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Downloaded from Brill.Com10/11/2021 03:27:50AM Via Free Access 338 IAWA Bulletin N.S., Vol 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. Downloaded from Brill.com10/11/2021 03:27:50AM via free access 338 IAWA Bulletin n.s., Vol. 11 (4), 1990 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. Downloaded from Brill.com10/11/2021 03:27:50AM via free access Deng Liang & P. Baas - Wood anatomy of Theaceae from China 339 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.
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