IAWA Journal, Vol 14 (4), 1993: 391-412 WOOD ANATOMY OF
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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. Downloaded from Brill.com10/06/2021 05:49:47PM via free access 392 IAWA Journal, Vol. 14 (4),1993 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- Downloaded from Brill.com10/06/2021 05:49:47PM via free access Chen, Baas, Wheeler & Wu - Wood anatomy of Magnoliaceae from China 393 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