3 (4) + -

Section Rytidospermum M. hypoleuca (M. obovata) dcd 58 55 74 0 2 (3) native to Japan, FHOw 2102 M. officinalis dcd NK 151, Hunan, 700 m 85 32 60 0 7(j) 1080 2 NK 1(j), Hunan, 700 m 66 35 55 0 670 1510 2-3

Section Oyama M. globosa dcd YIP s.n., Yunnan 67 58 58 0 842 1300 2-3 NK 111, Yunnan, 1500 m 36 22 51 704 1264 2-5 + - M. sieboldii, NK 176, Jilin dcd 80 20 50 13 630 810 2-3

Section Theorhodon M. grandiflora (U.S.A. native) evg NFC s.n., Nanjing 158 34 49 6 950 1300 2-3 + + FHOw 11245 60 42 54 7 2-3 + + NK 204, Guangzhou 52 21 58 880 1948 2-5 + +

Section Gynopodium M. nitida evg NK 119, Yunnan, 1550 m 60 49 67 8 1479 1890 2-3 (4) + - NK 177, Sichuan 41 22 81 4 658 1613 2-3 + - var. lotungensis evg GIP97020 47 53 70 8 1170 2030 2 (3) + + NK 128, Hunan 127 14 64 9 1155 2100 2-3 + -

Section Alcimandra M. cathcartii, NK 125, Yunnan evg 55 30 60 4 755 1500 2-3 + -

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(Table 2 continued)

Ph VF %sol TD B VEL FL RW H OC

Subgenus Yulania

Section Yulania (deciduous, temperate) M. amoena, NK 173, Hunan dcd 80 65 50 0.2 605 990 2 (+) + M. dawsoniana, NK 173, Hunan dcd 47 36 71 4 600 1380 2-3 (+) - M. heptapeta dcd NK 106, Yunnan 62 44 73 0.3 860 1723 2 Zu s.n., Hunan 40 49 84 0 900 1418 2 (+) - M. sargentiana, NK 171, Sichuan dcd 25 50 80 0.1 920 1450 2 + - M. sprengii, NK 131, Hunan dcd 36 45 80 0 679 1298 2 (3) M. zenii, NK 175, Hunan * dcd 60 40 50 0.2 685 1385 2-3

Section Buergeria M. cylindrica, NK 172, Anhui * dcd 45 30 65 6 730 1440 2-3 M. kobus (native to Japan) dcd NFC S.n. 84 54 50 0 2-3 Uw253 100 43 40 0 2-3 + -

Section Tulipastrum M. quinquepeta dcd NFC s.n. 68 50 65 0 2-3 H - NK 152, Henan 113 71 56 0 813 1300 2-3 + - M. soulangiana, NSw RI130-129 dcd 84 47 56 0 2-3 - +

Ph =phenology; evg =evergreen; dcd =deciduous; VF = vessel frequency; %sol =per cent solitary vessel; TD = mean tangential diameter (Jlm); B =average number of bars per scalariform perforation; VEL = mean vessel element length (Jlm); FL = mean fibre length (Jlm); RW = multiseriate ray width; H = helical thickenings in vessels, OC = oil cells in rays. * =juvenile wood sample. se1s 40-134/mm2, 42-65% solitary, re­ Fibres 1010 (730-1350) Jlm long, mostly mainder in radial multiples of 2-6, occasion­ thin-walled, with small bordered pits, 2-3 ally in oblique to tangential multiples of 2-4, Jlm in diameter, mainly confined to the radial and clusters, round to oval or slightly angu­ walls. lar, tangential diameter 55 (40-80) Jlm, ra­ Parenchyma mostly in marginal or zonate dial diameter up to 20-95 Jlm, walls 1.5-2.5 bands of 2-4 cells wide, rarely scanty para­ ).tm thick. Vessel element length 740 (460- tracheal, in strands of 2-8 cells. 870) Jlill. Perforations scalariform in oblique Rays 5-7/mm, 1-4 cells wide, 11-19 end walls, averaging 11 (7-28) bars, occa­ (10-39) cells high, up to 1240 ).tm high, all sionally scalariform perforations combined body ray cells procumbent or mixed with a with reticulate plates. Intervessel pits nonves­ few upright and/or square cells, with 1-4 tured, scalariform, sometimes opposite, 16- rows of upright and/or square marginal 56 ).tm in horizontal diameter, with slit-like cells. apertures. Vessel-ray and vessel-parenchyma Oil cells absent. pits opposite to scalariform, bUf with strong­ Minute acicular crystals present in some of ly reduced borders, occasionally unilaterally the ray cells, several to many per cell. compound. Very fine helical thickenings Note: To our knowledge, the only other (barely visible with the light microscope) published description of wood of Kmeria is present throughout the vessel elements. that of Canright (1955) who examined a 1.5-

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year-old twig of Kmeria duperreana from In­ and narrow (1-2 cells wide) marginal bands. donesia. This species is similar in vessel diam­ Wood diffuse-porous. Vessels 19-158/mm2, eter (52 Ilm), but has fewer bars per perfora­ 12-75% solitary, remainder in radial multi­ tion plate (8-12). In other Magnoliaceae, the ples of 2-4 (6-17), occasionally in oblique number of bars per perforation plate decreas­ to tangential multiples of 2-3, and clusters, es with increasing stem diameter (Canright angular or rounded in cross section, angular 1955), and so the lower number of bars per outlines more obvious in narrow stems; mean perforation plate in the twig of Kmeria du­ tangential diameters 30-84 (20-100) Ilm, perreana compared to K. septentrionalis may radial diameter up to 50-160 Ilm, walls 3 be a consistent difference between the species. (4-5) Ilm thick. Mean vessel element length 570-1480 (range 310-1790) Ilm. Perfora­ tions exclusively scalariform in M. alboseri­ Magnolia L. (Figs. 1,2,9,15,18,24; Table 2) cea, M. cathcartii, M. championii, M. cylin­ drica, M. coco, M. dawsoniana, M. delavayi, Material examined (21 species, 1 variety; M. grandijlora, M. nitida, M. nitida var. 10- 39 samples): tUngensis, M. phanerophlebia and M. sie­ Subgenus Magnolia: boldii, with 1-13 (1-16) bars, distance be­ Section Gwillimia - M. albosericea Chun tween bars 11-20 (5-60) Ilm; perforations & C.H. Tsoong; M. championii Benth.; M. scalariform and/or simple in M. amoena, M. coco (Lour.) DC.; M. delavayi Franchet; M. heptapeta, M. sargentiana and M. zenii; per­ henryi Dunn; M. phanerophlebia B. L. Chen; forations exclusively simple in M. globosa, Section Rytidospermum - M. hypoleuca M. hypoleuca, M. officinalis, M. kobus, M. Sieb. & Zucc.; M. officinalis Rehder & E.H. quinquepeta and M. soulangiana, the simple Wilson; perforation plates with oval outline. Interves­ Section Oyama - M. globosa Hook. f. & sel pits nonvestured, scalariform or scalari­ Thomson; M. sieboldii K. Koch.; form-opposite, mostly opposite in M. hypo­ Section Theorhodon - M. graiuJijlora L. leuca and M. officinalis of sect. Rhytidosper­ (native to eastern United States); mum and M. sieboldii of sect. Oyama; oblong, Section Gynopodium -M. nitida W.W. 10-43 (5-80) Ilm in horizontal diameter, Smith; M. nitida var. lotungensis (Chun & with slit-like apertures. Vessel-ray parenchy­ C.T. Tsoong) B.L. Chen & Noot.; ma pits often restricted to marginal rows in Section Alcimandra -M. cathcartii (Hook. evergreen species including sections Gwilli­ f. & Thomson) Noot.; mia, Theorhodon and Gynopodium, gener­ ally throughout the entire ray in the remainder Subgenus Yulania: of the species, and mostly large and simple or Section Yulania - M. amoena Cheng; M. with reduced borders, typically scalariform dawsoniana Rehder & E.H. Wilson; M. hep­ oval-oblong to rounded, often unilaterally tapeta (Buchoz) Dandy; M. sargentiana Reh­ compound with a row of round ray paren­ der & E.H. Wilson; M. sprengeri Pampan.; chyma pits subtended by a single elongate M. zenii Cheng; vessel pit. Vessel-axial parenchyma pits more Section Buergeria - M. cylindrica E.H. or less similar to vessel-ray pits. Helical thick­ Wilson; M. kobus DC. (M. praecocissima enings distinct throughout vessel elements in Koidz.); the species of sections Gwillimia, Theorho­ Section Tulipastrum - M. quinquepeta don, Gynopodium and Alcimandra, absent Buchoz; M. soulangiana Soulange ex Thieb. from species of sect. Rytidospermum. Heli­ Evergreen or deciduous trees or shrubs cal thickenings, if present, very faint in some from tropical to temperate regions, from low­ species of sections Yulania and Buergeria of lands up to 2800 m. subgenus Yulania. Thin-walled tyloses usu­ Growth rings distinct or faint, marked by ally present. marginal or zonate parenchyma; in temperate Mean fibre length 810-2100 (range 540- species (e.g. all of sect. Yulania) rings dis­ 2840) Ilm long, FIV ratio 1.28-2.30, thin­ tinct and marked by radially flattened fibres to medium thick-walled, with bordered pits

Downloaded from Brill.com10/06/2021 05:49:47PM via free access Chen, Baas, Wheeler & Wu - Wood anatomy of Magnoliaceae from China 403 of 4-5 (11) 11m, mainly confined to the ra­ 6. Based on the samples available, wood dial walls. Fine helical thickenings present in of section Rytidospermum is distinct because some or many fibres in M. albosericea (SCCFC intervessel pits are predominantly or com­ 2880), M. officinalis (NK 160), M. nitida monly opposite and oil cells are absent. Some (NK 119), M. cathcartii (NK 125), M. daw­ additional samples were examined to see if soniana (NK 206), M. sargentiana (NK 171), this combination of character states occurred M. zenii (NK 175), M. cylindrica (NK 172) in other species in the section. Magnolia tri­ and M. kobus (NFC s.n.) petala (Uw 354, BWCw 8418), M. fraseri Parenchyma mostly in marginal or zonate (BWCw 8050, BWCw 8359), and the Asian bands of 1-11 cells wide, in addition very M. macrophylla subsp. ashei (Aw 8979) and rarely also diffuse apotracheal and scanty M. macrophylla (Aw 1627) also have predom­ paratracheal in 2-4 (8) celled strands. inantly opposite pits and lack oil cells, but Rays 4-12/mm, 1-3 (5) cells wide, 6-37 they differ from M. hypoleuca and M. offici­ (3-60) cells or up to 770-2540 11m high, nalis in having exclusively scalariform perfo­ typically heterocellular with procumbent body rations (data from Stark 1954: 1-17 bars in ray cells and 1-2 (3-14) rows of square to M.fraseri; 2-10 bars in M. tripetala). upright marginal cells. The number of margi­ 7. A recent phylogenetic analysis of the nal rows is greater in the narrower stems. temperate species of Magnolia (Qiu et aI., Oil cells occur in varying frequency, gen­ unpub. ms.) using data for chloroplast DNA erally as marginal cells, in rays of M. albo­ restriction sites indicated that sect. Rhytido­ sericea (SCCFC 2880), M. championii (com­ spermum was not a monophyletic group. The mon, B.L. Chen 87F163, SCCFC 2720), distribution of perforation plate type within M. coco (B.L. Chen 21, 86S561), M. gran­ the section is not entirely in accord with the diflora (FHOw 11245, NFC s.n.), M. hep­ cpDNA analysis. The cladistic analysis found tapeta (ZU s.n.; absent from NK 106), M. that M. tripetala (from the southeastern U. S.) nitida var. lotungensis (GFSI 97020, SCAU was sister to the three Asian taxa, M. rostrata, 2612, but absent from M. nitida, NK 119), M. officinalis, M. hypoleuca. Magnolia tripe­ M. soulangiana (not common, NSW RI130- tala has exclusively scalariform perforations; 129). M. officinalis has predominantly simple per­ Crystals absent. forations (a few scalariform present); M. hy­ Notes: 1. Exclusively simple perforation poleuca has exclusively simple perforation plates are restricted to deciduous species of plates. It is possible that within this group, sections Rytidospermum, Oyama, Yulania, the distribution of simple and scalariform per­ Buergeria and Tulipastrum. forations is not phylogenetically meaningful. 2. Mixed simple and scalariform perfora­ tion plates were observed in 4 of the 6 spe­ Manglietia Blume (Figs. 6, 7, 10, 12, 13, cies of sect. Yulania. 17, 20-23; Table 3) 3. Ten of the 12 species with helical thick­ enings in vessels are from the tropical and Material examined: 12 species, 1 variety; subtropical regions, almost exclusively in 26 samples; M. aromatica Dandy; M. conifera evergreen species. Dandy; M. dandyi (Gagnep.) Dandy ex B.L. 4. Oil cells mostly occur in the tropical and Chen & Noot.; M. fordiana Oliver; M. for­ subtropical species. They were not observed diana var.forrestii (W.W. Smith ex Dandy) in any samples of sections Rytidospermum, B. L. Chen & Noot.); M. grandis Hu & Oyama, Yulania, and Buergeria. Oil cells are Cheng; M. insignis (Wall.) Blume; M.lucida of variable occurrence in some species (pres­ B.L. Chen & S.C. Yang; M. megaphyUa Hu ent in some samples, absent in others). & Cheng; M. mota Dandy; M. pachyclada 5. Canright (1955) reported that oil cells c.Y. Wu, spec. nov.; M.pachyphylla Chang; only occur in the West Indian species of Mag­ M. sinica (Law) B.L. Chen & Noot. nolia, but they were observed in 7 of the 20 Evergreen trees from tropical to subtropi­ species examined for this study. Canright did cal regions, extending into warm temperate not have material of these species. regions, at altitudes of 500-2000 m.

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Table 3. Variation in selected wood anatomical features of Manglietia.

VF %sol TD B DB VEL FL RW H

M. aromatica NK S.n. 25 36 82 6 13-19 750 1390 2-3 M. conifera B.L. Chen 87FI49 27 57 68 4 16-27 850 1950 2 NK 164 72 49 57 12 8-16 1380 1990 2 (+) M. dandyi B.L. Chen 87F184 * 40 74 40 5 13-19 980 1530 2 NK200 79 88 52 5 15 555 1299 2-4 M.fordiana YFB W617 52 40 56 4 8-16 890 1540 2 GlF 89455 14 56 87 4 16-35 920 1840 2 NK 165 73 15 65 6 10 853 1707 2-4 NK 186 63 6 55 2 20 816 1402 2 (3-4) NK 189 47 16 91 3 28 888 1660 2 (3-4) M.fordiana var.jorrestii NK 115 43 33 71 6 13-14 1080 1830 2 NK 110 75 40 60 4 16-27 840 1470 2 M. grandis B.L. Chen GS86191 26 23 79 8 13-16 880 1920 2 + NK 117 33 27 83 7 19 867 1927 2 M. insignis B.L. Chen 87F185 * 46 54 67 7 8-16 960 1440 2 B.L. Chen GS86186 * 50 53 61 4 19-27 1040 1800 2 (3) NK 114 58 33 70 10 19 911 2182 2 M.lucida B.L. Chen 87T002 23 21 93 6 8-16 820 1390 1-2 M. megaphylla B.L. Chen 87F181 35 31 88 5 19-24 960 1770 2 NK 126 30 15 77 6 16-28 730 2050 2 M. mota GSF 1266 38 52 53 4 16-32 950 1690 2 + NK 138 80 30 55 3 13-32 840 1390 2 (3) (+) M. pachyclada B.L. Chen GS86185 34 29 56 0-8 19-29 860 1860 2 (3) M. pachyphylla GHSI1246 68 55 55 5 8-27 810 1660 2 M. sinica B.L. Chen 87F180 19 58 93 7 21-24 1210 2010 2-3 + NK 123 20 40 100 5 16-21 1080 1920 2-3 (4) +

VF = vessel frequency; %sol = per cent solilary vessel; TD = mean tangential diameter (/lm); B = average number of bars per scalariform perforation; DB = dislance between bars (/lm); VEL = mean vessel clement length (/lm); FL = mean fibre length (Jlill); RW = multiseriate ray width; H = helical thickenings in vessels. * = juvenile wood sample.

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Growth rings indistinct to faint, marked Notes: 1. Septate fibres, which were re­ by marginal or zonate parenchyma. Wood corded in M. mota by Ho (1949), were absent diffuse-porous. Vessels 14-80/mm2, 6- from our material. It is possible that there 88% solitary, remainder in radial multiples of were tyloses in the fibres, which easily can 2-4(-5), occasionally in oblique to tangen­ be mistaken for septae. tial multiples of 2-3, and clusters, usually 2. Simple perforations (in M. pachyclada) angular, more rarely rounded in cross sec­ are here recorded for the first time in Mang­ tion, mean tangential diameter 40-100 (range lietia. However, simple perforations are ex­ 25-160) j.l.m, radial diameter up to 65-190 tremely rare and likely to be overlooked in j.l.m, walls 3-6 j.l.m thick. Mean vessel ele­ sections, so that this characteristic would not ment length 555-1380 (range 460-1950) be of value in distinguishing M. pachyclada j.l.m. Perforations exclusively scalariform in from other species of Manglietia. most species, with 3-7 (1-13) bars, bars oc­ 3. The vessel diameters (120-155 j.l.m in casionally irregularly branched, distance be­ mature) Canright (1955) reported for Mang­ tween bars 14-22 (8-35) j.l.m; perforations lietia are greater than observed here. rarely simple in M. pachyclada. Intervessel pits nonvestured, scalariform or scalariform­ Michelia L. (Figs. 5, 14, 16, 19; Table 4) opposite, oblong, 11-57 (5-83) j.l.m in hori­ Material examined: 25 species; 48 sam­ zontal diameter, with slit-like apertures. Ves­ ples. sel-ray pits mostly large and simple or with Section Michelia - M. aenea Dandy; M. very reduced borders, typically scalariform alba DC.; M. cavaleriei Finet & Gagnep.; M. oval-oblong to rounded, often unilaterally champaca L.; M. compressa (Maxim.)Sarg.; compound with a row of round ray paren­ M. coriacea Chang & B. L. Chen; M. doltso­ chyma pits subtended by a single elongate pa Buch.-Ham. ex DC.; M. floribunda Finet vessel pit, vessel-parenchyma pit more or & Gagnep.; M. foveolata Merr. ex Dandy; less similar to vessel-ray pits. Helical thick­ M.fulva Chang & B.L. Chen; M. macclurei enings present in the middle portion of vessel Dandy; M. maudiae Dunn; M. mediocris Dan­ elements in M. sinica, one sample of M. con­ dy; M. shiluensis Chun & Y. Wu; M. velu­ i/era (NK s.n.) shows complete fine helical tina DC.; M. wi/sonii Finet & Gagnep.; thickenings throughout the vessel element. Section Anisochlamys - M. hypolampra Thin-walled tyloses usually present. Dandy; Fibres 1300-2080 (530-2980) j.l.m long, Section Dichlamys -M. balansae (A. DC.) F/V ratio 1.44-2.29, thin to medium thick­ Dandy; M. chapensis Dandy; M. martinii (H. walled, with bordered pits of 4-11 j.l.m, main­ Lev.) Finet & Gagnep. ex H. Lev.; ly confined to the radial walls. Thin-walled, Section Micheliopsis - M. figo (Lour.) tyloses present in M. insignis, M. pachycladn Sprengel; M. yunnanensis Franchet ex Finet and M. sinica (NK s.n.), some fibres septate & Gagnep.; (or seemingly so due to appressed tyloses?) Section Tsoongiodendron - M. odara in M. insignis. Trabeculae found in M. coni­ (Chun) Noot. & B.L. Chen; fera. SectionParamichelia-M. baillonii (Pierre) Parenchyma mostly in marginal or zonate Finet & Gagnep. bands of 2-7 cells wide, in addition very rarely also diffuse apotracheal and scanty pa­ Evergreen trees or shrubs from tropical to ratracheal in 2-10 celled strands. subtropical regions, except M. campressa Rays 4-9/mm, 1-2(-3) cells wide, 13- which extends northwards into temperate re­ 23 (7-48) cells or up to 800-3400 J.I.ffi high, gions, at altitudes of 60-2700 m. typically heterocellular with procumbent body Growth rings distinct to faint, marked .by ray cells and 1-4 rows of square to upright marginal or zonate parenchyma. Wood dif­ marginal cells. In juvenile wood square to fuse-porous. Vessels 12-135/mm2, 32-80 upright cells also present among the body ray % solitary, remainder in radial multiples of cells. 2-4 (8), occasionally in oblique to tangential Crystals and oil cells absent. multiples of 2-3, and clusters, usually an-

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Table 4. Variation in selected wood anatomical features of Michelia.

VF % sol TD B DB VEL FL RW H

Section Michelia M. aenea B.L. Chen GS86187 Yunnan, 1790 m * 21 76 60 5 15-20 860 1710 3 + NK 191, Yunnan * 66 35 78 3 26 842 1409 2 M. alba FPC 124 18 65 5 730 1355 2-4 + NK 183, Guangzhou * 63 36 72 5 17 737 1452 2-3 M. cavaleriei NK 184, Guangdong * 70 30 70 3 10-20 820 1410 2 (3) + NK 194, Hunnan 126 20 33 3 14 475 929 2 + FHOw 1831 20 46 110 2 (3) + M. champaca GlF 380 13 57 90 4 20-30 850 1410 2-4 + FHOw 3857 15 59 107 6 670 1460 2-3 + NK 192, Guangdong 26 27 106 4 18 630 1050 2-4 M. compressa FHOw 8212 43 58 64 3 20-25 2-3 + NK 142, Taiwan * 78 20 45 3 20-25 590 1325 2-3 (+) GJF A206 44 71 60 4 10-20 2 (3) + M. coriacea B.L. Chen 87F178, Yunnan, 1450 m 45 52 60 4 10-20 930 1580 2-3 + M. doltsopa FHOw 166 41 60 73 5 15-20 650 2-3 + NK 203, Yunnan * 135 35 45 3 20-25 650 1200 2-3 + M. floribunda B.L. Chen GS86188, Yunnan * 35 80 55 4 20-35 840 1560 2 (3) + NK 102 24 59 76 7 21 931 1752 2-3 + M. foveolata B.L. Chen 87T066, Yunnan, 1585 m * 37 79 60 2 35-40 930 1630 2-3 + GBlF012 52 64 3 766 1450 2-4 + NK 124, Yunnan, 1500 m 38 19 72 4 26 775 1274 2-4 + M. fulva B.L. Chen GS86193, Yunnan 31 69 52 4 24-32 770 1610 2 + M.lacei NK 118 18 58 110 2 35-45 2-3 + M. maclurei GlF600 37 53 65 3 25-50 830 1440 3-4 + NK 163, Guangzi 82 32 75 3 21 644 1202 2-3 + NK 195, Hunan 134 31 56 4 14 632 998 2-3 + M. maudiae GlFOl5 38 63 60 3 10-20 800 1430 2-4 + NK 132, Hunan, 1800 m * 100 55 40 3 15-20 670 1130 2-3 + M. mediocris GlF90679 18 58 80 3 20-25 860 1420 2-3 + NK 202, Hunan 35 40 80 2 20-50 950 1910 2 (3) +

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(Table 4 continued) VF %sol TD B DB VEL FL RW H

M. shiluensis NK 134, Hainan * 40 45 80 3 15-30 780 1360 2 (3) + M. velutina B.L. Chen 87F183, Yunnan 12 67 90 2 24-32 880 1610 2-3 (4) + FHOw955 15 36 128 7 21 931 1752 2-3 + M. wilsonii NK 136, Hunan * 75 30 55 4 10-30 800 1350 2 (3) + NK 178, Sichuan 75 50 50 4 10-35 850 1340 2 + Section Anisochlamys M. hypolampra NK 127, Yunnan, 1800m 20 45 75 0.3 25-50 810 1580 2-3 (4) + Section Dichlamys M. balansae GIF90627 24 69 60 2 10-30 770 1430 2-3 + NK 122, Yunnan, 1500 m * 45 35 60 5 10-15 780 1210 2-4 (5) + M. chapensis RL. Chen 87F186, Yunnan, 1650 m 12 54 75 4 20-30 830 1460 2-4 + NK 193, Yunnan * 56 45 40 2 20-45 2-3 + M. martinii NK 135, Hunan 53 51 75 2 20-35 850 1760 2 + Section Micheliopsis M. Jigo GIF246 59 78 4 730 1250 2-4 + NK 182 88 51 39 4 17 741 1169 2-3 + NK 196, Hunan * 124 28 30 5 11 552 873 2-4 + NK 198, Nanjing * 95 45 45 4 10-25 640 1345 2-3 (4) + M. yunnanensis NK 197, Yunnan, BOOm 70 28 60 4 16-40 850 1540 2 (+/-) Section Tsoongiodendron M.odora NK 166, Guangxi 49 35 70 2 20-30 830 1335 2-3 (4) + Section Paramichelia M. baillonii NK 101, Yunnan, 1800m 20 30 100 4 25-30 950 1780 2-3 (4) (+/-) FHOw2349 17 61 105 2-3 + VF = vessel frequency; %sol = per cent solitary vessel; TD = mean tangential diameter ijun); B = average number of bars per scalariform perforation; DB = distance between bars (~); VEL = mean vessel element length (jJm); FL = mean fibre length ijun); RW = muItiseriate ray width; H = helical thickenings in vessels. * = juvenile wood sample.

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gular, more rarely rounded in cross section, ensis and M. hypolampra are extremely fine mean tangential diameter 30-128 (range 20- and barely visible using phase contrast. 190) 11m, radial diameter up to 60-250 11m, 2. There are no obvious differences in the walls 3-4 (5) 11m thick. Mean vessel element wood anatomy of the sections. In each sec­ length 475-950 (range 350-1490) 11m. Per­ tion there are woods both with and without forations exclusively scalariform in most spe­ oil cells in the rays. cies, with 2-5 (1-10) bars, bars occasionally irregularly branched, distance between bars Discussion 16-30 (10-50) 11m; perforations rarely sim­ ple in M. balansae and M. velutina. Interves­ Classification and identification of Chinese sel pits nonvestured, scalariform or scalari­ Magnoliaceae form-opposite, oblong, 9-45 (5-100) 11m in Although Barkley (1975) questioned the horizontal diameter, with slit-like apertures. inclusion of Liriodendron in the Magnolia­ Vessel-ray pits present in all cross fields, ceae, a recent cladistic analysis of nucleotide mosty large and simple or with reduced bor­ sequences of the plastid gene rbcL of the ders, typically scalariform oval-oblong to Magnoliidae showed that while Liriodendron rounded, frequently unilaterally compound was distinct, there was a well supported rela­ with a row of round ray parenchyma pits sub­ tionship of Liriodendron to other Magnolia­ tended by a single elongate vessel pit, vessel­ ceae (Qiu et aI., in press). Wood of the Lirio­ parenchyma pits more or less similar to ves­ dendroideae is characterised by opposite in­ sel-ray pits. Helical thickenings present tervessel pits and vessel-ray pits, while wood throughout vessel elements in all species, dis­ of the Magnolioideae generally is character­ tinct in most species. Thin-walled tyloses usu­ ised by scalariform and scalariform-opposite ally present. pits. However, this distinction does not Mean fibre length 873-1910 (range 580- always hold because species of Magnolia 2540) 11m long, F/V ratio 1.62-2.20, thin section Rhytidospermum have opposite pits. to medium thick-walled, with bordered pits of Thus, wood anatomically there is overlap 4-8 11m, mainly confined to the radial walls. between Liriodendron and Magnolia. Stark Parenchyma mostly in marginal or zonate (1954) in his key to Magnoliaceae wood bands of 1-8 cells wide, in addition very of the United States, did not distinguish be­ rarely also diffuse apotracheal and scanty pa­ tween Liriodendron and Magnolia fraseri and ratracheal, in 2-6 celled strands. M. tripetala. Rays 4-7/mm, 1-3 (4) cells wide, 14- The genus Manglietia has long been in 23 (3-64) cells or up to 350-1600 11m high, dispute. Bentham & Hooker (1862), Hutchin­ typically heterocellular with procumbent body son (1964), Dandy (1927,1974), Law (1984) ray cells and 1-3 (1--9) rows of square to and many authors recognise this genus; on upright marginal cells. the other hand, Prantl (1891) and some more Oil cells, generally as marginal cells, occur recent authors, reduced this genus to Magno­ in the rays of 15 species (60% of the species lia. Magnolia has two ovules in each ovary; studied); they are common in M. balansae, M. Manglietia usually has more than two. Mang­ chapensis (B.L. Chen 87FI86), M. foveola­ lietia, which is evergreen, and the evergreen ta, M. hypolampra, M. mediocris, M. odora, species of Magnolia range from tropical to M. velutina, rare in M. compressa, M. mar­ subtropical regions. Most species of Mang­ tinii, and intermediate in occurrence in other lietia do not have helical thickenings and species. Oil cells occasional in banded paren­ when they do the thickenings are very faint; chyma in M. chapensis (B.L. Chen 87FI86), oil cells are absent. Evergreen species of M. mediocris, M. hypolampra. Magnolia have helical thickenings and oil Crystals absent. cells. Manglietia has exclusively scalariform Notes: 1. Helical thickenings in vessels perforations, deciduous Magnolia species are well-developed in most species of Mich­ have at least some simple perforations. Thus, elia examined. However, the helical thicken­ although many characters overlap, woods of ings in M. alba, M. champaca, M. yunnan most species of Manglietia can be separated

Downloaded from Brill.com10/06/2021 05:49:47PM via free access Chen, Baas, Wheeler & Wu - Wood anatomy of Magnoliaceae from China 409 from both the evergreen and the deciduous Ecological trends in the Magnoliaceae species of Magnolia. woods from China Wood anatomy of Kmeria is similar to Magnoliaceae from China cover a wide Manglietia, but differs in having crystals, its ecological range, from tropical to temperate rays have more marginal rows than Mang­ areas, from about 18-46° N, and range from lietia, but this may be due to the material of low altitudes to more than 2000 m. They range Kmeria being of young stems. The Magno­ in habit from trees to shrubs. There is con­ liaceae have long been considered difficult siderable variation in length, diameter and taxonomically because they are relatively ho­ frequency of vessel elements. However, it is mogeneous (e.g. Dandy 1974; Nooteboom difficult to assess precisely the relationships 1985), and the wood anatomy is also rela­ between altitude, latitude, habit, and wood tively homogeneous with overlap between anatomy. For example, although Magnolia the genera. A dichotomous key to Chinese cathcartii is confined to the tropical-subtropi­ Magnoliaceae woods follows. cal forests, it usually grows above 2000 m altitude. On the other hand, some species have a wide altitudinal range in the same geo­ Generic wood anatomical key to the graphic area, such as Magnolia championii Magnoliaceae from China which ranges from 500 up to about 2000 m. 1a. Intervessel pits and vessel-ray pits ex­ In this paper, ecological trends can only be clusively or predominantly opposite .. 2 discussed in broad geographic terms because b. Intervessel pits and vessel-ray pits most­ of the lack of reliable and detailed data on ly scalariform, sometimes scalariform- altitude of provenances. opposite, rarely opposite ...... 3 Temperate species have distinct growth 2a. Perforations exclusively or predominant­ rings with growth ring boundaries marked by ly simple both marginal parenchyma and radially flat­ Magnolia hypoleuca, M. officinalis tened fibres; tropical species have indistinct b. Perforations exclusively or predominant- growth rings and marginal and zonate paren­ ly scalariform ...... Liriodendron chyma. This observation agrees with that of 3a. Perforations exclusively or predominant­ Van der Graaff and Baas (1974) for five spe­ ly simple Magnolia (deciduous species), cies of Magnolia. Figures 25-27 (see next sections Oyama, Buergeria, Tulipastrum page) show that vessel elements tend to be lon­ b. Perforations exclusively or predominant­ ger and wider and less frequent in the tropical­ ly scalariform ...... 4 subtropical species (47 wood samples) than 4a. Vessel walls without helical thicken- in the temperate species (12 wood samples). ings ...... 5 Perhaps, the results also are related to habit, b. Vessel walls with helical thickenings . 6 because temperate Magnoliaceae are mostly 5a. Intervessel pits never opposite Manglietia small trees and shrubs while most tropical b. Intervessel pits rarely opposite species of Manglietia and Michelia are trees. Magnolia (evergreen species) Perforation plates tend to be simple in the 6a. Helical thickenings weakly developed 7 temperate species of Chinese Magnoliaceae, b. Helical thickenings well-developed ... 9 except for M. sieboldii; perforations are sca­ 7 a. Rays without oil cells ...... 8 lariform in the tropical-subtropical species. b. Rays with oil cells ..... Michelia (p. p.) Van der Graaff and Baas (1974) found that in 8a. Rays without crystals Manglietia (p. p.) Magnolia (5 species), in contrast to Drypetes, b. Rays with minute, acicular crystals Gordonia, Styrax, and Weinmannia, there Kmeria was a decrease in number of bars with increas­ 9a. Vessel-ray pits usually restricted to the ing latitude or altitude. The present study con­ marginal rows of cells, occasionally uni­ firms Van der Graaff and Baas's observa­ laterally compound ... Magnolia (p. p.) tions for Magnolia. b. Vessel-ray pits throughout the rays, of­ Magnolia can be separated into two groups ten unilaterally compound on wood anatomical characters. The first Michelia (p. p.) group, ranging from 18-30° N, includes all

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% of species the evergreen species which possess exclu­ 30 sively scalariform perforation plates; the sec­ ond group, ranging from 30-c. 40° N, in­ cludes mainly deciduous members with simple 20 perforations; M. cylindrica, M. dawsoniana and M. sieboldii are exceptions as they are deciduous but have scalariform plates. Helical thickenings occur predominantly 10 in evergreen species, and but a few deciduous species. In the dicotyledons as a whole, heli­ cal thickenings are more common in tem­ 0 perate species than in tropical species (Baas 0 0 0 0 0 II) ..... Cl 0 '"6 6 6 6 & Schweingruber 1987; Carlquist 1988; II) ..... 6 '" Cl Wheeler & Baas 1991); Magnolia is an ex­ 25 vessel frequency (mm2) ception to this generalisation. Incidence of helical thickenings is apparently more strong­ ly related to phenology in the Magnoliaceae. % of species 80 Acknowledgments We wish to acknowledge the financial 60 support of the Ministry of Education and Sci­ ence of the Netherlands, which pnabled the first author to study the wood anatomy of 40 Magnoliaceae at the Rijksherbarium/Hortus Botanicus, Leiden. We are very grateful to the following institutes and universities for 20 providing wood samples and help: Zhong­ shan University (Guangzhou); Nankai Uni­ o versity (Tianjing); Guangdong Institute of Forestry (Guangdong); South China Agricul­ tural University (Guangzhou); Southwestern 26 vessel element length (mm) Forestry College (Yunnan); South & Central China Forestry College (Hunan); Yunnan In­ stitute of Forestry (Yunnan); Nanjing Forestry % of species College (Nanjing); Fujian Forestry College 50 (Fujian); Wenshan Forestry Bureau (Anhui) and Yuyuan Forestry Bureau (Guangdong); 40 Rijksherbarium/Hortus Botanicus, Leiden (The Netherlands). We thank Yin-Long Qiu 30 for providing copies of manuscripts on 20 cladistic analyses of the Magnoliaceae and Magnoliidae. 10

o References o o 0 ... Baas, P. & F.H. Schweingruber. 1987. Eco­ 6 '"o

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