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Downloaded from Brill.Com09/27/2021 06:35:29AM Via Free Access 334 IAWA Bulletin N.S., Vol IAWA Bulletin n.s., Vol. 12 (3),1991: 333-353 THE WOOD ANATOMY OF THE THEACEAE by Deng Liang and Pieter Baas Rijksherbarium/Hortus Botanicus, P.O. Box 9514, 2300 RA Leiden, The Netherlands Summary A general wood anatomical description of Table 1. Genera of the Theaceae * and their the Theaceae is given on the basis of a previ­ approximate numbers of species. ous study of species from China (Deng & Baas 1990) and additional observations on Subfamily Number of Species genera and species outside China. The wood Genus species studied anatomy of Archboldiodendron, Balthasaria, Ternstroemioideae Ficalhoa, Franklinia, Freziera, and Visnea Adinandra 70 8 are described separately. Although the Thea­ Anneslea 4 3 ceae in the delimitation followed here (Table Archboldiodendron 1 1) are wood anatomically relatively homo­ Balthasaria 2or3 2 geneous, the recognition of three subfamilies Cleyera 17 4 Camellioideae, Ternstroemioideae and Sla­ Eurya 70 16 denioideae is supported by anatomical fea­ Euryodendron 1 1 tures (vessel grouping, bar number, type of Freziera** 38 1 vessel-ray pits, intervessel pit arrangement; Killipiodendron 1 o cf. Table 4). The controversial position of *** Symplococarpon *** 1 o Ficalhoa is discussed; its wood anatomy T ernstroemia 85 8 closely resembles that of the Camellioideae. Visnea 1 The Theaceae are wood-anatomically markedly different from the Bonnettiaceae. Camellioideae Quantitative and some qualitative wood Apterosperma 1 1 anatomical features follow general latitudinal Camellia 88 45 trends, previously established for other fam­ Franklinia 1 1 ilies or genera with a largely mesic ecology. Gordonia 70 9 Key words: Systematic wood anatomy, eco- Hartia 14 5 logical wood anatomy, Theaceae, Fical­ Parapyrenaria 2 1 hoa. Pyrenaria 20 5 Schima 1 (30) 6 Introduction Stewartia 6 5 A detailed account of the wood anatomy of the Theaceae native to China was given in Sladenioideae a previous paper (Deng & Baas 1990). In Sladenia that paper, generic descriptions were present­ Insertae sedis ed, as weil a~ discussions on the wood ana­ tomical diversity and its bearing on generic Ficalhoa and subfamily delimitation, together with an * Delimitation of the Theaceae according to analysis of ecological trends within China. In Takhtajan (1987, see also discussion). the present paper we extend the observations ** Freziera includes Freziera and Patascoya to species and genera outside China and dis­ according to Weitzman (1987). cuss implications for classification and eco­ *** Material not available for study. logical trends of the whole family. Downloaded from Brill.com09/27/2021 06:35:29AM via free access 334 IAWA Bulletin n.s., Vol. 12 (3),1991 Figs. 1 & 2. TS, x 32. - 1: Archboldiodendron merrillianum, wood diffuse-parous, growth rings absent. - 2: Balthasaria schliebenii, ibid. - Figs. 3 & 4. Ficalhoa laurifolia. - 3: TS, x 32. - 4: RLS, x 80, heterogeneous ray type III (Kribs), with one or two rows of square to upright marginal cells; note also large and simple vessel-ray pits. Downloaded from Brill.com09/27/2021 06:35:29AM via free access Deng & Baas - Wood anatomy of Theaceae 335 For easy reference the genera of the Thea­ with 40-41 (30-50) bars, distance between ceae, their approximate total number of the bars 3-6 11m. Intervessel pits nonvestured, species and number of speeies studied wood scalariform to opposite, 10-35 (8-50) 11m anatomically are listed in Table 1. The assign­ in horizontal diameter, with sJit-like apertures. ment of genera to subfamilies largely follows Vessel-ray pits half-bordered, scalariform to Keng (1962), with some modifications ex­ opposite and alternate. Vessel-parenchyma plained in our previous paper (Deng & Baas pits half-bordered to simple, scalariform, 1990). For a comprehensive listing of earlier opposite to alternate, or even diffuse. Spiral wood anatomicaJ literature on the Theaceae thickenings absent. Thin-walled tyloses pres­ the reader is referred to Deng and Baas (1990). ent in a few vessels. Fibre-tracheids 2740-2930 (2210-3530) Materials and Methods 11m long, F/V ratio 1.32-1.38, thin- to thick­ The bulk of the material on which the con­ walled, with distinctly bordered pits of 9-12 clusions of this paper are based is the same 11m, numerous in radial and tangential walls. (210 samples, belonging to 95 species and Parenchyma scanty paratracheal and dif­ 15 genera) as used in our paper on Theaceae fuse apotracheal, in 4-7 (2-8)-celled strands. from China (Deng & Baas 1990). In addition, Rays 8-IO/mm, 1-2(-3) cells wide, c. 90 specimens belonging to 28 species 0.38-0.86 (0.10-1.63) mm high, hetero­ were studied. Most of these represented spe­ geneous type II-III, with 1-7 rows of square eies from outside China and belonged to or upright marginal cells. genera already studied; 14 species represent­ Crystals absent. ed 6 genera without representatives in China. Notes: 1. Keng (1962) described the wood Data from the literat ure and from slides in the structure of Archboldiodendron based on Rijksherbarium reference collection (including juvenile wood. Quantitative values in Keng's a duplicate set from the FHOw slide collec­ material are much lower (e.g. vessel diameter tion at Oxford) were also studied. No material 23-39 11m, vessel member length 860-1450 was available of two genera, Killipiodendron 11m, fibre-tracheid length 960-1750 11m). and Symplococarpon. Citation of wood sam­ Keng moreover recorded uniseriate rays of pIes in generic descriptions, microtechnical Kribs type III while in our material the rays procedures, measurements, and descriptive are 1-2(-3) cells wide and belong to Kribs conventions are the same as those explained type II & III. These differences are probably in detail by Baas & Zhang (1986) and Deng caused by the age of the sampies. & Baas (1990). 2. Dur material incIuded one sampie (FP A w 30318) labelIed A. merrillianum but aberrant in its wood structure in having a Results much lower number of bars per perforation and large and simpe vessel-ray pits; presum­ Wood anatomical descriptions 0/ genera out­ ably this sampie belongs to a genus of the side China Camellioideae. Archboldiodendron Kobuski (Fig. 1) Material studied: A. merrillianum Kobuski, Balthasaria Verdc. (Fig. 2) New Guinea: FPAw 5399, FPAw 34962. Material studied: B. schliebenii Verdc., Evergreen trees in tropical montane rain Burundi: Pouilloux 781 (CTFT 20701) - B. forests (alt. c. 2400 m) in New Guinea. schli{!benii Verdc. var. intermedia (Boutique Growth rings indistinct. W ood diffuse­ & Troupin) Verdc., ZaIre: Bouxin 863 (Tw porous. Vessels 31-35/mm2, 84-96% soli­ 24172), Lewalle 6106 (Tw 24306). tary, remainder in tangential and radial or Evergreen trees in tropical montane rain oblique multiples of 2-3, angular in cross forests (alt. 2200-2400 m) in West and East section, tangential diameter 70-95 (40-120) Africa. 11m, radial diameter up to 240 11m, walls 2-3 Growth rings indistinct. Wood diffuse­ 11m thick. Vessel member length 2070-2120 porous. Vessels 27-32/mm2, 83-97% soli­ (1520-2650) 11m. Perforations scalariform tary, remainder in tangential and radial to Downloaded from Brill.com09/27/2021 06:35:29AM via free access 336 IAWA Bulletin n.s., Vol. 12 (3),1991 oblique pairs, angular in cross section, tan­ horizontal diameter, with slit-like apertures. gential diameter 75-80 (45-110) J.UTI, radial Vessel-ray pits simple to half-bordered with diameter up to 160 11m, walls 3-5 11m thick. much reduced borders, scalariform, opposite Vessel member length 1470-1890 (1060- or diffuse. Vessel parenchyma pits half-bor­ 2760) 11m. Perforations scalariform with 30- dered to simple, often in single vertical rows. 40 (19-50) bars, distance between bars 3-5 Spiral thickenings absent. Thin-walled tyloses 11m. Intervessel pits nonvestured, scalariform present in a few vessels. to opposite, 6-30 (4-38) 11m in horizontal Fibre-tracheids 2670 (2340-2870) 11m diameter, with slit-like, occasionally coales­ long, F/V ratio 1.26, thin- to thick-walled, cent apertures. Vessel-ray pits half-bordered, with distinctly bordered pits of 7-10 11m, opposite to alternate. Vessel-parenchyma pits numerous in radial and tangential walls. mainly in single vertical rows, sometimes op­ Parenchyma scanty paratracheal to va si­ posite or alternate. Spiral thickenings absent. centric and apotracheally diffuse and diffuse­ Tyloses absent. in-aggregates, in 1-6-celled strands. Fibre-tracheids 2610-2710 (2210-3310) Rays 9/mm, of two distinct sizes: narrow 11m long, F/Vratio 1.43-1.77, thin- to thick­ rays 1-2-seriate, wide rays (3-)5-7-seriate, walled, with distincdy bordered pits of 8- 1.10 (0.20-2.20) mm high, heterogeneous 10 11m, numerous in radial and tangential type I-III, with 1-6 rows of square and up­ walls. right marginal cells. Parenchyma scanty paratracheal and dif­ Crystals absent. fuse apotracheal, in 4-7-celled strands. Note: Metcalfe and Chalk (1950) gave an Rays 12-15/mm, of two more or less dis­ account of Ficalhoa wood anatomy under tinct sizes: narrow rays 1-2-seriate and wide Ericaceae. See also separate discussion of the rays 3-5-seriate, 0.85-0.94 (0.1-2.0) mm position of Ficalhoa. high, heterogeneous type I, with 1-7 rows of square and upright marginal cells. Franklinia Bartr. ex Marshall (Figs. 5-7) Crystals absent. Material studied: F. alatamaha Marshali, Note: Keng (1962) noted that the rays are USA, Philadelphia (cult.): Madw 3551 (= of two sizes: uniseriate and biseriate, but in Tw 38660), AFw 2309 (= Usw 21142). our material of Balthasaria wide rays are 3-5- Deciduous trees or shrubs, native to SE. seriate. Quantitative values in Keng's material Georgia, U.S.A., now known only in cul­ are higher (e.g. vessel diameter 162 11m, ves­ tivation. sei member length 2340 11m and fibre-tracheid Growth rings distinct, marked by flattened length 2880 11m) than in our material.
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