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(Araliaceae). Ii IAWA Journal, Vol. 22 (3), 2001: 301–330 WOOD ANATOMY OF SCHEFFLERA AND RELATED TAXA (ARALIACEAE). II. SYSTEMATIC WOOD ANATOMY OF NEW CALEDONIAN SCHEFFLERA by Alexei A. Oskolski1 & Porter P. Lowry II2a & b SUMMARY The wood anatomy of 22 of the 26 species of Schefflera occurring in New Caledonia was studied. Only two features (the presence of scalari- form perforation plates and scanty paratracheal axial parenchyma) ap- pear to be constant throughout the species examined. The pattern of wood structure diversity was analyzed using PCA; the results generally agree with the current recognition of four groups of species among New Caledonian Schefflera based on macromorphology. Three of these groups (Dizygotheca, “Canacoschefflera” and “Gabriellae”) represent natural assemblages closely related to one another. The fourth group (Schefflera sect. Schefflera) is isolated from the others, as indicated by its very large rays and abundant septate fibres. The occurrence of crys- tals in chambered cells of axial parenchyma was observed for the first time in Araliaceae. The wood structure of Schefflera plerandroides, previously placed in the segregate genus Octotheca, shows no essential differences from that of the other members of the Dizygotheca group, supporting the hypothesis that polymerous flowers have evolved inde- pendently at least twice within the Schefflera alliance. Key words: Araliaceae, Schefflera, Dizygotheca, Canacoschefflera, New Caledonia, systematic wood anatomy, crystals, chambered cells. INTRODUCTION Schefflera J.R. Forst. & G. Forst. is the largest genus of Araliaceae, with an estimated number of species ranging from 400 (Grushvitzky et al. 1985) to 650–700 (Lowry 1989), or as high as 900 (Frodin 1995), and making up more than half of the family. Schefflera is currently circumscribed broadly (Frodin 1975, 1982, 1989, 1993; see also Lowry 1989) to include several segregates usually recognized as synonyms (e.g., Agalma, Brassaia, Cephaloschefflera, Crepinella, Geopanax, Heptapleurum, Neocus- sonia, Scheffleropsis, and Sciadophyllum), plus several others that have been main- 1) Botanical Museum, V.L. Komarov Botanical Institute of the Russian Academy of Sciences, Prof. Popov str. 2, 197376 St. Petersburg, Russia [E-mail: [email protected]]. 2a) Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299, U.S.A. [E-mail: [email protected]]. 2b) Laboratoire de Phanérogamie, Muséum National dʼHistoire Naturelle, 16 rue Buffon, 75005 Paris, France [E-mail: [email protected]]. Downloaded from Brill.com10/09/2021 01:50:26AM via free access 302 IAWA Journal, Vol. 22 (3), 2001 Oskolski & Lowry II — Wood anatomy of Schefflera 303 tained as distinct in some treatments (e.g., Didymopanax, Dizygotheca, Octotheca, and Plerandra). This broad interpretation of Schefflera has, however, been called into question in a recent study (Wen et al. 2001) in which molecular data suggest that the genus, as currently defined, may be polyphyletic. Species of Schefflera (sensu lato) occur in most tropical and subtropical regions, with especially high concentrations in Malesia, Southeast Asia, New Caledonia, the Andes and the Guyana Highlands. Among these areas, the small southwest Pacific island of New Caledonia (c. 17,000 km2) is remarkable because of its high levels of diversity and endemism, both within Schefflera (26 species) and among Araliaceae as a whole, with a total of eight genera and over 90 native species recorded, only two of which occur elsewhere (Lowry 1986b, 1989, unpubl. data; Lowry et al. 1986). The New Caledonian Schefflera species, along with a few close relatives on neighboring islands (cf. Smith 1985; Lowry 1989), also appear to be rather isolated systematically and their relationships to other groups in the genus are somewhat obscure. The diversity and distinctiveness of the New Caledonian flora in general, and of its Araliaceae in particular, are due in large part to the wide variety of habitat types present on the island, which result from the complex interactions of topography, cli- mate, and a special set of edaphic features, including large areas of ultramafic soils (cf. Morat 1993; Lowry 1998). This is compounded by the islandʼs long geological history, during which it has been isolated from Australia for approximately 80 million years (Kroenke 1996), serving as a refuge for many “primitive” vascular plants (Raven & Axelrod 1972, 1974; Raven 1980), including representatives of some basal line- ages of Araliaceae (Plunkett & Lowry 2001). All 26 species of New Caledonian Schefflera are endemic to the island (Lowry, unpubl. data). Their habit varies from small monocaulous treelets to large, well branched trees up to 20 m in height. Species occur primarily in moist evergreen for- ests; the genus is less well represented in sclerophyllous forests and maquis vegeta- tion (cf. Jaffré et al. 1993; Lowry 1998). Four well-defined groups can be distin- guished within the genus in New Caledonia based on morphological differences in the leaves, inflorescences, flowers, and fruits, as follows: 1) Schefflera sect. Schefflera is represented by three species in New Caledonia, with five additional members occurring in other island groups of the southwest Pacific (one species each in Vanuatu, Samoa, and New Zealand, and two species in Fiji; Lowry 1989). The type of the genus (S. digitata J.R. Forst. & G. Forst.) from New Zealand belongs to this section, which could thus be interpreted as “Schefflera in the strictest sense” (Lowry 1989). This group is distinguished by its paniculate in- florescences bearing numerous umbellules or racemules of small flowers with 5 stamens and an equal number of carpels and free styles, and its thin, often mem- branaceous leaves and almost succulent stems and petioles. 2) The Dizygotheca group, previously recognized as a distinct genus Dizygotheca N.E. Brown (including Octotheca R. Vig.) by several authors (e.g., Guillaumin 1948; Hutchinson 1967), has twelve species in New Caledonia (all endemic) and two additional representatives in Vanuatu (Lowry 1989). Members of this group are characterized by anthers with 8 thecae (vs. 4 thecae in all other Schefflera) and Downloaded from Brill.com10/09/2021 01:50:26AM via free access 302 IAWA Journal, Vol. 22 (3), 2001 Oskolski & Lowry II — Wood anatomy of Schefflera 303 a distinctive paniculate-umbellate inflorescence structure in which the lateral axes usually have a median pseudo-whorl of umbellules (cf. Lowry 1989). They also have a chromosome number of n = 24 (as compared to n = 12 for nearly all other Schefflera examined to date) (Lowry 1986b, unpubl. data). Most Dizygotheca in New Caledonia have free styles (united in two species), and a majority are found on non-ultramafic substrates. Octotheca was originally described as a monotypic genus on the basis of its polymerous flowers (Viguier 1906; see also Viguier 1909, 1925; Hutchinson 1967; Eyde & Tseng 1971; Tseng & Hoo 1982) with 10–15 carpels and 10, 15 or 20 stamens (vs. 5–10 carpels and 5 stamens in Dizygotheca), but it otherwise shares all of the features that characterize the group, and should probably not be retained. 3) The “Canacoschefflera” group is an informally recognized assemblage of nine en- demic species distinguished by their once- or twice-umbellate inflorescences bear- ing 2–5-carpellate flowers with an equal number of free or basally united styles, and leaves that are usually quite coriaceous. Species of “Canacoschefflera” occur primarily on ultramafic substrates at higher elevations. 4) The “Gabriellae” group is represented by two species in New Caledonia, and two others occurring elsewhere in the southwest Pacific,S. vanuatua Lowry in Vanuatu (Lowry 1989) and S. seemanniana A.C. Smith in Fiji (Smith 1985). Members of this group are characterized by strictly compound umbellate inflorescences bear- ing generally 5-merous flowers with the styles fused into a short beak, and leaflets with numerous parallel secondary veins. The New Caledonian species are often large trees that can play an important role in the composition of forest communi- ties, with one (S. gabriellae Baill.) occurring primarily on ultramafic substrates and the other (S. pancheri Baill.) found almost exclusively on other soil types. Wood anatomy has been shown to be a useful tool for examining infrageneric rela- tionships within both Schefflera (Oskolski 1995) and other groups of Araliaceae (Rodriguez 1957; Oskolski 1996, 2001; Oskolski et al. 1997; Oskolski & Lowry 2000). Oskolski (1995), using a representative sample of members of the genus from through- out the world, recognized seven broad groups within Schefflera on the basis of their wood structure, and proposed interpretations of their possible systematic relation- ships. However, data on the wood anatomy of New Caledonian Schefflera were scanty, and hence insufficient to formulate any conclusive interpretations about the position of the four groups represented on the island. In fact, only two (or perhaps three) of these species have been examined previously [S. gabriellae (Sarlin 1954; Oskolski 1994, 1995, 1996); S. elegantissima (Veitch ex Masters) Lowry & Frodin (Oskolski 1994, 1996 as Dizygotheca elegantissima), and Dizygotheca sp. (Record & Hess 1944; Metcalfe & Chalk 1950)]. The present study, which is part of a general survey of wood anatomy throughout Araliaceae (Oskolski 1994, 1995, 1996; Oskolski et al. 1997; Oskolski & Lowry 2000), surveys the wood structure of 22 of the 26 species of Schefflera represented in New Caledonia. The results are considered
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