IAWA Bulletin n.s., Vol. 11 (4), 1990: 337-378
WOOD ANATOMY OF TREES AND SHRUBS FROM CIllNA II. THEACEAE
by
Deng Liang! and Pieter Baas Rijksherbarium, P.O. Box 9514, 2300 RA Leiden, The Netherlands
Summary The wood anatomy of 95 species belong particularly in the Malesian Archipelago, con ing to fifteen genera of the Theaceae native to tinental Southeast Asia, and the Neotropics. China is described. Despite the wood anatom The delimitation of the family is still a matter ical homogeneity of the family it is possible of debate. For this study we have adopted to key out individual genera (p. 373) as long a concept of Theaceae, comprising the Tern as the unknown material is confined to Chi stroemioideae, Camellioideae (= Theoideae), nese species. In general the wood of Theaceae and Sladenia (Sladeniaceae of some authors). can be characterised by exclusively solitary This delimitation will be argued in a subse vessels, scalariform perforations, opposite to quent paper on the wood anatomy of Thea scalariform vessel wall pitting, ground tissue ceae from outside China (Deng & Baas, in of long fibre-tracheids, parenchyma scanty preparation). In the tropical and subtropical paratracheal and apotracheally diffuse, and southern part of China, the Theaceae are re heterocellular rays. presented by not less than 15 of its c.24 The two subfamilies Camellioideae and genera. The family includes a few economi Ternstroemioideae differ from each other in cally important timber trees (Schima, Adinan type of vessel-ray pits and number of bars. dra, Ternstroemia). The genus Camellia not Sladenia, often treated as a genus of the Thea only includes the tea plant (mainly Camellia ceae, is aberrant in having long vessel mul sinensis) but also many ornamental shrubs tiples and alternate intervessel pits. This in and some oil seed species. dicates an isolated position of the genus. For easy reference the genera of the Thea Within the genus Camellia, of which over ceae occurring in China, their approximate 40 species were studied, 4 wood anatomical total number of species in the world and in groups can be recognised. Only one of these China, and the number of species studied corresponds with a section (Chrysantha) rec wood anatomically are listed in Table 1. In ognised on the basis of macromorphological this table the genera are classified in the same characters. manner as by Keng (1962). Estimated num The taxa from tropical forests tend to have bers of species for China follow the Icono wider and longer vessel members and a graphia Cormophytorum Sinicorum, Sup lower vessel frequency than those from sub plement II (Zhang 1983a) and other Chinese tropical provenances. literature sources (Zhang 1981, 1983b), and Key words: Systematic wood anatomy, identi do not reflect any personal opinion on spe fication, ecological wood anatomy, Thea cific delimitation. Numbers for the whole ceae, Sladenia, Camellia, China. world are taken from Mabberley (1987) and How (1982; between brackets). The tenden cy to adopt very narrow species concepts Introduction among Chinese taxonomists becomes clear The Theaceae are shrubs and small to when the total number of species in Camellia large trees of wide distribution in the tropics, estimated for the whole world (82, in Mab-
1) Permanent address: Department of Biology, Peking University, Beijing, People's Republic of China.
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Table 1. Enumeration of genera of the Theaceae occurring in China.
Subfamily World China Number of species studied Genus Ternstroemioideae Adinandra 70 20 6 Annesfea 4 6 3 Cfeyera 17 6 4 Eurya 70 (130) 80 13 Euryodendron 1 1 1 Sfadenia 1 1 1 1 Ternstroemia 85 (100) 20 4 Camellioideae Apterosperma 1 1 1 Camellia 82 (220) 190 43 Gordonia2 70 7 2 Hartia 4 (14) 13 5 P arapyrenaria 1 (2) 1 1 Pyrenaria3 20 (40) 8 3 Schima 1 (30) 19 6 Stewartia4 6 (11) 6 2
1) Sfadenia is treated in a separate family, the Sladeniaceae, by some authors; in Dilleniaceae, Actinidiaceae or Linaceae by others. 2) Gordonia includes Lapfacea according to Keng (1980). 3) Pyrenaria includes Tutcheria according to Keng (1972). 4) Stewartia = Stuartia. berley 1987) is contrasted with the much Chiang (1962, 1964), Chowdhury (1951) higher estimate of 190 for China alone by Chowdhury & Ghosh (1958), Cutler et af. Zhang (1981). The same discrepancy is found (1987), Dechamps (1985), Desch (1954), for several other genera. Detienne et af. (1982), Detienne & Jacquet A great number of papers on Theaceae (1983), Edlmann & del Monaco (1981), wood anatomy have been published since the Espinoza de Pernia (1987), Furuno (1979), pioneering work by Solereder (1885) and Gasson & Cutler (1990), Ghosh et af. Hitsemann (1886, cited by Solereder 1899). (1963), Van derGraaff & Baas (1974), Herat Solereder (1899 & 1908) and Metcalfe and & Theobald (1977), Ho (1985), Jutte (1959, Chalk (1950) summarised the earlier work, 1964), Kramer (1974), Kribs (1968), Linde and Keng (1962) made a comprehensive man et af. (1963), Luo (1989), Ogata (1975- comparative morphological study of the 1983, 1985), Peraza Oramas & Lopez de Theaceae, including accounts of the wood Roma (1967), Van der Slooten (1968), Sudo anatomy of many genera. Apart from the (1959, 1963, 1988), Tang (1973), Vargas older literature listed by Solereder, Metcalfe (1962), Versteegh (1968), Wong (1975), Wu & Chalk, and Keng, wood anatomical data & Wang (1976), Xie & Mo (1987), Xie et af. on a limited number of Theaceae have been (1987), Xu etal. (1985, 1989), Yamauchi published more recently by Anonymous (1980), Yang & Huang-Yang (1987), Yao (1961), Balan Menon (1955), Baretta-Kuipers (1988), and Ye (1982). Despite these numer (1976), Bascope (1954), Chattaway (1955, ous publications, literature on the wood anat 1956), Cheng et af. (1979, 1980, 1985), omy of Theaceae from China is limited.
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This paper deals with the systematic and and constitutes at the same time a basis for ecological wood anatomy, and microscopic the systematic and ecological sections at the wood identification of the Theaceae native to end of this paper. China, and is the second instalment of the series on the wood anatomy of trees and Growth rings (Figs. 1-6) shrubs from China, a co-operative project Growth rings in the Theaceae from China between the Department of Biology of Peking are distinct or distinct to faint. Specimens University (Beijing) and the Rijksherbarium/ lacking growth rings have only been noted in Hortus Botanicus (Leiden). The first issue Anneslea (Fig. 1). Growth ring boundaries was on Oleaceae (Baas & Zhang 1986). A can be of two types: subsequent paper reviewing the systematic 1) Marked by radially flattened late wood and ecological wood of the Theaceae from fibres, which mayor may not be more thick their entire distribution area is in preparation. walled than the subsequent earlywood fibre tracheids. This type is typical for diffuse Material and Methods porous Theaceae (Figs. 2, 4, 6). Wood samples were collected by the first 2) As 1, but in addition with more or less author and obtained from various institutional marked differences in vessel diameter in wood collections in China. Herbarium vouch early- and latewood (sometimes varying in ers associated with the former were identified different growth rings). This characterises the by Prof. Luo and Prof. Liang. Many of the semi-ring-porous (to diffuse-porous) species other wood samples are unfortunately un (Fig. 3). vouchered, or data referring to the herbarium voucher were not available. As far as possi Vessel distribution and grouping ble this has been remedied by confirming their (Figs. 1-6) identity with matching additional authenti Theaceae are typically diffuse-porous, but cated collections by various Chinese botanists intergradation towards semi-ring-porosity oc (see Acknowledgements). In the nomencla curs in Adinandra, Apterosperma, Camellia, ture we largely follow Iconographia Cormo Gordonia, Hartia, Parapyrenaria, Pyrenaria, phytorum Sinicorum, Supplement II (Zhang and Stewartia (Fig. 3). Semi-ring-porosity is, 1983a) and the study by Keng (1972) on however, never a constant feature at the Pyrenaria. genus level, and often varies even within Light microscopic studies were carried out species or a single specimen from ring to on sections and macerations obtained in the ring. usual manner (cf. Baas & Zhang 1986). Sam Except in Sladenia, Theaceae have mainly ples for scanning electron microscopy were (over 65%) to (almost) exclusively solitary prepared according to Exley et al. (1977). vessels, the remaining vessels being in radial, Conventions for descriptions and determin oblique or tangential pairs or short multiples ing quantitative values are the same as those of at most 3(-4). In Sladenia (Fig. 4) most explained by Baas and Zhang (1986) or were vessels (65%) are in radial multiples of slightly modified to bring them in line with 2-6(-10). recommendations by the IAWA Committee (1989). Vessel frequency and element size Survey of wood anatomical character (Figs. 5 & 6) states in the Theaceae from China Vessel frequency ranges from 27 to 420/ mm 2 in the Theaceae studied; average tangen Preceding the generic descriptions, the tial vessel diameter from 24 to 88 j.lm, and range of wood anatomical character states average vessel member length from 660- within the Theaceae from China will be sur 1890 j.lm. Despite the large ranges of varia veyed and discussed with reference to diag tion of quantitative characters, their diagnos nostic and systematic value. This should fa tic value is limited. Individual species or gen cilitate the interpretation of the descriptions, era of which numerous samples were studied
Downloaded from Brill.com10/11/2021 03:27:50AM via free access 340 IAWA Bulletin n.s., Vol. 11 (4), 1990 sometimes cover a very large portion of the characteristically round to oval and elongate. variation in the family as a whole. For in The apertures are usually slit-like. Average stance, in Eurya, vessel frequency ranges horizontal pit chamber diameter ranges from from 39-201/mm2 (Figs. 5 & 6). Part of 8-22 11m (total range 3-35 11m) and is of the variation recorded is related to ecological very little diagnostic value. factors, another part probably to age and The type of vessel-ray pits is a very impor diameter of the stem samples studied and to tant taxonomic character in Theaceae which the tree or shrub habit of the species. can be used to define the two subfamilies. In Camellioideae vessel-ray pits are large and simple or half-bordered with reduced borders Vessel perforations (Figs. 7-9, 29) to almost simple, and in various arrangements All Theaceae have scalariform perfora from scalariform, opposite or transitional, to tions with on average 8-108 bars per perfo alternate, and occasionally unilaterally com ration (total range 3-139). The lowest num pound (Fig. 13). In Ternstroemioideae, ves bers were found in Pyrenaria (3) and the sel-ray pits are always half-bordered (Fig. highest in Eurya (139). Irregularly forked or 12), usually small, and opposite to alternate. crossed bars commonly occur in Adinandra, Vessel to axial parenchyma pits show the Cleyera, Eurya, Sladenia, and Ternstroemia. same but slightly weaker difference between One or a few elongated bordered pits often the two subfamilies: half-bordered to simple occur next to the opening at one or both ends in Camellioideae and always half-bordered in of the perforation plate, which sometimes Ternstroemioideae. They are mostly arranged makes it difficult to distinguish perforations in a single vertical row, sometimes scalari from scalariform bordered pits (Fig. 9). De form, alternate to opposite (Fig. 20), or even spite fairly large ranges of variation within diffuse, and occasionally unilaterally com specimens, species and genera, the number pound. of bars has some diagnostic and taxonomic value. Most genera of Camellioideae (except Vessel wall thickness and sculpturing Hartia) have much fewer bars per perforation (Figs. 12-17) than the genera of the Ternstroemioideae (ex The vessel walls are always very thin cept Euryodendron and Sladenia). A number (1.5-3 11m). Spiral (helical) thickenings on of genera can also be separated from each the vessel walls are found in most genera ex other using bar number (cf. the key at the end cept in Euryodendron, Parapyrenaria, and of this paper on page 373). The distance be Pyrenaria. In Camellia, Eurya, and Schima tween bars varies greatly and can-in some presence or absence of spiral thickenings cases also be used to separate taxa. varies depending on the species or even spec In many species of Eurya, the perforation imen. However, for most Chinese represen plates may extend from the tip of a vessel tatives of the Theaceae the various expres member nearly to its centre. The end walls of sions of vessel wall sculpturing provide vessel members are oblique with short to highly diagnostic markers. Three character long tails. The genera that have relatively few states are represented: bars (e.g. Parapyrenaria, Pyrenaria) have 1) Spiral thickenings (distinct or faint), shorter tails and less oblique end walls than present throughout the vessel elements (body genera with many bars (e.g. Cleyera, Eurya). and tails) or present in the body only, charac terise Adinandra, Apterosperma, Camellia Vessel wall plttmg (Figs. 10-13, 20, 29) p.p., Cleyera, Eurya p.p., Hartia p.p., and Intervessel pits in the Theaceae are non Sladenia (Fig. 14). vestured and generally scalariform (Fig. 10), 2) Spiral thickenings restricted to the tails scalariform-opposite and opposite to occasio of the vessel elements (Annes lea, Schima, nally alternate. Only in Sladenia intervessel Stewartia, Gordonia, Hartia p. p., Fig. 15). pits are always opposite to alternate in ar 3) Wall thickenings present around pit rangement (Fig. 11), thereby reinforcing the apertures throughout the vessel members isolated position of the genus. Pit shape is (Ternstroemia, Figs. 16 & 17).
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It should be stressed that presence or ab variation within individual genera limit the sence of spiral thickenings sometimes is re diagnostic value of this character in the same lated to ecological factors (cf. Baas 1973; way as is the case for vessel member length. Van der Graaff & Baas 1974). For instance, The ratio of fibre-tracheid length and vessel in Schima superba the tropical specimen member length (F /V ratio) ranges from (Hainan) lacks spirals and the subtropical 1.23-2.50, and is of equally limited diag specimen has distinct spiral thickenings in nostic value in the Theaceae. For instance the vessel element tails. It can be anticipated that genus Camellia covers almost this entire range for some genera of which all Chinese species of variation, and many genera of which have spiral thickenings, species of lower lati several species and many specimens were tudes (e. g. from Indo-China and Malesia) studied cover major parts of it. will lack spirals or have them less prominent ly developed. Axial parenchyma (Fig. 20) The axial parenchyma is generally scanty Tyloses and vessel contents (Fig. 18) paratracheal and apotracheally diffuse or dif Tyloses are ofrare occurrence in the Thea fuse-in-aggregates. Only Euryodendron has ceae from China; they have only been noted in addition vasicentric parenchyma. Most in Adinandra, Apterosperma, Camellia (Fig. commonly strands are 4-6 cells long (total 18), Eurya, Euryodendron, Gordonia, Har range 2-10); this variation appears to be of tia, Parapyrenaria, Schima, and Stewartia, very little or no diagnostic and systematic but not as a constant feature of those genera. value. In most cases the tyloses are thin-walled. In one specimen (Hartia cordifolia) sclerotic ty loses were noted. In accordance with hypoth Ray tissue (Figs. 21-29) eses put forward by Chattaway (1949) and Rays are typically heterocellular. Body ray Bonsen (1990) the tyloses are more common cells of multiseriate (and some uniseriate) in Theaceae wi th large apertures in the vessel rays are procumbent and the ray margins ray /parenchyma pits (i.e. in the Camellioi have 1 to over 4 rows of square to upright deae with reduced pit borders or almost sim marginal cells. In Camellia group I the rays ple pits) than in the Temstroemioideae which have procumbent, square and upright cells predominantly have half-bordered pits with present throughout the rays (Figs. 28 & 29). small apertures on the vessel side. Gums or For most Theaceae from China, Kribs' clas other deposits are typically absent or of very sification (1968) is applicable and has been infrequent occurrence. adopted as a matter of convenience in the de scriptions, with the number of rows of up Fibres (Fig. 19) right to square marginal cells given as well. The fibres are typical fibre-tracheids Kribs' heterogeneous types I, II and III oc (sensu Baas 1986a) and non septate, gener cur. Usually each genus or even species and ally with bordered pits arranged in a single specimen shows a range of two to all inter row or occasionally in two or three rows in grading types (e.g. in Camellia). both the radial and tangential walls. Average Ray frequency ranges from 6-25/mm, pit chamber diameter ranges from 5-8 ~m most commonly 8-13/mm, and is of very (total range 3-11 ~m). limited diagnostic value. Ray size only shows Fibre-tracheid wall thickness varies from a limited variation; width ranges from 1-8 thin to very thick. Most Theaceae have me cells, but is most commonly 1-3 cells. Nar dium thick-walled to thick-walled fibre-tra row 1-2(-3)-seriate rays characterise Adi· cheids. The systematic value at the genus nandra, Camellia group I, Cleyera, and Schi level of fibre-tracheid wall thickness is ex ma p. p. Some species in these genera may tremely limited. have predominantly to almost exclusively Average fibre-tracheid length ranges from uniseriate rays (Fig. 22). Multiseriate rays of 880-2730 ~m (total range 700-3370 ~m). Overlapping ranges between genera and huge (text continued on page 349)
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Legends to Figures 1-35:
Figs. 1-4. TS, x 32. - 1. Anneslea rubiflora, wood diffuse-porous with indistinct ring boun dary. - 2. Schima argentea, wood diffuse-porous with distinct growth ring boundary. - 3. Stewartia sinensis, wood semi-ring-porous. - 4. Sladenia celastrifolia, most vessels in long ra dial multiples.
Figs. 5 & 6. TS, x 32, ecological trends in Eurya. - 5. Tropical E. trichocarpa with wide ves sels in relatively low frequency. - 6. Subtropical E. macartneyi with narrow vessels in high frequency. - Figs. 7-9. Radial split surfaces (SEM) showing perforation plates and wall or namentation. - 7. Eurya macartneyi, bars branched and anastomosing, x 170. - 8. Apterosper ma oblata, less than 20 bars and spiral (helical) thickenings, x 340. - 9. Cleyera japonica, per foration plates with 40 or more bars, faint spiral thickenings, x 255. - Figs. 10 & 11. TLS, x 320. - 10. Schima superba, scalariform intervessel pits. - 11. Sladenia celastrifolia, opposite to alternate intervessel pits.
Figs 12 & 13, RLS, x 320. - 12. Anneslea rubiflora, vessel-ray pits half-bordered. - 13. Ste wartia sinensis, vessel-ray pits with much reduced borders to almost simple, note also unilater ally compound pits on the left and in bottom of photograph. - Fig. 14. Apterosperma oblata, spiral thickenings throughout vessel member, SEM, x 460. - Fig. 15. Hartia naiyuangensis, spiral thickenings restricted to tails of vessel members, SEM, x 310. - Figs. 16 & 17. Tern stroemia gymnanthera, vessel wall thickenings restricted around pit apertures; 16. TLS, x 80; 17. Radial surface, SEM, x 1900. - Fig. 18. Camellia sinensis, vessel with thin-walled tyloses (arrow), SEM, x 265. - Fig. 19. Schima wallichii, distinctly bordered fibre-tracheid pits, ra dial surface, SEM, x 475.
Fig. 20. Anneslea rubiflora, tangential surface showing parenchyma strand, and fibre-tracheids with distinctly bordered pits, SEM, x 225. - Figs. 21-25. TLS, x 80. - 21. Anneslea rub i flora, rays of two distinct sizes. - 22. Schima argentea, almost all rays uniseriate. - 23. Para pyrenarea multisepala, rays 1-5-seriate. - 24. Camellia chrysantha, rays with alternating uniseriate and equally narrow biseriate portions. - 25. Camellia oleifera, rays with enlarged crystalliferous cells.
Figs 26-29. Ray composition, RLS, x 80. - 26. Pyrenaria microcarpa, rays heterogeneous II III, with one or two rows of marginal square to upright cells. - 27. Camellia chrysantha, ray heterogeneous I, with 2-6 rows of square to upright marginal cells. - 28. ibid., ray with alter nating portions of procumbent and square to upright cells. - 29. Camellia sinensis, ibid., note also scalariform perforations and large vessel-ray pits.
Figs. 30-35. Crystals. - 30. Sladenia celastrifolia, long chains of chambered crystals in axial parenchyma, TLS, x 80. - 31. Camelliafurfuracea, crystals in enlarged ray cells, RLS, x 80.- 32. Camellia oleifera, integumented crystal in enlarged ray cell, SEM, x 600. - 33. Camellia gigantocarpa, non-integumented crystal in enlarged ray cell, SEM, x 405. - 34. Camellia ter minalis, nonintegumented crystals in normal ray cells, SEM, x 685. - 35. Camellia spec., in tegumented crystal in normal ray cell, SEM, x 1200.
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5-8 cells wide are found only in Anneslea, Generic wood anatomical descriptions Euryodendron, and Ternstroemia. Rays are of two distinct sizes in Anneslea (Fig. 21), Subfamily: Ternstroemioideae Eurya p.p., Euryodendron, and Ternstroe mia. In some cases it is difficult to decide Adinandra Jack (Table 2) whether the size classes are distinct or wheth Material studied: A. bockiana Pritzl ex er the intermediate sizes are sufficiently num Diels, Mt. Leigong in Guizhou: Guizhou erous to conclude in favour of absence of this Wood Company w 114 - A. bockiana var. feature; e.g. Parapyrenaria (Fig. 23). In Ca acutifolia (Hand.-Mazz.) Kobuski, Guang mellia group I (sect. Chrysantha) rays with dong: Guangdong Institute of Forestry w 749 multi seriate portions as wide as the uniseriate and w 78 - A. glischroloma Hand.-Mazz., portions occur (Fig. 24). Ray height shows Guangdong: Guangdong Institute of Forestry very wide ranges of variation. In genera with w 307 - A. hainanensis Hayata, Hainan: rays of two distinct sizes average multi seriate Guangdong Institute of Forestry w 76; Gu ray height is usually greater (0.22-4.0 mm) angdong: Guangdong Institute of Forestry than in genera with intergrading ray sizes w 572 - A. mil/ettii (Hook. et Am.) Benth. (0.06-1.4 mm; mostly 0.2-0.5 mm). Per et Hook. f., Guangdong: Guangdong Insti forated ray cells are of rare occurrence in the tute of Forestry w 216, w 1797; Anhui: An family, and have only been noted in Aptero hui Agricultural College w 3061; Guangxi: sperma, and Camellia p.p. Sheath cells have Liuzhou Wood Company s.n.; Fujian: Fujian been noted in Eurya p. p., Euryodendron, Forestry College s. n. - A. nitida Merr. ex Stewartia p.p., Ternstroemia p.p. but are Li, Guangxi: Liuzhou Wood Company w 77; never very distinct. In all taxa studied the ray Guangxi Forestry College s. n. parenchyma cells are moderatel y thick -walled, Evergeen trees from tropical or subtropical densely pitted, with interparenchyma pits of forests. ten weakly bordered, and the cell walls often Growth rings faint to distinct, marked by disjunctive. flattened and thick-walled latewood fibre-tra cheids. Wood diffuse-porous or semi-ring Crystals (Figs. 25, 30-35) porous to diffuse-porous. Vessels 91-2941 Crystals are of rare occurrence in the Thea mm 2, 63-93% solitary, remainder in radial ceae; they have only been noted in Camellia, and tangential to oblique multiples of 2-3, Schima, Sladenia, and Stewartia, but mostly angular in cross section, tangential diameter not as a constant feature at the genus level. 35-55 (20-70) 11m, radial diameter up to When present there is usually one large pri 110 11m, walls I-211m thick. Vessel member matic crystal per cell or chamber. Crystals in length 1130-1890 (670-2320) 11m. Perfora chambered axial parenchyma cells occur in tions scalariform with 31-48 (21-67) bars, Schima, Sladenia (Fig. 30), and Stewartia. bars sometimes irregularly forked or crossed, The chambers are sometimes enlarged. The distance between bars 3-4 (2-5) 11m. Inter crystalliferous chains are 4-12 (2-51) vessel pits nonvestured, scalariform and op chambers long. In Schima wallichii some posite to alternate, oval to elongate, 5-29 11m chambers have additional vertical division in horizontal diameter, with slit-like aper walls. Chambered crystalliferous ray paren tures. Vessel-ray pits half-borded, opposite chyma cells are limited to some specimens of and alternate, occasionally scalariform in A. Group II of Camellia (Fig. 31). Crystals in hainanensis (w 76); vessel-parenchyma usu normal upright and/or square ray cells occur ally in a single vertical row. Spiral thicken in Group I of Camellia; crystals in normal ings present throughout body of vessel ele procumbent ray cells occur in Group IV of ments, distance between coils 2-7 11m. Ty the same genus. It is common to find crystals loses usually absent, but present in A. bocki enclosed in a sclerotic membrane Cintegu ana (w 749), A. hainanensis (w 76) and A. mented crystals ') in Camellia p. p., Schima, mil/ettii (w 1797, and Guangxi s.n.). Sladenia, and Stewartia (Figs. 25, 31, 32 and Fibre-tracheids 1860-2690 (1520-2840) 35). 11m long, FIV ratio 1.25-1.69, medium to
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Table 2. Wood anatomical diversity in Adinandra.
E ~ E .6 til o;l .6 8 .!01 ~ ...... oj ~ :a ~ c ~ '" 4-< c 3' a '[ 0 ~ '0 til ~ '0 a til til U til til 4-< ... a.l 1a --- "Cl a.l ...... ;> .0 1a :g .~ 0 4-< .0 u .s:: ... c ~ o;l 0 ~a.l a.l ;>, •.::l c C .0 u til ... a.l a c --- c ] tl a.l a a.l '0 a.l '0 ~ ::l til 0.0 ~ til C ::l til a ~ til ~ 0' a.l ::l ] ] a.l .0 ;> § c ;> t.::l .s--- a.l 0 a.l a.l a.l a.l "Cl J: 0.0 0.0 u u 0.0 0.0 '0 oj c § 0:1 .~ til ~ z:l ~ ... til .~ a.l ~ til tl a.l ;>, a.l '0 ;> Material ;> til 0:1 ~ :a :a ~ ~ ~ A. bockiana 235 90 35 36 2-4 4-6 1130 1860 1-2 A. bockiana var. acutifolia w749 294 63 35 41 2-4 4-5 1340 2130 1(-2) w78 197 85 45 48 2-4 4-6 1360 2100 1(-2) A. glischroloma 243 90 35 43 2-4 4-5 1440 2240 1(-2) A. hainanensis w572 151 91 50 33 3-5 4-6 1310 2170 1-2 w76 107 87 55 43 2-5 3-6 1890 2690 1-3 A. millettii w216 164 85 45 32 3-4 4-6 1320 2240 1-3 w 1797 235 72 40 39 2-4 4-5 1550 1940 1(-2) Guangxi s. n. 123 82 45 40 3-4 4-6 1400 2330 1-2(-3) Fujian s.n. 178 89 40 39 3-4 4-6 1610 2390 1-2 w 3061 160 93 40 37 2-4 2-4 1490 2480 1(-2) A. nitida Guangxi s.n. 200 84 40 39 3-5 5-7 1400 2150 1-2(-3) w77 91 92 50 31 3-5 4-6 1270 2130 1-2(-3)
thick-walled, with distinctly bordered pits of 2. Adinandra hainanensis (w 76, Hainan) 6-8 ~m, numerous in radial and tangential has the longest vessel members and fibre-tra walls. cheids, the widest vessels and lowest vessel Parenchyma scanty para tracheal and diffuse frequency, and thereby follows a general apotracheal, in 4-6 (2-8)-celled strands. trend for tropical Theaceae. Rays 10-13 (9-18)/mm, 1-3 cells wide, 3. The wood structure of Adinandra is 0.20-0.49 (0.14-0.82) mm high, hetero very homogeneous, the wood anatomical geneous type I to III, with 1-6(-8) rows of variation in quantitative and some qualitative square to upright marginal cells or with pro characters within the genus is recorded in cumbent, square and upright cells mixed Table 2. throughout the ray. Crystals absent. Anneslea Wall. (Figs. 1, 12,20; Table 3) Notes: 1. Herat & Theobald (1977) noted Material studied: A.fragrans Wall., Xi homogeneous-heterogeneous rays in A.lasio mong in Yunnan: Southwest Forestry College petala from Sri Lanka. In our material all rays w 538; Guangdong: South China Agricultural observed were heterogeneous. University s. n.; Mt. Darning in Guangxi:
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Table 3. Wood anatomical diversity in Anneslea. E ~ E ~ ::1. 8 ..c:: ~ 0; biJ ..c:: N""' ;e \: biJ 8 -0 ~ '";>.. 8 ~ .§ 0; .... '" a'" :g .... .. .... .0 ] ~ ~ e;; 0 § ~ 'i=: .--.. ;>.. ~ .J:) .... \: U 8 ~ "';::::: <1'1 8 > .. .~ > ~ ~ ;e'" ~ 0; ~5 A. fragrans '" w 538 41 90 55 41 2-4 1180 2110 1-2 3-6 Guangdong s. n. 69 94 65 33 3-4 1510 2580 1(-2) 3-5 Guangxi s.n. 117 80 45 36 2-5 1130 2070 1 3-5 A. hainanensis w9 129 90 55 39 2-5 1640 2580 1-2 3-6 w276 82 90 60 39 3-5 1550 2130 1-2 3-5 A. rubiflora 101 76 65 55 2-4 1710 2340 1-2 3-5
Guangxi Forestry College s. n. - A. hainan walled, with distinctly bordered pits of 4-7 ensis (Kobuski) Hu, Hainan: Guangdong In ~m, numerous in radial and tangential walls. stitute of Forestry w 9, w 276 - A. rubiflora Parenchyma scanty paratracheal and apo Hu et Chang, Guangdong: Guangdong Insti tracheal diffuse, in 4-6 (2-7)-celled strands. tute of Forestry w 77. Rays 11-14/mm, of two distinct sizes, Evergreen trees from tropical seasonal rain 1 (-2)-seriate and 3-6 (2-8)-seriate, up to 3 forests to subtropical forests. mm high, heterogeneous type II to III. Growth rings absent or faint and marked Crystals absent. by weak changes in fibre-tracheid wall thick Notes: 1. Keng (1962) recorded rays of ness. Wood diffuse-porous. Vessels 41-129/ type heterogeneous I in A. fragans var. lan mm 2,76-94% solitary, remainder in tan ceolata, but in our material of A.fragrans gential, or radial to oblique pairs, angular in rays are heterogeneous II to III. cross section, tangential diameter 45-65 2. Anneslea rubiflora has the highest num (35-90) ~m, radial diameter up to 140 ~m, ber of bars (Table 3). It remains to be tested walls 1.5-2 ~m thick. Vessel member length whether this feature can be used for identifi 1130-1710 (760-2180) ~m. Perforations cation of this species. In other respects the scalariform with 33-55 (21-71) bars, dis genus Anneslea is wood anatomically very tance between bars 2-4 ~m. Intervessel pits homogeneous. non vestured, scalariform, scalariform-oppo site, or opposite to alternate, oval to elongate, 4-17 ~m in horizontal diameter, with slit-like Cleyera Thunb. (Fig. 9; Table 4) apertures. Vessel-ray and vessel-parenchyma Material studied: C. incornuta Y. C. Wu, pits opposite to alternate, half-bordered and Mt. Darning in Guangxi: Guangxi Forestry smaller than intervessel pits. Spiral thicken College s.n. - C. japonica Thunb., Guang ings only present in vessel element tails. dong: Guangdong Institute of Forestry w 82, Fibre-tracheids 2070-2580 (1770-2980) w 51-C. obovata H. T. Chang, Guangdong: ~m long, F/V ratio 1.37-1.83, thin- to thick- Guangdong Institute of Forestry w 127 -
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Table 4. Wood anatomical diversity in Cleyera.
~ 6' '-'~ 6' '" ~ :::t E ..c '-' ,--. oj til ..c ,--. N '8 I c bh 8 .....'" c .:!l "0 0 2 "0 8 '" ~ ... 2 u ... '"0 til '" ..8 "d ..... 0 ,--. '" .D '"til '0 .~ 0 8 > ...... D u 8 ..c tf. 0 0 u I:i >. '-' ca c c oj .D u .~ 0 8 c c ] ~ 0 8 0 "0'" 0 "0 l? ::l ::l b.I) l l C 8 '" ] '-' 0' '"0 c ::l '"0 0 '"> £3 c ..8 ..8 > t+= -S
C. pachyphy/la Chun ex H. T. Chang, Guang tance between coils 8-16 (2-30) Ilm. Thick dong: Guangdong Institute of Forestry w 74; walled tyloses present in C. obovata. Xingning in Hunan: Deng s.n. - Cleyera Fibre-tracheids 1620-2380 (990- 2880) spec., Guangxi: Guangxi Wood Company Ilm long, FIV ratio 1.32-1.73, thick-walled, s. n. with distinctly bordered pits of 5-7 Ilm, Evergeen trees from subtropical mountain numerous in radial and tangential walls. forests (alt. 800-1200 m). Parenchyma scanty paratracheal and apo Growth rings distinct, marked by flatten tracheal diffuse, in 4-5 (3-7)-celled strands. ed and thick-walled latewood fibre-tracheids. Rays 11-20/mm, 1-2(-3) cells wide, 6- Wood diffuse-porous. Vessels 177-258/ 13 (2-30) or up to 0.5 mm high, hetero mm 2, 70-96% solitary, remainder in radial geneous type I to III, with 1-3(-9) rows of and tangential to oblique multiples of 2-3 square to upright marginal cell. (-4), angular in cross section, tangential di Crystals absent. ameter 35-40 (25-60) Ilm, radial diameter Notes: 1. Cleyera pachyphylla has the low up to 85 Ilm, walls 1.5-2.5 Ilm thick. Ves est value with vessel member length and tra sel member length 940-1580 (560-2130) cheid length, and can be separated from other Ilm. Perforations scalariform with 33-42 species studied. In other aspects the genus (20-68) bars, distance between bars 3-4 is wood anatomically quite homogeneous Ilm. Intervessel pits nonvestured, scalariform (Table 4). to opposite-alternate, oval to elongate, 4-12 2. In its general wood anatomy the genus Ilm in diameter, with slit-like apertures. Ves Cleyera is very close to Adinandra. Only the sel-ray and vessel- parenchyma pits opposite distance between the coils of the spiral thick to alternate or diffuse, mostly half-bordered, enings on the vessel walls between two gen occasionally with reduced borders and almost era is different. simple. Spiral thickenings present throughout 3. The specimen of Cleyera spec. (Guangxi body of vessel members, fine to coarse, dis- Wood Company s.n.) was originally receiv-
Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. Baas - Wood anatomy of Theaceae from China 353 ed as a Camellia. Wood anatomically it is opposite to alternate, occasionally unilaterally completely out of place in this genus, and a compound. Spiral thickenings absent in E. perfect fit with Cleyera, and we re-identified ciliata and E. trichocarpa, present in vessel it accordingly. element tails in some specimens and through out the vessel elements in others (see Table Eurya Thunb. (Figs. 5, 6, 7; Table 5) 5). Tyloses mostly absent, noted in E. brevi Material studied: E. acuminatissima Merr. styla and E. hebeclados only. et Chun., Xingning in Hunan: Deng s.n. - Fibre-tracheids 1590-2310 (1330-2760) E. alata Kobuski, Guilin in Guangxi: Guang j.Lm long, FlY ratio 1.23-2.09, thin- to thick xi Forestry College s.n.; Longling County in walled, usually medium thick-walled, with Yunnan: Southwest Forestry College w 359 distinctly bordered pits of 5-7 j.Lm, numerous - E. brevistyla Kobuski, Hunan: Deng 74 - in radial and tangential walls. E. ciliata Merr., Hainan: Guangdong Institute Parenchyma scanty paratracheal, diffuse of Forestry w 158 - E. hebeclados Ling, Gu apotracheal, and diffuse-in-aggregates, in 4-6 angdong Institute of Forestry w 1656, w 603; (3-7)-celled strands. Hunan: Deng 73 - E. impressinervis Kobus Rays 13-21 (l0-25)/mm, of two dis ki, Guangdong: Guangdong Institute of For tinct sizes or all ray sizes intergrading: 1(-2) estry w 158 - E. japonica Thunb., Anhui, seriate and (2-)3-4-seriate, 0.35-1.24 Anhui Agricultural College w 1380 - E. mac (0.13-1.76) mm high, heterogeneous type I artneyi Champ., Guangdong: Guangdong to II (-III), with 1-7 (1-18) rows of square Institute of Forestry w 623 - E. mega to upright marginal ray cells. Weakly differ trichocarpa Chang, Guangdong: South China entiated sheath cells occasionally present. Agricultural College s. n. - E. muricata Dunn, Crystals absent. Hunan: Deng 2, 64, 65, 71, 74, 75 - E. niti Notes: 1. The material studied included a da Korth., Fujian: Fujian Forestry College specimen labelled Eurya groffii Merr. (Guang s.n.; Lipo in Guizhou: Guizhou Wood Com dong Institute of Forestry w 501). This sam pany w 279; Guangdong: Guangdong Insti ple is aberrant in Eurya because it has a signi tute of Forestry w 730; Hainan: Guangdong ficantly lower number of bars per perforation Institute of Forestry w 230 - E. trichocarpa (average 34) and vessel wall thickenings re Korth., Ximeng in Yunnan: Southwest For stricted to the pit aperture regions as in Tern estry College w 528 - E. tsaii H. T. Chang, stroemia. We have provisionally reidentified Lushui in Yunnan: Yunnan Wood Company this sample as a Ternstroemia. w 59. 2. Spiral thickenings are absent in the trop Evergreen small trees and shrubs from ical species (E. ciliata and E. trichocarpa) and tropical and subtropical forests. thereby follow a general ecological trend. Growth rings distinct, marked by flatten Despite this, expression of helical sculpture is ed and thick-walled fibre-tracheids in the late fairly diagnostic at the species level within wood. Wood diffuse-porous. Vessels 39- Eurya. 20l/mm2, 80-98% solitary, remainder in tangential, radial to oblique multiples of2-3, Euryodendron H. T. Chang angular in cross section, tangential diameter Material studied: E. excelsum H.T. Chang, 30-75 (20-100) j.Lm, radial diameter up to Guangdong: Zeng & Huang 12305. 160 j.Lm, walls 1-2 j.Lm thick. Vessel member Evergeen trees from subtropical forests. length 830-1690 (640-2180) j.Lm. Perfora Growth rings faint, marked by weakly tions scalariforrn with 51-108 (36-139) bars, flattened latewood fibre-tracheids. Wood dif bars sometimes forked and anastomosing fuse-porous. Vessels 43/mm2, 90% solitary, (Fig. 7), distance between bars 1-5 j.Lm. remainder in tangential, or radial to oblique Intervessel pits nonvestured, scalariforrn, pairs, angular to rounded, tangential diameter scalariforrn-opposite and opposite to alter 80 (55 -110) j.Lm, radial diameter up to 155 nate, oval to elongate, 5-27 j.Lm in horizontal j.Lm, walls 1.5-2.5 j.Lm thick. Vessel member diameter, with slit-like apertures. Vessel-ray length 1370 (880-1880) j.Lm. Perforations and vessel-parenchyma pits half-bordered, scalariforrn with 20 (12-34) bars, distance
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Table 5. Wood anatomical diversity in Eurya.
1: ~ 0/) 8' ~ 0-::s '" -6 £ ... C ~ 0/) '""' £
between bars 6-11 Jlm. Intervessel pits non pits, but half-bordered. Spiral thickenings ab vestured, scalariform, scalariform-opposite sent. Thin-walled tyloses occasionally pres to alternate, 5-12 Jlm in horizontal diameter, ent. with slit-like apertures. Vessel-ray and ves Fibre-tracheids, 2650 (2320-3170) Jlm sel-parenchyma pits similar to intervessel long, FIV ratio 1.94, thin- to thick-walled,
Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. Baas - Wood anatomy of Theaceae from China 355 with distinctly bordered pits of 6-8 J.1m, with distinctly bordered pits of 6-7 J.1m, numerous in radial and tangential walls. numerous in radial and tangential walls. Parenchyma scanty paratracheal to vasi Parenchyma diffuse-in-aggregates and centric, and apotracheal diffuse to diffuse-in scanty paratracheal, in 4-6 (3-7)-celled aggregates, in 3-9 (2-1O)-celled strands. strands. Rays 12 (9-15)/mm, oftwo distinct sizes, Rays 11-12/mm, 1-3(-4) cells wide, up 1-2-seriate and (3-)4-6-seriate, 1-2-seri to 1.4 mm high, heterogeneous type I to II, ate rays 0.48 (0.22-1.18) mm high; multi with 3-6(-10) rows of square to upright seriate rays 0.89 (0.48-1.85) mm high, het marginal cells. erogeneous type II to III, with 1-5(-9) rows Crystals solitary prismatic and integument of square to upright marginal cells; weakly ed in chambered axial parenchyma; chambers differentiated sheath cells common. slightly enlarged, in chains of 4-10(-18). Crystals absent. Notes: 1. Keng (1962) described the wo'od Note: The wood anatomy of Euryodendron structure of Sladenia based on juvenile wood. is here described for the first time. Zhang Quantitative values in Keng's material are (Chang) published this genus in 1963 and much smaller (e.g. vessel frequency 95- suggested that it is close to Eurya. Wood 134/mm2, vessel diameter 32-49 J.1m, ves anatomically Euryodendron can easily be sel member length 560-780 J.1m, fibre-tra separated from Eurya by its much lower bar cheid length 820-1040 J.1m) than that in our number, wider bar spacing, and wider multi material. Keng's account of vessel grouping: seriate rays. Although these differences may "solitary or 2-5 in short tangential chains" be trivial in other plant groups, in Theaceae may be a misprint for 'radial' chains. Luo's they constitute a large enough discontinuity account of Sladenia wood anatomy (1989, to support the separate generic status. Wood see especially his plate 93) based on two ma anatomically Euryodendron is at least as ture wood specimens is in full agreement close to Ternstroemia as to Eurya. with our observations. 2. The anatomical description of mature wood of Sladenia provides new information Sladenia Kurz (Figs. 4, II, 30) about the taxonomic position of this genus. Material studied: S. celastrifolia Kurz, Xi See also the separate discussion on the posi shuang Banna in Yunnan: Deng s.n. tion of Sladenia. on page 371. Evergreen trees from tropical seasonal rain forests. Ternstroemia Mutis ex L. f. (Figs. 16 & 17; Growth rings faint, marked by flattened Table 6) latewood fibre-tracheids. Wood diffuse-por Material studied: T. gymnanthera (Wight ous. Vessels 38/mm2, mostly in radial mul et Am.) Sprague, Guangdong: Guangdong tiples of 2-6(-10), remaining 35% solitary, Institute of Forestry w 612, w 1664; South angular in cross section, tangential diameter China Agricultural College s.n.; Hainan: Gu 65 (50-100) J.1m, radial diameter up to 120 angdong Institute of Forestry w 342, w 353, J.1m, walls 2-2.5 J.1m thick. Vessel member w 286; Rongshui in Guangxi: Liuzhou Wood length 1540 (880-2210) J.1m. Perforations Company s. n.; Guangxi, Guangxi Forestry scalariform with 17 (11-26) bars, occasion College s.n.; Fujian: Fujian Forestry College ally irregularly branched, distance between s. n.; Jiangxi: Anhui Agricultural College w bars 4-9 J.1m. Intervessel pits nonvestured, 1653 - T. gymnanthera Sprague var. wightii opposite to alternate or diffuse, 6-9 Jlm in (Choisy) Hand.-Mazz., Guangdong: Guang diameter, with slit-like apertures. Vessel-ray dong Institute of Forestry w 42, w 65 - T. and vessel-parenchyma pits similar to inter macrophylla S. Y.Liang, Mt. Damao in Gu vessel pits, but half-bordered and slightly angxi: Liuzhou Wood Company' s.n. - T. smaller. Spiral thickenings weakly develop nitida Merr., Rongshui in Guangxi: Liuzhou ed. Tyloses absent. Wood Company s.n. - Ternstroemia spec., Fibre-tracheids 2380 (1930-2760) J.1m Guangdong: Guangdong Institute of Forestry long, F/V ratio l.54, medium thick-walled, w 501.
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Table 6. Wood anatomical diversity in Ternstroemia.
6' ~ ~ 6' ..c ~ ~ 00 ..c "'" ;a c 00 '"8 ~ C til ~ ~ ;;." 8 til til '-' .... ~ C Evergreen trees from tropical and subtrop seriate and half-bordered. Spiral thickenings ical forests. usually restricted to vessel element tails, but Growth rings faint to distinct, marked by local thickenings mostly present around pit radially flattened late wood fibre-tracheids. apertures throughout vessel elements (Figs. Wood diffuse-porous. Vessels 93-240/mm2, 16 & 17), or even extending to form irregular 68-95% solitary, remainder in tangential spirals (in T. microphylla). Tyloses absent. and radial to oblique multiples of 2-3, tan Fibre-tracheids 1740-2520 (1380-2820) gential diameter 35-65 (30-85) J.lm, radial J.lm long, F/V ratio 1.53-2.01, very thick diameter up to 125 J.lm, walls 1.5-2 J.lm walled, with distinctly bordered pits of 5-7 thick. Vessel member length 910-1420 11m, numerous in radial and tangential walls. (660-1710) J.lm. Perforations scalariform Parenchyma scanty paratracheal and dif with 24-38 (18-54) bars, distance between fuse apotracheal, in 4-6 (2-8)-ceUed strands. bars 2-6 11m. Intervessel pits nonvestured, Rays 8-15/mm, of two distinct sizes, scalariform, opposite to alternate, 6-8 (3- 1(-2)-seriate and 3-6-seriate, 2-4 mm high, 20) 11m in horizontal diameter, with slit-like heterogeneous type II (I-III), with 1-6(-12) apertures. Vessel-ray and vessel-parenchy rows of square to upright marginal cells. ma pits usually opposite to alternate or uni- Weakly differentiated sheath cells common. Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. Baas - Wood anatomy of Theaceae from China 357 Crystals absent. scalariform or scalariform-opposite, oval to Notes: 1. Yang & Huang-Yang (1987) elongate, 12-15 (7-30) 11m in horizontal noted vestured pits in Ternstroemia. In our diameter, with slit-like apertures. Vessel-ray light microscopic and SEM studies vestured pits large and simple, scalariform or scalari intervessel pits were not observed. There are form-opposite, or even diffuse, occasionally also no other records of vestured pits in the unilaterally compound. Vessel-parenchyma family, and we suspect that the above obser pits round to oval, half-bordered to simple, vation is erroneous or based on soluble in usually in a single vertical row. Fine spiral crustations on the pit membranes which often thickenings present throughout vessel ele create the false impression of vesturing in a ments, distance between coils 3-5 11m. Ty number of woods. loses thick-walled, present in a few vessel 2. Herat and Theobald (1977) reported elements. homogeneous rays in T. emarginata from Sri Fibre-tracheids 1980 (1710-2210) 11m Lanka. However, their illustrations show long, FIV ratio 1.78, medium thick to very heterogeneous rays, as found in all material thick-walled, with distinctly bordered pits of studied from China. 4-5 (3-6) 11m, numerous in radial and tan 3. Some of the wood anatomical variation gential walls. in Ternstroemia seems associated with eco Parenchyma scanty paratracheal and dif logical factors. The three tropical specimens fuse apotracheal, in 4-6-celled strands. (Hainan) have relatively long vessel members Rays 14/mm, 1-2(-4) cells wide, 8-14 and the lowest vessel frequencies (Table 6). 0-18) cells or up to 0.22 (0.09-0.38) mm 4. Ternstroemia stands out within the Thea high, heterogeneous type II to III, with 1-3 ceae from China on account of its local wall (-5) rows of square to upright marginal thickenings around vessel pit apertures, cells, occasionally with sheath cells. which sometimes extend to form irregular Crystals absent. spirals. Notes: 1. Xie and Mo (1987) studied the 5. Chattaway (1955) noted crystals in ray same sample of A. oblata. Their description cells in three species of Ternstroemia outside largely agrees with the above, but they erron China, indicating that the total range of wood eously referred to the intervessel pits as op anatomical variation in the genus is larger posite. than covered here. 2. Zhang (Chang) first described Aptero sperma in 1976 and suggested a close rela Subfamily Camellioideae tionship with Schima. Wood anatomically the genera are slightly different, viz. in vessel Apterosperma H. T. Chang (Figs. 8, 14) wall sculpturing and some quantitative val Material studied: A. oblata H. T. Chang, ues, e.g. bar number. However, these minor Guiping County in Guangxi: Guangxi For differences do not preclude close phylo estry College s. n. genetic affinity. Small evergeen trees from subtropical for ests at 300-500 m altitude. Camellia L. emend. Sweet (Figs. 18,24, Growth rings distinct, marked by radially 25,27-29,31-35; Table 7) flattened and thick-walled fibre-tracheids in Material studied (listed according to wood latewood. Wood diffuse-porous to weakly anatomical groups recognised during this semi-ring-porous (varying in different rings). study): Vessels 282/mm2, 80% solitary, remainder in tangential and radial to oblique pairs, an Group I gular in cross section, tangential diameter 30 Camellia chrysantha (Hu) Tuyama, Fang (25-45) 11m, radial diameter up to 75 11m, cheng in Guangxi: Guangxi Institute of For walls 1.5-2 11m thick. Vessel member length estry s.n. - C. euphlebia Merr. ex Sealy, 1110 (840-1330) 11m. Perforations scalari Dongxin in Guangxi: Guangxi Institute of form with 2307-31) bars, distance between Forestry s.n. - C. impressinervis Chang et bars 3-7 11m. Intervessel pits nonvestured, s. Y. Liang, Longzhou in Guangxi: Guangxi Downloaded from Brill.com10/11/2021 03:27:50AM via free access 358 IAWA Bulletin n.s., Vol. 11 (4), 1990 Institute of Forestry s. n. - C. longgangensis - Camellia spec., Xiashi in Guangxi: Guang C.F. LiangetS.L. Mo, Longzhou in Guang xi Institute of Forestry s.n. - Camellia spec., xi: Guangxi Institute of Forestry s. n. - C. Longzhou in Guangxi: Guangxi Institute of longgangensis var. grandis C.F. Liang et Forestry s. n. - Camellia spec., Anhui: Anhui S.L. Mo, Ningming in Guangxi: Guangxi Agricultural College w 3891. Institute of Forestry s.n. - C. longruiensis S. Y. Liang et C.Z. Deng, Longrui in Guang Group III xi: Guangxi Institute of Forestry s. n. - C. Camellia acutissima Chang, Hunan: Deng longzhouensis J. Y. Luo, Longzhou in Gu s.n. - C. caudata Wall., Yunnan: Yunnan angxi: Guangxi Institute of Forestry s.n. - Wood Company w 439 - C. cordifolia (Metc.) C. multipetala S.Y. Liang et C.Z. Deng, Nakai, Guangxi: Guangxi Institute of Fores Fangcheng in Guangxi: Guangxi Institute of try s.n. - C. cuspidata Wright, Hunan: Forestry s. n. - C. pubipetala Y. Wan et S. Z. Deng 13, 72, 81; Guangxi: Guangxi Wood Huang, Dongxing in Guangxi: Guangxi In Company s.n. - C. dehongensis Chang, stitute of Forestry s.n. - C. terminalis S. Y. Yunnan: Deng s.n. - C. handelii Sealy, Liang et Z.M. Su, Fangcheng in Guangxi: Hunan: Deng s.n. - C. irrawadiensis Barua, Guangxi Institute of Forestry s.n. - C. wu Yunnan: Deng s. n. - C. rhytidocarpa Chang, mingensis S. Y. LiangetC.R. Fu, Wumingin Guangxi: Guangxi Forestry College s. n. - Guangxi, Guangxi Institute of Forestry s. n. C. rosthorniana Hand.-Mazz., Hunan: Deng s. n. - C. sinensis O. Ktze, Guangxi: Gu Group II angxi Wood Company s.n. and Deng s.n.; Camellia brevistyla (Hay.) Coho St., Fujian: Yunnan: Deng s.n.; Jiangxi: Deng s.n.; Fujian Forestry College s.n. - C. chekiango Hunan: Deng 16,90 (root- and stemwood), laeosa Hu, Hunan: Deng s. n. - C.furfuracea 199, 200, 201, 205, 210, 213, 215, 218, (Merr.) Coho St., Guangdong: Guangdong 219, 222, and 227; Zhejiang: Deng 198, Institute of Forestry w 547 - C. furfuracea 207, 211, 212, and 216; Anhui: Anhui Agri var. lutea Hu, Guangxi: Guangxi Forestry cultural College 209, 226; Guangxi: Deng College s.n. - C. gigantocarpa Hu et T.C. 109 (root and stemwood); Jiangxi: Deng 110 Huang, Hunan: Deng s. n. - C. grijsii Hance, - C. sinensis var. assamica Kit., Yunnan: Hunan: Deng 108 - C. japonica L., Zhejiang: Deng s.n.; Hainan: Guangdong Institute of Deng s.n. - C. kissi Wall., Hunan: Deng Forestry w 294; Hunan: Deng s.n. - C. sub s. n. - C. lapidea Hu, Guangdong: Guang acutissima Chang, Hunan: Deng s.n. - C. dong Institute of Forestry s.n. - C. longi taliensis Melch., Yunnan: Deng s.n. caudata Chang, Hunan: Deng s.n. - C. mi crophylla (Merr.) Chien, Hunan: Deng s.n. Group IV - C. oleifera Abel, Guangdong: Guangdong Camellia spec., Kunrning in Yunnan: Yun Institute of Forestry w 796; Jianghua in Hu nan Institute of Forestry w 155. nan: Deng s.n.; Daoxiang in Hunan: Deng s. n.; Fujian: Fujian Forestry College s.n.; Evergreen shrubs or small trees from Anhui: Anhui Agricultural College s.n. - C. tropical to subtropical regions; some species, pitardii Coho St., Sichuang: Sichuang Agri especially widely cultivated for tea or oil cultural College w 8459; Hunan, Deng s. n. seeds, or as ornamental. - C. polyodonta How ex Hu, Guilin in Gu Growth rings always distinct, marked by angxi: Deng s.n.; Wangian in Guangxi: Deng flattened latewood fibres. In some species s.n. and Guangxi Forestry College s.n. - C. marked by differences in vessel diameter. reticulata Lindl., Hongkong: KJw 1347 - C. Wood diffuse-porous in some species, dif saluenensis Stapf ex Bean, Yunnan: Deng fuse- to semi-ring-porous in others. Vessels s.n. - C. semiserrata Chi, Guangdong, Gu 113-454/mm2,70-97% solitary, remain angdong Institute of Forestry w 771 - C. der in radial and oblique to tangential multi vietnamensis T. C. Huang ex Hu, Guang ples of 2-3(-4), tangential diameter 30-50 dong: Guangdong Institute of Forestry w 491, (20-65) 11m, radial diameter up to 95 11m, w 671 -C. villosa Chang, Hunan: Deng s.n. walls 1-2.5 11m thick. Vessel member length Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. Baas - Wood anatomy of Theaceae from China 359 660-1400 (440-1740) j.l.m. Perforations sca tion with presence or absence of spiral thick lariform with 11-42 (8-55) bars, distance enings of vessel, and ray type within Camellia between bars 6-9 (5-12) j.l.m. Intervessel seems to be of taxonomic interest. As appar pits nonvestured, scalariform to opposite ent from the descriptions and Table 7, the scalariform, in some species intervessel pits four groups of Camellia can be characterised scalariform to alternate, oval to elongate, 9- by the following features: 16 (3-25) j.l.m in horizontal diameter, with slit-like apertures. Vessel-ray pits simple or I. Crystals prismatic to elongate in 'normal' half-bordered with much reduced borders, upright and square ray parenchyma cells. large, usually scalariform, sometimes uni Spiral vessel wall thickenings usually laterally compound. Vessel-parenchyma pits absent. Rays 1-2(-3)-seriate with alter half-bordered to almost simple, uniseriate or nating uniseriate and narrow 2-3-seri,ate diffuse. Spiral thickenings absent in Group I; portions which are equally wide. weak spiral thickening present in some spe II. Crystals prismatic to elongate, usually in cies of Group II & III and in Group IV (see tegumented in weakly to distinctly en Table 7). Tyloses only noted in low frequen larged and often sclerified idioblastic ray cies in C. acutissima, C. brevistyla, C. cordi cells, which are often chambered. Spiral folia, C .furfuracea p. p., and a few specimens thickenings usually present. Rays 1-3 of C. sinensis. (-5)-seriate with multiseriate portions Fibre-tracheids 1020-2560 (700-2870) virtually always wider than the uniseriate j.l.m long, FIV ratio 1.24-2.50, medium portions. thick to very thick-walled, in most species III. Crystals absent. Faint spiral thickening thick-walled, with distinctly bordered pits of present in some species. Rays 1-3(-4) 5-7 (4-8) j.l.m, numerous in radial and tan seriate with multi seriate portions mostly gential walls. wider than uniseriate portions, only in Parenchyma scanty paratracheal and dif some species (or specimens) with equal fuse apotracheal, in some species also dif ly wide multi seriate and uniseriate por fuse-in-aggregates, in 3-6 (2-7)-celled tions (Table 7). strands. IV. Prismatic crystals integumented, in 'nor Rays 1O-21/mm, 1-3(-5) cells wide, mal' procumbent, upright and square ray 0.31-0.55 (0.21-0.74) mm high, hetero cells. Spiral thickenings absent. Rays 1- geneous type I to III, with 1-5(-7) rows of 2(-3)-seriate with multi seriate portions upright and square marginal cells or in some wider than uniseriate portions. species with procumbent, square and upright cells mixed throughout the rays. In Group I 2. Sealy (1958) divided Camellia into 12 usually 1-2(-3)-seriate with alternating por sections. Zhang (1981) recognised not less tions of uniseriate and narrow 2-3-seriate than 19 sections. Wood anatomically only portions (Fig. 24). section Chrysantha can easily be separated Crystals absent in Group III; prismatic to and coincides with our Group I. Species of elongate in other groups; present in non Group II belong to sections Oleijera, Furfur chambered 'normal' upright and square ray acea, Paracamellia, Tuberculata, and Camel parenchyma cells in Group I; in Group II lia of subgenus Camellia. Species of Group usually integumented and present in enlarged III are from section Thea (subgenus Thea) and often sclerified idioblastic ray cells, and sections Theopsis and Camelliopsis of which may be chambered in C. furfuracea subgenus Metacamellia. The single species of var. lutea and Camellia spec. (w 3891); in 'Group' IV cannot be assigned to a section Group IV, integumented crystals present in because it remained unidentified. normal ray cells. Usually there is only one 3. Several specimens received as Eurya or crystal per ray cell; two or three crystals per even Prunus (Rosaceae) [in Table 7: Camellia cell were found in some species of Group I. spec. (from Anhui) and Camellia spec. (from Notes: I. The diversity in presence or ab Yunnan)] were assigned to Camellia on the sence of crystals, type of crystals in associa- basis of their wood anatomy onlv. Downloaded from Brill.com10/11/2021 03:27:50AM via free access 360 IAWA Bulletin n.s., Vol. 11 (4), 1990 Table 7. Selected wood anatomical characters of Camellia. *ell ~ :;a ,* 11)... 6 ""..c::<"l_ 6 ~ 6 ~ ell ..c:: ell 6 ~~ ~«:I ell ~ bi> 11) ..c:: ell ;a I: ell ..!.o; ell > bi> "" ",;::l 11) ~ 11) I: N ~ ell ~ 1:0' "" > ell ... ~ U «:I 11) ell -5 .... 6 11) ~ ~ .... .0 I: '0 0 ~o > 6 .... 0 '" «:I ell E- ~ 11) ell ..c:: ... . .: I: 0; 0 U >. ;g 'l:j bJl ~ ~.g U ell 6 ... I: I: I: 1'; 6 .~ t: 11) .... ~ ~ '2 ;::l I: 0 0 ell 11) ;::l ~ ell 6 ;::lo.. 0' 11) 11) ;::l .!o. 11) bJl § U > I: t+=: -5 I: «:I * . ~ ' 5 C Material > > ell > U > 0.. ~ '" '" '" ...'" "'ell Group I C. chrysantha 150 91 40 1250 21 2220 1-2(-3) + n C. euphlebia 167 82 40 1400 28 2460 1-2(-3) + n C. impressinervis 186 90 30 950 30 2220 1-3 + n C. longgangensis 177 87 30 910 21 1530 1-2(-3) + n var. grandis 160 96 35 1190 20 1590 1-2(-3) + n C. longruiensis 147 92 35 1200 23 2190 1-2(-3) + nc C. long zhouensis 176 93 30 1020 18 1820 1-2(-3) + n C. multipetala 220 82 35 750 21 1540 1-2(-3) + n C. pubipetala 224 90 35 1200 22 1710 1-2(-3) + n C. terminalis 160 94 30 920 20 1450 1-3 + n C. wumingensis 187 91 30 850 19 1490 1-3 + n Group II C. brevistyla 190 90 40 1231 15 ± 2086 1-5 ei C. chekiangolaeosa 124 85 40 850 14 + 2070 1-3 ei C. fur/uracea 250 88 35 1020 15 1880 1-3 ei var.lutea 153 88 35 820 13 1820 1-3 eci C. gigantocarpa 165 77 45 820 12 ± 1600 1-5(-6) e(i) C. grijsii 175 80 35 890 14 2070 1-3 ei C. japonica 373 86 30 660 14 + 1420 1-2 ± ei C. kissi 155 96 30 850 19 + 1860 1-3 ei C. lapidea 117 84 40 1070 16 ± 2560 1-3 ei C. longicaudata 900 18 ± 1890 1-3 ei C. microphyl/a 280 85 35 ± 1-3(-4) ei C. oleifera Guangdong s.n. 172 77 45 1080 17 + 2290 1-3 ei Jianghua s. n. 191 90 35 910 16 + 1930 1-3 ei Daoxiang s. n. 144 87 35 920 17 ± 1890 1-3 ei Anhui S.n. 160 95 30 980 15 + 1960 1-3 ± ei C. pitardii 170 90 40 780 15 + 1950 1-3 ei C. polyodonta Wangtian s. n. 166 84 40 800 15 ± 1890 1-3 e(i) Guangxi 235 89 30 920 15 + 2010 1-3 e(i) Guilin s.n. 210 93 35 750 15 + 1830 1-3 ± e(i) C. eticulata 187 87 30 930 15 + 2090 1-3 ei Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. Baas - Wood anatomy of Theaceae from China 361 (Table 7 continued) *~ ~ Downloaded from Brill.com10/11/2021 03:27:50AM via free access 362 IAWA Bulletin n.s., Vol. 11 (4). 1990 (Table 7 continued) * j -5 0.0 H * * + = presence, - = absence. ** - = absent; n = in nonnal ray cells; e = in enlarged ray cells ('idioblasts'); c = in chambered ray cells; i = integumented. *** rootwood. Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. Baas - Wood anatomy of Theaceae from China 363 4. Two rootwood samples (Deng 90 and compound. Vessel-parenchyma pits half 109) of C. sinensis were studied; they had bordered to almost simple, round to oval longer, wider and more thin-walled vessel elongate, usually in a single vertical row. members, longer fibre-tracheids, and larger Spiral thickenings absent. Tyloses absent in ray cells and total ray volume, while vessel most samples, but thin-walled tyloses present frequency was lower than in the stem wood in w 669 of G. axillaris. of the same plants (Table 7), thereby follow Fibre-tracheids 1650-2680 (1380-3370) ing general differences between the wood 11m long, F/Vratio 1.29-1.95, medium thick anatomy of stems and subterranean organs. to thick-walled, with distinctly bordered pits Rays with alternating uniseriate and bi- or of 5-8 11m, numerous in radial and tangential triseriate portions of equal width are much walls. more common in the rootwood. The root Parenchyma very scanty paratracheal, and wood of C. cuspidata and C. japonica de diffuse apotracheal in 3-5-celled strands. scribed and pictured by Cutler et al. (1987) is Rays 9-15/mm, 1-3 cells wide in G. axil very similar to our rootwood samples of laris, 1-4(-5) cells wide in G. hainanensis, C. sinensis. and 0.29-0.48 (0.06-0.98) mm high. Het erogeneous type (I-)II-III, with 1-3(-5) Gordonia Ellis (Table 8) rows of square to upright marginal cells. Material studied: G. axillaris D. Dietr., Hai Crystals absent. nan: Guangdong Institute of Forestry w 699 Notes: 1. In G. hainanensis the rays are & w 285; Hainan: South China Agricultural usually wider [1-3(-4)-seriate] and have University s. n.; Muchuang County in Sichu more (up to 5-6) rows of square to upright ang: Sichuang Agriculture College w 8248; marginal cells than in G. axillaris [1-2(-3) Jingxiu County in Guangxi: Liuzhou Wood seriate with 1-3(-4) rows of square to up Company s. n.; Rongshui County in Guang right marginal cells]. xi: Guangxi Forestry College s.n. - G. hai 2. In G. axillaris the tropical specimens nanensis Chang, Hainan: Guangdong Insti (w 285, w 669, Guangdong s.n.) have rela tute of Forestry w 669 and w 718. tively high values for vessel diameter and low Evergreen trees from tropical or subtropi vessel frequencies compared with the sub cal forests. tropical specimens. Growth rings distinct to faint, marked by 3. Chattaway (1955, 1956) recorded crys radially flattened and thick-walled latewood tals in axial parenchyma of Gordonia; in our fibre-tracheids and sometimes also by differ materials they were absent. ences in vessel diameter. Wood diffuse-por ous to semi-ring-porous (varying in different Hartia Dunn (Fig. 15; Table 9) rings or samples of the same species). Ves Material studied: H. cordifolia Li, Ping sels 96-213/mm2, 79-94% solitary, re tang County in Guizhou: Guizhou Wood mainder in tangential and radial to oblique Company s.n. - H. kwangtungensis Chun, multiples of 2-3, angular in cross section, Guangdong: Guangdong Institute of Forestry tangential diameter 35-50 (25-70) 11m, ra w 292; Rongshui County in Guangxi: Liuzhou dial diameter up to 95 11m, walls 1.5-2 11m Wood Company s.n. - H. naiyuangensis thick. Vessel member length 1010-1750 Hu, Guangdong: Guangdong Institute of (580-2240) 11m. Perforations scalariform Forestry s. n. - H. sinensis Dunn, Longlin with 14-25 (11-39) bars, bars occasionally County in Yunnan: Southwest Forestry Col irregularly branched, distance between bars lege w 392; Mt. Daming in Guangxi: Guang 5 -7 (3-10) 11m. Intervessel pits nonvestured, xi Forestry College s.n. - H. yunnanensis scalariform or scalariform-opposite, oval to Hu, Jinping County in Yunnan: Southwest elongate, 10-19 (8-30) 11m in horizontal Forestry College s. n. diameter, with slit-like apertures. Vessel-ray Evergreen trees from subtropical forests. pits small to large, simple to half-bordered, Growth rings distinct, marked by flattened scalariform or scalariform-opposite, oppo and thick-walled latewood fibre-tracheids and site, or even diffuse, occasionally unilaterally differences in vessel diameter. Wood semi- Downloaded from Brill.com10/11/2021 03:27:50AM via free access 364 IAWA Bulletin n.s., Vol. 11 (4),1990 Table 8. Selected wood anatomical characters of Gordonia. 6' ~ -.::, 6' ..c -.::, 8o;j """' bh -5 ';;) :.s c bI) '"8 j:l c d:l <1.l d:l 8 % .... ~ u .... '" ~ "Cl '"<1.l .D '"~ E '03 0 > .D 8 ..c ""'.... 8 ~ til 0 <1.l u >-. '--' ""' C o;j u '0 .... <1.l a c C <1.l b <1.l d:l'" <1.l E d:l ~::: ::: bI) 8 ~ '" c ::: <1.l '"<1.l E- O' '"<1.l .D '" ~ <1.l > S C > ~ -5 ring-porous or diffuse-porous. Vessels 75- Fibre-tracheids 2050-2740 (1660-3090) 160/mm 2, 80-97% solitary, remainder in 11m long, F/Vratio 1.39-1.91, medium thick tangential and radial to oblique pairs, tangen to thick-walled, rarely thin-walled, with dis tial diameter 40-55 (30-70) 11m, radial di tinctly bordered pits of 6-8 11m, numerous in ameter up to 95 11m, walls 1.5-2 11m thick. radial and tangential walls. Vessel member length 1160-1720 (680- Parenchyma scanty paratracheal and apo 2150) 11m. Perforations scalariform with 42- tracheally diffuse or diffuse-in-aggregates, in 57 (30-70) bars, distance between bars 3-5 3-5 (2-7)-celled strands. (2-6) 11m. Intervessel pits nonvestured, sca Rays 8-13/mrn, 1-3(-5) cells wide, lariform or scalariform-opposite, oval to elon 0.30-0.54 (0.08-0.91) mrn high, hetero gate, 5-24 11m in horizontal diameter, with geneous type I-lI(-III), with 1-4(-6) rows slit-like apertures. Vessel-ray pits simple and of square to upright marginal cells. large, scalariform to scalariform-opposite, Crystals absent. occasionally unilaterally compound. Vessel Notes: 1. Hartia has been treated as a sec parenchyma pits round to oval, simple or tion of Stewartia by several authors following half-bordered with much reduced borders Airy Shaw (1936). Yan (1981) and Ye (1982) usually in a single vertical row, occasionally excluded the section from Stewartia, as did unilaterally compound. Spiral thickenings Merrill (1938). The wood anatomy supports usually present in vessel element tails only, the latter view. See also separate discussion but in H. naiyuangensis also present in body on the relationships of Hartia and Stewartia. of some of the vessel elements. Tyloses usu 2. All specimens of Hartia have long sca ally absent, infrequent sclerotic tyloses noted lariform perforation plates with more than 40 in H. cordifolia and thin-walled tyloses in H. bars, thereby exceeding the other genera in sinensis. the Camellioideae. Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. Baas - Wood anatomy of Theaceae from China 365 Table 9. Wood anatomical diversity in Hartia. 8' E, N"" ... "" 8 '" ~c: ~ "0 i::.8 "0 8 ~ ... ~ ..!:l () '"~ 8 ..... ~ '"0 .8 '";> .D '"~ ] 0 ,3- ...... D 8 [8 ..c: oJ 0 0 ",- () ~ ;... ~ 'p c: bIl O .D () ... 0 8 c: ~ g .~ 0 , 8 c: "0'" c:"" .8 "0 ::l ::l 1;'08 8 ~° ~ ;> ° '" ::l 0 '" ..1(- ] 5 '"0 §E, .D () ::l t;:l ir ;> c: r[ .~ 0 £ * ... 0 0 0'-" 0 ..J:: ;... - () £..c: '1:l .q 1;'0~ bIl bIl..c: bIl ~ ro c: ro_ ro .~ 0 "0 oJ~ '" .~ ... n ~bIl ...0 ... '" 0 ;> ;... 0 '"0 "0- ~ ~ ;> '" c: ;> p. ;> ~:.a ro :.a ro..!:l 'a~ ro ~ Material '" '" - H. cordifolia SD 105 84 50 46 3-5 1490 2050 1-4 H. kwangtungensis w 291 SD 105 81 50 44 2-4 1160 2070 1-4 Guangxi s. n. SD 78 80 50 42 2-4 1170 2230 1-4(-5) H. naiyuangensis SD 75 97 45 46 3-5 1270 + 2150 1-3 H. sinensis w 392 D 160 85 40 47 3-5 1650 2050 1-3 Guangxi s. n. D 84 89 55 49 3-5 1720 2640 1-3 H. yunnanensis D 130 91 40 71 3-4 1620 2740 1-3 * D = diffuse-porous; SD = semi-ring-porous. 3. The diversity in vessel distribution, ex tance between bars 5-12 J..lm. Intervessel pits tent of spiral thickenings, and ray width can nonvestured, scalariforrn to scalariforrn-op perhaps be used to separate species within posite, oval to elongate, 10-15 (7-25) J..lm in Hartia (see Table 9). Studies of more sam horizontal diameter, with slit-like apertures. ples are, however, required to test the diag Vessel-ray pits large and elongate to round, nostic value of these characters at the species scalariforrn, scalariforrn-opposite or diffuse, level. simple to half-bordered with reduced bor ders, sometimes unilaterally compound. Ves Parapyrenaria H. T. Chang (Fig. 23) sel-parenchyma pits half-bordered to almost Material studied: P. muitisepaia (Merr. et simple, in single vertical rows or diffuse. Spi Chun) Chang, Hainan: South China Agricul ral thickenings absent. Medium thick-walled tural University s.n. tyloses only present in a few vessel elements. Evergreen trees in tropical seasonal rain Fibre-tracheids 1790 (1460-2240) J..lm forests at 600-900 m altitude. long, F/V ratio 1.66, thin- to thick-walled, Growth rings faint to distinct, marked by usually medium thick-walled, with distinctly radially flattened latewood fibres and differ bordered pits of 6-9 J..lm, numerous in radial ences in vessel diameter. Wood diffuse- to and tangential walls. weakly semi-ring-porous, varying in differ Parenchyma scanty paratracheal and apo ent rings. Vessels 91/mm2, 85% solitary, tracheally diffuse to diffuse-in-aggregates, in remainder in radial and oblique to tangential 4-6 (3-7)-celled strands. pairs, angular in cross section, tangential di Rays 9 (7-12)/mm, 1-5 cells wide, 0.3 ameter 55 (30-70) J..lm, radial diameter up to (0.1-1) mm high, heterogeneous type II to 95 J..lm, walls 1.5-2 J..lm thick. Vessel mem III, with 1-2(-3) rows of square to upright ber length 1180 (840-1460) J..lm. Perfora marginal cells. tions scalariforrn with 8 (5 -16) bars, dis- Crystals absent. Downloaded from Brill.com10/11/2021 03:27:50AM via free access 366 IAWA Bulletin n.s., Vol. 11 (4), 1990 Table 10. Wood anatomical diversity in Pyrenaria. E E 2> ::i. .c: '--" ;;' bo .c: 8 c:: bo .0 0) 0 ~ E .... 0) & .... .0 0) .c: .... ~ 0 () 0) o () ;>, '--" 'pca .... c:: () :J 0) 0) .0 '--" ":::a ;> ...... c:: .0 ;> <.C1 o E 0) 0) 0) 0) -5 ~ O)~ () "0 0l)0) Ol) Ol) Ol) [):c '0 OIl c:: OIl .~ .oOl) , Eh:: 0) 0 ;> OIl :Jo.. ;> Note: According to Zhang (1963b), Parapy Growth rings faint to distinct, marked by renaria is a close relative of Pyrenaria. Wood differences in vessel diameter and fibre wall anatomically the two genera are indeed very thickness. Wood diffuse-porous to semi-ring similar; in our material studied there is only a porous, varying in different rings. Vessels small difference in ray width (l-5-seriate in 49-277 Imm2, 70-89% solitary, remainder Parapyrenaria; 1-4-seriate in Pyrenanaria); in radial, tangential and oblique multiples of this difference could easily break down when 2-3, tangential diameter 30-65 (20-100) more material of both genera is studied. Ilm, radial diameter up to 150 Ilm, walls 1-3 Ilm thick. Vessel member length 730-1740 (450-2040) Ilm. Perforations scalariform Pyrenaria Blume (Fig. 26; Table 10) with 7-17 (2-25) bars, distance between Material studied: P. championi (Nakai) H. bars 7-15 (3-18) Ilm. Intervessel pits non Keng, Longlin County in Yunnan: Southwest vestured, scalariform or scalariform-oppo Forestry College w 375; Guangdong: Guang site, oval to elongate, 7-22 (4-35) Ilm in dong Institute of Forestry w 331, w 367, w horizontal diameter, with slit-like apertures. 607; Fujian Forestry College s.n.; Guangxi: Vessel-ray pits large and simple, scalariform Liuzhou Wood Company s.n.; Rongshui in or scalariform-opposite, occasionally unila Guangxi: Guangxi Forestry College s.n. - terally compound and more or less reticulate. P. mengiongensis Tao G. T., Xishuang Ban Vessel-parenchyma pits similar to intervessel na in Yunnan: Deng s.n. - P. microcarpa pits, but half-bordered or with reduced bor (Dunn.) H. Keng, Guangdong: Guangdong ders, and smaller. Spiral thickenings absent. Institute of Forestry w 175 and w 710. Tyloses absent. Evergreen trees from tropical and SUbtrop Fibre-tracheids 890-2440 (79 -2760) Ilm ical fores ts. long, FIV ratio 1.22-2.24, medium thick- Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. Baas - Wood anatomy of Theaceae from China 367 walled, with distinctly bordered pits of 6-11 Evergeen trees from tropical and subtropi 11m, numerous in radial and tangential walls. cal forests. Parenchyma abundant apotracheally dif Growth rings faint to distinct, marked by fuse-in-aggregates and diffuse, and scanty radially flattened and thick-walled latewood paratracheal, in 3-6 (2-7)-celled strands. fibre-tracheids. Vessels diffuse, 27-166/ Rays 8-13/mm, 1-4 cells wide, 0.22- mm2, 68-95% solitary, remainder in radial 0.45 (0.03-0.8) mm high, heterogeneous to tangential multiples of 2-3(-5), weakly type I to II, with 1-3(-6) rows of square to angular or angular in cross section, tangential upright marginal cells. diameter 40-90 (30-110) 11m, radial diame Crystals absent. ter up to 190 11m, walls 1.5-2 11m thick. Ves Notes: 1. Pyrenaria is treated here in the sel member length 940-1820 (770-2240) delimitation of Keng (1972), who combined 11m. Perforations scalariform with 9-18 (5- Tutcheria (in our material P. championi and 21) bars, distance between bars 5-15 11m. P. microphylla) with Pyrenaria. The merger Intervessel pits nonvestured, scalariform to can be supported by the shared wood ana scalariform-opposite, oval to elongate, 8-29 tomical features of the three species studied; Jlffi in horizontal diameter, with slit-like aper the lower vessel frequency of Pyrenaria s. s. tures. Vessel-ray pits large and simple to half (P . meng/ongensis, Table 10) has no taxo bordered with reduced borders, mostly sca nomic significance in this respect. lariform, occasionally sCalariform-opposite or 2. In P. microcarpa there are usually few even diffuse, sometimes unilaterally com er [1-2(-4)] rows of square to upright mar pound. Vessel-parenchyma pits simple to ginal cells than in the other species. half-bordered, in a single vertical row or oc casionally diffuse. Spiral thickenings present Schima Reinw. ex Blume (Figs. 2, 10, 19, in vessel element tails (except in one speci 22; Table 11) men of S. superba, w 8). Thin-walled tyloses Material studied: S. argentea E. Pritz. ex present in some specimens of S. argentea and Diels, Xinning County, Hunan: Deng s.n.; S. wallichii. Rongshui County in Guangxi: Liuzhou Wood Fibre-tracheids 1580-2590 (1520-3260) Company s.n.; Guangdong: South China Ilm long, F/V ratio 1.32-1.78, thin- to thick Agricultural University; Lushui County in walled, with distinctly bordered pits of 5-10 Yunnan, Yunnan Wood Company w 20; Ilm, numerous in radial and tangential walls. Lipo County in Guizhou: Guizhou Wood Parenchyma scanty paratracheal and apo Company s.n.; Guangxi: Deng s.n.; Hunan: tracheal diffuse and diffuse-in-aggregates, in Deng s. n. - S. bambusi/olia Hu, Shangsi 4-6 (2-1O)-celled strands. County in Guangxi: Guangxi Wood Company Rays 10-13 (9-18)/mm, 1-2(-3) cells w 472. - S. crenata Korth., Yilian County in wide, 0.2-0.49 (0.14-0.82) mm high, het Yunnan: Southwest Forestry College w 425. erogeneous type II to III, with 1-3(-6) rows - S. noronhae Reinw. ex Blume, Longlian of square to upright marginal cells. County in Yunnan: Southwest Forestry Col Crystals absent in S. crenata, present in lege w 385 - S. superba Gardn. et Champ., the other species, but not always as a con Lipo County in Guizhou: Guizhou Wood stant feature (see Table 11). Crystals integu Company w 282; Longling County in Yun mented and prismatic in chambered axial pa nan: Southwest Forestry College w 393; Mu renchyma cells; chambers enlarged in some chuang County in Sichuang: Sichuang Agri species, sometimes weakly sclerified, number cultural College w 8250; Hainan: Guangdong of chambers per chain 4-10 (2-51); some Institute of Forestry w 8; Fujian: Fujian For chambers with vertical division walls in S. estry College s. n.; Anhui: Anhui Agricultural wallichii. College w 1287 - S. wallichii Choisy, Nan Notes: 1. Keng (1978) reduced all Malayan ning in Guangxi: Guangxi Forestry College species to S. wallichii. Mabberley (1987) con s.n.; Nanning in Guangxi: Guangxi Forestry sidered this variable species to comprise the College S.n.; Baishe County in Guangxi: entire genus. For practical reasons we still Guangxi Wood Company w 80126. have adopted narrow species concepts. The Downloaded from Brill.com10/11/2021 03:27:50AM via free access 368 IAWA Bulletin n.s., Vol. 11 (4), 1990 Table 11. Wood anatomical diversity in Schima 6' $; ~ 6' OJ ..t:: ~ r-. ~ to r-. ~ en a:l '" ~ til a a:l ~ a:l til a en en ~ .... ] u 0 .... en til :> 0 ta 1l .... 0 :> .c ta ] 0 ~ 8- .... .c a ..t:: .... 0 ~ 0 0 0 til '-' OJ ~ u :>. t * c = chambered axial parenchyma cells; e = crystalliferous chambers enlarged; i = integu- mented. wood anatomical variation can partly be used This is probably a manifestation of the gen- in support; e. g., S. wallichii sensu stricto eral ecological trend for spiral thickenings to can be separated from the other species by be less common in the tropics. its low vessel frequency, wider vessels, and wider rays (Table 11); the species moreover Stewartia L. (Figs. 3 & 13; Table 12)) stands out on account of the infrequent, yet Material studied: S. rubiginosa Chang, constant occurrence of vertical anticlinal divi- Xingning in Hunan: Deng s.n. - S. sinensis sion walls in some of the crystal chambers. Rehd. et Wils., Anhui, Anhui Agricultural 2. In Schima, only one specimen (w 8 of College 1480, s.n.; Mt. Darning in Guangxi: S. superba) from tropical origin (Hainan) Guangxi Forestry College; Guangdong: Gu- lacks spiral thickenings on the vessel walls. angdong Institute of Forestry s. n. Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. Baas - Wood anatomy of Theaceae from China 369 Table 12. Wood anatomical diversity in Stewartia. ';i;' 0 ~ la", "0 .... u :::= '" "d .... 0'0 '" '"ta 1l .Q3 0 "'u ~ .0 8 ..c ...... 0; ~ 0; 0 0 Or:: >.. '-' .p r:: g .0 "'.~ u 0 8 p r:: r:: 0 8 ~Oll 0 "0'" '"la "0 ::: ota ~a .0 ~ ....~ ,-.. ::: '" ~ -8 .::8 0' 0 1a~ .... '" 0 '"> ...... 0 .0 ~~ @~ -5 0- * .. '" 0 '"0 ~ ::: > 0 > r:: ::: 0 0. > 1;;;0 r:: ;0 oj- ~ oj~ ~ r:: .... t Material '" S. rubiginosa SD 36 83 65 14 5-12 1100 1760 1-4 1-7 S. sinensis w 1437 SD 41 94 65 13 4-8 1000 + 2090 1-3(-4) 1-6 + Anh. s.n. SD 90 89 60 19 3-6 1090 1880 1-3(-4) 1-4 Gngd. s.n. D 35 93 100 13 7-19 1380 + 2640 1-3(-4) 1-5 + w 139 D 40 80 85 14 4-12 1220 + 2420 1-3(-4) 1-5 + * SD = semi-ring-porous; D = diffuse-porous. Deciduous trees from subtropical forests. Parenchyma abundant, scanty paratracheal Growth rings faint to distinct, marked by and apotracheally diffuse to diffuse-in-aggre radially flattened and thick-walled latewood gates, in 4-6 (2-1O)-celled strands. Rays fibre-tracheids. Wood diffuse-porous to weak- 8-15/mm, 1-4 cells wide, 0.34-0.47 1y semi-ring-porous. Vessels 35-105/mm2, (0.12-1.04) rom high, heterogeneous type 1- 80-95% solitary, remainder in radial or tan II(-III), with 1-3(-6) rows of square to gential to oblique multiples of 2-3, weakly upright marginal cells. angular to rounded in cross section. tangen Crystals absent from S. rubiginosa and one tial diameter 60-100 (35-120) Jlm, radial specimen of S. sinensis; prismatic, often in diameter up to 195 Jlm, walls 2-2.5 Jlm tegumented, in normal or slightly enlarged thick. Vessel member length 1090-1380 chambered axial parenchyma cells, usually in (690-1850) Jlm. Perforations scalariform chains of 4 in most material of S. sinensis. with 13-19 (7-25) bars, distance between Note: In the limited material studiedS. rubi bars 7-15 Jlm. Intervessel pits nonvestured, ginosa can be separated from S. sinensis on scalariform or scalariform-opposite, oval to the basis of its stronger ray heterogeneity (1-7 elongate, 13-20 (5-29) Jlm in horizontal rows of square to upright marginal cells). diameter, with slit-like apertures. Vessel-ray pits scalariform to opposite, half-bordered Discussions with weakly reduced borders to almost sim Taxonomic implications ple, often unilaterally compound. Vessel Anticipating the results of a generic wood parenchyma pits simple to half-bordered, op anatomical survey of the Theaceae from their posite or diffuse. Spiral thickenings restricted entire distribution area (Deng & Baas, in pre to vessel element tails. Thin-walled tyloses paration) we will discuss the systematic sig usually present in S. sinensis. nificance of the present resul ts on Chinese spe Fibre-tracheids 1760-2640 (970-3040) cies only. This is at the present stage only pos Jlm long, F/V ratio 1.59-2.19, thin- to thick sible for genera which are largely confined to walled, usually medium thick-walled, with China. In addition wood anatomical argu distinctly bordered pits of 6-8 (5-10) Jlm, ments will be brought forward on the contro numerous in radial and tangential walls. versial delimitation of Stewartia and Hartia. Downloaded from Brill.com10/11/2021 03:27:50AM via free access 370 IAWA Bulletin n.s., Vol. 11 (4), 1990 Table 13. Selected wood anatomical features of the Theaceae from China. Vl ~ 0. *Vl 'l:l 11 Vl 13 ~ (1) 8 u 0. Vl "3 ::l Vl .!:l 8 1:l (1) 'iil ] a'" ;. ....'" ~ .0 u 0 -5 ...... S .::: ~ 8: 0 'l:l .... 0 .~~ .... '"00 ",0. .::: ;a'" .S ~ II '8 's, (1) +1 ·a·~ .;.:: 0 ~'" ;...... § .::: Vl ~ roO .::: 0 .~ 'l:l * ~ '"(1) ~.!:l (1) ro -5 * ...... ;. (1) ro 1~ 00 .... '" 0 .... (1)<1) ~ '§ t 5 "'''' \3~ .~ ;.... '" ~ts ;. (1) 0. '"~ '"ro .5~ ;'.0 roO. u .... Ternstroemioidae S '" Adinandra 31-48 + Anneslea 33-55 t + Cleyera 33-42 + Eurya 51-108 ±/- + Eyryodendron 20 + Ternstroemia 24-36 p + Sladenia + + 17 ± aci Camellioideae Apterosperma + 23 + Camellia + 11-42 ±/- r(eci) Gordonia + 14-25 t Hartia + 42-57 ±/t Parapyrenaria + 8 Pyrenaria + 7-17 Schima + 9-18 t/- aci(e) Stewartia + 13-19 aci * t = restricted to the tip of vessel members; p = restricted to pit apertures. ** r = in ray cells; a = in axial parenchyma cells; c = in chambered cells; e = in enlarged cells; i = integumented. Conclusions on affinities and classifica supported by marked differences in vessel tion of higher categories (above the genus ray pitting (half-bordered in the former; with level) can only be preliminary at this stage. reduced borders to large and simple in the Nevertheless three conclusions, partly con latter); 3) the genus Sladenia is distinct on ac firming earlier studies on a worldwide sam count of its high degree of vessel grouping in ple (Metcalfe & Chalk 1950; Keng 1962) can long radial multiples and its opposite to alter be made (see also Table 13): 1) despite varia nate intervessel pits (the other Theaceae have tion in a number of qualitative and quantita predominantly solitary vessels and scalari tive features, the family Theaceae is wood form to opposite intervessel pits). The taxo anatomically a very homogeneous one, char nomic position and status of Sladenia is dis acterised by scalariform vessel perforations, cussed separately below. fibre-tracheids, diffuse axial parenchyma, There is a tendency for some other fea and heterocellular rays; 2) The two subfami tures to be associated with the above group lies Ternstroemioideae and Camellioideae are ing: e. g. the number of bars per perforation Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. Baas - Wood anatomy of Theaceae from China 371 plate is usually higher in the Ternstroemioi The taxonomic status of Hartia and Stewartia deae than in Sladenia and the Camellioideae Several taxonomists have advocated the and rays of two more or less distinct sizes are reduction of Hartia to Stewartia (Airy Shaw confined to the Temstroemioideae. As can be 1936; Cheng 1934; Sealy 1958; Spongberg seen from Table 13, there are, however, not 1974). However, on the basis of extensive able exceptions to these trends. morphological and chemotaxonomic studies Yan (1981) and Ye (1982) supported the re The position of Sladenia instatement of Hartia. Wood anatomically The monotypic genus Sladenia occurs in there are two striking differences between Southwest China (Yunnan and Guangxi) and Hartia and Stewartia, viz., in average number in the border regions with Burma and Thai of bars per scalariforrn perforation and pres land. From the time of its first description by ence or absence of chambered crystals (Table Kurz in the Theaceae its affinities have been a 13). These differences were confirmed by in matter of debate. Gilg (1893) assigned it to cluding some species of Stewartia from out the Dilleniaceae, but later (Gilg & Werder side China (S. monodelpha and S. pseudo mann 1925) transferred it to the Actinidia camellia). Therefore we have provisionally ceae. Hallier (1923) incorporated Sladenia in kept Hartia separate from Stewartia. The dif his wide concept of the Linaceae. Airy Shaw ference of bar number may be related to dif (1965) raised the genus to family rank, but ferences in phenology: Stewartia is decidu considered its closest affinities to be with ous, and probably has higher demands for Theaceae. Kobuski (1951) and Keng (1962), conductive efficiency (reduced resistance to however, advocated to retain Sladenia in the flow) in the early part of the growing sea Theaceae. The latter view can be supported son, which might explain the low number by our wood anatomical evidence. of widely spaced bars. Hartia is evergreen The wood of Actinidia differs from that of and probably has more moderate and less Sladenia in the presence of two types of ves fluctuating transpiration rates throughout the sel members in alternating zones, and in hav year. ing simple perforations in at least part of the vessel elements (cf. Metcalfe & Chalk 1950, Ecological trends in the Theaceae woods confirmed by personal observations). If Sau from China rauia and Clematoclethra are included in the Within China the Theaceae cover a rather Actinidiaceae the differences become less, but restricted ecological range in tropical to sub are still larger than between Sladenia and the tropical forests to the South of the Yangzhi Theaceae. The same applies when one com river between 19° and 32° N from low altitude pares the wood anatomy of Sladenia with that up to 2000 m. Van den Oever et al. (1981) of the Dilleniaceae (cf. Metcalfe & Chalk 1950; and Baas (1982) have warned against the Dickison 1967). The Linaceae are wood ana hazards of searching for ecological trends in tomically even more different from Sladenia. material from limited ecological ranges, Especially the chambered crystals of Sla because any ecological tendencies may be denia, shared with a number of Camellioi overshadowed by noise variation or by other deae (and absent from Actinidiaceae and Dil factors. Because of this, and also because leniaceae) may represent a synapomorphic precise data on important parameters such as trait. This would invalidate Keng's (1962) altitudinal range or ecological niche in the treatment of Sladenia as a tribe in the Tern forest were usually not available, we have stroemioideae. To do full justice to the only recognised two different ecological cate unique features of Sladenia within the Thea gories for the Theaceae from China: tropical ceae (apart from the high degree of vessel and subtropical. Examples of typical tropical grouping and the opposite to alternate inter provenances are Hainan Island and Xi shu an vessel pits, also its dichasial cymes, dilated Banna in Yunnan with a latitudinal range of filaments and lack of foliar sclereids, cf. 19-22° N, annual rainfall of 1400-1800 Keng 1962) a status of subfamily, Sladenioi mm, and average maximum temperatures of deae, within the Theaceae is advocated by us. 24°-26° C. For subtropical provenances the Downloaded from Brill.com10/11/2021 03:27:50AM via free access 372 IAWA Bulletin n.s., Vol. 11 (4), 1990 40 -- - subtropical species -- - subtropical species -- tropical species 50+ r---, -- tropical species I I I I I I I I 30 40 I I I I "'c "'c I I I I E" 1---' "E I I '0 I I '0 I L __ I I I "g. "g- 30 I r---1 L __ -, I "0 20 I "0 I I I " I 0> I I I ---, "ro I I ~ I C 20 I I I I I I e I L __ -, "~ I I I I I Co "Co 10 I " I I I I I I I I I I I 10 I I L __ I I I 1--...1 L __ -, I I I I I L __ ,-__ I I I \---.., VL , VD 600 800 1000 1200 1400 1600 1800 2000 (Jun) 20 30 40 50 60 70 80 90 (~m) average vessel member length average vessel diameters 45 -- - subtropical species -- tropical species 40 35 ~ 30 ---, E I .~ I I g- 25 I I "0 I I ~ 20 r- I I I C I L __ -, I "~ 15 I I Co I I " I I I I 10 I L __ -, I I I Figs. 36-38. Frequency diagrams of quan __ ...r--_.J I I ,---, titative vessel attributes in tropical (n = 20) I L __ -' I I and subtropical (n = 160) samples from Chi 50 100 150 200 250 300 350 400 I na. - 36. Vessel member length. - 37. Vessel average vessel frequency mm 2 diameter. - 38. Vessel frequency. latitudinal range is 22-32° N, with 1100- Stewartia, and Ternstroemia, and conform to 1500 rnrn of precipitation and average maxi well established general dependencies of mum temperatures from 15°_22° C (Wu wood anatomy on ecology (Baas 1973,1976, 1980). 1982, 1986b; Baas et al. 1983; Baas & Zhang Although there is a tremendous overlap in 1986; Baas & Schweingruber 1987; Carlquist quantitative values between species and spe 1975, 1980, 1984a & b; Carlquist & Hoek cimens of tropical or subtropical origin, Fig man 1985; Van der Graaff & Baas 1974; Van ures 36-38 show that in the tropical material den Dever et aI, 1981). (20 specimens) vessel members tend to be For number of bars per scalariform perfo longer and wider, while vessel frequency is ration there are no clear differences between lower than in the subtropical sample (160 tropical and subtropical species (a similar ab specimens). These trends can also be retraced sence of latitudinal trends has been reported within individual genera in China such as by Van der Graaff & Baas (1974) and Van Adinandra, Eurya. Gordonia, Pyrenaria, den Dever et al. (1981). Other features whose Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. Baas - Wood anatomy of Theaceae from China 373 expression tends to differ in tropical and sub euphlebia) and the other 7 species from lime tropical material are growth rings (less dis stone forests. Averages for vessel member tinct in the tropical than in the subtropical length (1250-1400 11m) and vessel diameter specimens) and spiral vessel wall thickenings (40 11m) in the acid soil species are higher (more frequently present and more distinct in than in species from limestone substrates the subtropical sample). For the family as a (850-1190 11m, and 30-35 11m), while ves whole or for individual genera, the trends sel frequency tends to be slightly higher in would become much more obvious if mate the latter. Although all species have crystals rial from the lower latitude tropics would be in ray cells, their frequency is invariably included in the comparison (cf. Deng & higher in the samples from limestone sub Baas, in preparation). strates. These results agree with a more xeric Within the genus Camellia, of which num ecology on limestone than on acid soils and is erous species and samples were studied, a also in accordance with Rury (1985) who very weak latitudinal correlation was found found that wood inclusions reflected chemical for vessel member length, vessel diameter composition of soils and parent rocks in and vessel frequency in section Thea. Within Erythroxylum. section Chrysantha (Group I in Table 7) there In the notes following the generic descrip is an interesting difference between the spe tions some instances of ecological trends are cies from acid soils (c. chrysantha and C. further specified. Generic wood anatomical key to the Theaceae from China Cautionary note Using the dichotomous key below, it is possible to key out genera or groups of species within genera, as they were represented in our sample. Several qualitative and quantitative dif ferences between genera will doubtlessly break down if material from outside China were to be included (see also remarks in the chapter on ecological trends). The key should therefore only be used for Theaceae of unknown identity from China. l. Vessels mostly in radial multiples of2-6 ...... Sladenia 1. Vessels mostly solitary ...... 2 2. Vessel-ray pits always simple or with reduced borders...... 8 2. Vessel-ray pits mostly fully half-bordered...... 3 3. Rays up to 3-4(-6) cells wide...... 4 3. Rays up to 2(-3) cells wide or mainly uniseriate...... 7 4. Local wall thickenings associated with pit apertures throughout vessel members Ternstroemia 4. Local wall thickenings absent; normal spirals weakly developed or absent...... 5 5. Number of bars per perforation mostly less than 30 ...... Euryodendron 5. Number of bars per perforation mostly over 30 ...... 6 6. Multiseriate rays mostly 4-6 cells wide ...... Anneslaea 6. Multiseriate rays always 2-4 cells wide ...... Eurya 7. Distance between coils of spirals 8-15 (2-30) 11m ...... Cleyera 7. Distance between coils of spirals 3-4 (2-5) 11m ...... Adinandra 8. Average number of bars per perforation over 40 ...... 9 8. Average number of bars per perforation less than 40 ...... 10 9. Crystals present in ray cells ...... Camellia IV 9. Crystals absent ...... H artia 10. Spiral thickenings present on vessel walls ...... 11 10. Spiral thickenings weakly developed or absent ...... 16 Downloaded from Brill.com10/11/2021 03:27:50AM via free access 374 IAWA Bulletin n.s., Vol. 11 (4), 1990 11. Spiral thickenings present throughout vessel elements ...... Apterosperma 11. Spiral thickenings restricted to tails of vessel elements ...... 12 12. Rays 1-2 cells wide ...... , Schima p.p. 12. Rays wider ...... 13 13. Rays mostly 1-2(-3) cells wide ...... 14 13. Rays mostly 1-4(-5) cells wide ...... 15 14. Crystals present in chambered parenchyma cells ...... Schima wallichii 14. Crystals absent ...... Gordonia axillaris 15. Average vessel diameter over 55 ~m, vessel frequency less than 90/mm2 ..... Stewartia 15. Average vessel diameter less than 55 ~m, vessel frequency over 95 /mm 2 Gordonia hainanensis 16. Crystals present ...... 17 16. Crystals absent ...... 19 17. Crystals in chambered axial parenchyma cells ...... Schima p. p. 17. Crystals in ray cells ...... 18 18. Crystals not integumented, in normal ray cells ...... Camellia I 18. Crystals integumented, mostly in enlarged cells ...... Camellia II 19. Multiseriate rays mostly 4-6 cells wide ...... Parapyrenaria 19. Multiseriate rays mostly 2-4 cells wide ...... Pyrenaria, Camellia III Concluding remarks the section of plant morphology and anat Although relatively constant in many of omy, Department of Biology of Peking Uni their wood anatomical features, the Theaceae versity (Beijing); Mr. Su Zhonghai, Guang from China have allowed the construction of dong Institute of Forestry (Guangzhou); an artificial key to identify genera or spe Prof. Ho Tienxiang, Prof. Zhang Hongda cies groups within genera, and to contribute and Dr. Cheng Baoliang, Sunyixian Univer meaningfully to problems on generic delimi sity (Guangzhou); Prof. Xiao Shaoqiong, tation and suprageneric classification. The Southwest College of Forestry (Kunming); existence of ecological trends in the wood Ms. Hou Deshu, Sichuang Agricultural Col anatomy could be established, and enhances lege (Yaan, Sichuang); Mr. Zhang Shuyin, our understanding of the underlying causes Anhui Agricultural College (Heifei); Mr. Xu of the limited wood anatomical diversity of Feng, Guangxi College of Forestry (Nan the family. A survey of the wood anatomy of ning); Prof. Liang Shengye, Guangxi Insti all genera of the Theaceae from its entire dis tute of Forestry (Nanning); Mr. Zhu Junxing, tribution area is needed to analyse the phylo Guangxi Wood Company (Nanning); Mr. Lu genetic significance of the wood anatomical Yongqing, Liuzhou Wood Company in variation. Such a survey is in progress and Guangxi (Liuzhou); Prof. Luo Zhongchun, will also explore possible additional ecologi Xingning Institute of Forestry in Hunan cal trends and strategies. (Xingning); Prof. Pen Guangqian, Hunan Institute of Tea Science (Changsha); Mr. Luo Acknowledgements Wenji, Guizhou Wood Company (Guiyang); We wish to acknowledge the financial sup Prof. Zhang Shungao, Yunnan Institute of port under the agreement between Peking Tea Science (Xishuang Banna); Mr. Wang University and Leiden University, which en Hong, Xishuang Banna Botanical Garden in abled the first author to carry out this study at Yunnan (Xishuang Banna); Dr. D.F. Cutler the Rijksherbarium/Hortus Botanicus (Lei and Ms. M. Gregory (Jodrell Laboratory, den). We are very grateful to the following Kew). Prof. Dr. Hsuan Keng (Singapore) persons and institutes for providing wood kindly gave taxonomic advice. Our thanks are samples and help: Prof. Dr. Li Zhengli, Prof. also due to Bertie Joan van Heuven (Lei Zhang Xinying and other staff members of den) for wide-ranging technical assistance. Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. 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