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WOOD ANATOMY of SCHEFFLERA and RELATED TAXA (ARALIACEAE) by Alexei A

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WOOD ANATOMY of SCHEFFLERA and RELATED TAXA (ARALIACEAE) by Alexei A IAWA Journal, Vol. 16 (2),1995: 159-190 WOOD ANATOMY OF SCHEFFLERA AND RELATED TAXA (ARALIACEAE) by Alexei A. Oskolski V. L. Komarov Botanical Institute of the Russian Academy of Sciences, Botanical Museum, Prof. Popov str. 2, 197376 St. Petersburg, Russia SUMMARY The wood anatomy of 31 Schefflera species from Indochina, Australia, Oceania, Africa, and South America, 3 species of Didymopanax from South America, and Tupidanthus calyptratus and Scheffleropsis hemi­ epiphytica from Indochina (Araliaceae) are described. Seven groups of species can be recognised. Tupidanthus and Scheffleropsis are very similar to the Schefflera spe­ eies from sections Schefflera (subseetions Octophyllae and Heptapleu­ rum) and Brassaia (subsection Actinophyllae). These taxa have charac­ teristic vessel-ray and vessel-axial parenchyma piuing formed by solitary large pits among numerous distinctly smaller ones. Didymopanax cannot be separated from the South American Schefflera species with respect to wood anatomy. Seetion Agalma of Schefflera differs from other studied taxa by the presence of helical thickenings on the vessel walls. The relationships of the examined species is discussed. The presence of radial canals in Schefflera and allied taxa is regarded as a primitive feature. The mountain species tend to have more distinct growth rings, more numerous and narrower vessels, and wider and (or) higher rays than those in the lowland. Helical thickenings and abundant vascular tra­ cheids are restricted to mountain species. Schefflera digitata from New Zealand, the only temperate Schefflera species, shows these features in the most pronounced form, but lacks helical thickenings. It also has the shortest vessel elements in the genus. Key words: Araliaceae, Schefflera, Scheffleropsis, Tupidanthus, Didymo­ panax, systematic wood anatomy, ecology. INTRODUCTION Schefflera 1. R. & G. Forst. is the largest and geographically most widespread genus in the Araliaceae. According to different estimates it comprises from 400 (Grushvitzky et al. 1985) to 650-700 species (Lowry 1989). It is found in most tropical and subtropi­ cal regions, but is especially common in Southeast Asia, Madagascar, New Caledonia, the Andes and the Guyana Highlands (Frodin 1975). The genus is typified by the only species of New Zealand S. digitata J.R. & G. Forst. (which is the only species from Downloaded from Brill.com10/07/2021 09:20:01PM via free access 160 IAWA Journal, Vol. 16 (2),1995 Table 1. Correspondence between the subdivisions of the different arrangements of Schefflera (with important diagnostic characters) and the wood anatomical groups. Harms (1894) Viguier Hoo & Tseng (1965) Groups of Inflorescence Position (1909) species in type of styles (series) present study secr. Euschejjlera Ib secr.Agalma A racemes or united subsecr. Agalma spikes into column Ia, III secr. Schejjlera B, Cl umbels in subsecr. Octophyllae panicies -- -- - subsecr. IIb (a, ß. y) subsecr. Heptapleurum B none, Heptapleurum stigma sessile ? IIb (0) subsecr. Digitatae C2, C3 free or Dl, D2 united at the base secr. Cephaloschejjlera IIa secr. Brassaia capitula subsecr. Cephaloschejjlera subsecr. Actinophyllae B united into column I I temperate latitudes).The Schefflera species are mostly treelets with unbranched trunks or with few unarmed branches bearing terminal whorls of digitately compound leaves. Besides tall trees (up to 20 m high), shrubs, hemi-epiphytes, epiphytes and climbers occur within the genus. The classification of Schefflera is subject to continuous debate. Infrageneric group­ ings were proposed by Rarms (1894), Viguier (1909) and Roo and Tseng (1965); the correspondence between these divisions and the distribution of the main taxonomic features (inflorescence type, structure and position of styles and stigmas) is repre­ sen ted in Table 1. The generic rank of seetion Cephaloschefflera Rarms has been advo­ cated by Merrill (1923) and Rutchinson (1967, who called this genus Brassaia Endl.); moreover, Rutchinson (1967) after Miquel (1855) regarded seetion Agalma as a sepa­ rate genus. On the other hand, Frodin (1975) considered seetion Cephaloschefflera Rarms as an artificial group representing "an evolutionary grade (probably resulting from parallel retardation of pedicel growth) derived from various groups of umbelluliferous species throughout the geographical range ofthe [Cephaloscheffleral complex." Rowever, he recognised section Brassaia as coinciding approximately with subseetion Actinophyllae Tseng & Roo. There is a number of genera closely related with Schefflera whose rank is a matter of discussion. Three genera of that kind (Tupidanthus Rook. f. & Thoms., Scheffleropsis Ridley and Didymopanax Decne. & Planchon) are considered in this paper. Downloaded from Brill.com10/07/2021 09:20:01PM via free access Oskolski - Wood anatomy of Schefflera s.l. 161 Tupidanthus Hook. f. & Thoms. is a monotypie genus: the only speeies, T. calyptratus Hook. f. & Thoms., is a seandent shrub or treelet native to tropieal Asia from India to the Malay Peninsula. This genus is clearly distinguished by its numerous stamens (up to 70) and nearly 100-10eular ovary, while SchejJlera speeies as weIl as many other Araliaeeae usually have 3-5-merous flowers. That is why Tupidanthus was often eon­ sidered as the most primitive genus in the family (Harms 1894; Viguier 1909; Li 1942; Grushvitzky & Skvortsova 1973).Eyde and Tseng (1971) ehallenged this point ofview and regarded flower polymery as a result of seeondary multiplieation of flower or­ gans. Most authors recognise the generic rank of Tupidanthus, but Frodin (in Stone 1978) and Lowry et aL (1989) include this taxon in SchejJlera. SchejJleropsis Ridley eomprises 4 speeies whieh are treelets and hemi-epiphytes native to Indochina. This genus is closely related to Tupidanthus, but differs from it by less numerous flower parts (8-28 stamens, 8-23-loeular ovary). The idea of the ge­ nerie rank of SchejJleropsis has been proposed by Grushvitzky and Skvortsova (1973), whereas other authors (Hutchinson 1967; Eyde & Tseng 1971) did not support this concept. Didymopanax Decne. & Planchon numbers c. 25 species of trees and shrubs and is widespread in tropical America. This genus is distinguished by its bilocular ovary and has reeently been included in SchejJlera by Maguire et aL (1984) and Cannon and Cannon (1989). Wood anatomieal data on a limited number of the SchejJlera speeies were given by Moll and Janssonius (1918), Kanehira (1921), Leeomte (1926), Fujioka (1927), Benoist (1931), Tang (1932), Sarlin (1954), Rodriguez (1957), Lindeman et aL (1963), Ver­ steegh(1968), Braun (1970), Butterfieid and Meylan (1976), Meylan and Butterfieid (1978), Dechamps (1979), Ilie (1991), and Oskolski (1994). Many authors (Benoist 1931; Williams 1936; Record & Hess 1943; De Bastos 1946; Tortorelli 1956; Rodriguez 1957; Ragonese 1961; Lindeman et aL 1963; Sudo 1963; Dechamps 1979; Detienne & J aequet 1983; Ilie 1991; Oskolski 1994) studied the wood structure of Didymopanax morototoni (AubI.) Deene. & Planchon from the point of view of the economie signi­ ficanee of its timber; the wood anatomy of few other Didymopanax speeies was de­ scribed by Alves de Pinho (1966), Van der Slooten et aL (1970), and Mainieri and Chimelo (1989). Metcalfe and Chalk (1950) provided data on the wood anatomy of SchejJlera and Didymopanax (species not given). The data on the wood of Tupidanthus and SchejJleropsis have hitherto not been presented except by Oskolski (1994). This study surveys the wood anatomy of Schefflera and related taxa such as Tupidanthus, SchejJleropsis, and Didymopanax, and discusses both taxonomie and eco­ logical aspects. The work is made within the framework of the general study on the wood anatomy of Araliaceae (Oskolski 1994). MATERIALS AND METHODS Wood sampies were obtained from various institutional wood colleetions (SFCw, Bw, LEw, USw) and from Dr. LV. Grushvitzky and Dr. Nina T. Skvortsova who had col­ leeted them during their N orth Vietnam expeditions in 1963, 1966, and 1969 (these Downloaded from Brill.com10/07/2021 09:20:01PM via free access 162 IAWA Journal, Vol. 16 (2), 1995 Table 2. List of the species studied and their assignment to wood anatomical groups. Species Locality of collection Wood anatomical group Didymopanax Decne. & Planeh. D. angustissimum E. Marchai Brazil Dl D. macrocarpum Seem. Brazil Dl D. morototoni (Aub!.) Decne. & Planeh. South America Dl Schefflera J. R. & G. Forst Sect. Agalma (Miq.) Tseng & Hoo Subsect. Agalma Harms S. delavayi (Franch.) Harms & Diels North Vietnam A S. vietnamensis Grushv. & N. Skvorts. North Vietnam A Subsect. Glummea Grushv. & N. Skvorts. S. fasciculifoliolata Grushv. & N. Skvorts. North Vietnam A S. laxiuscula Grushv. & N. Skvorts. North Vietnam A Sect. Schefflera Tseng & Hoo Subsect. Schefflera (= Digitatae Tseng & Hoo) S. barteri (Hoch.) Harms Cameroon D2 S. digitata J. R. & G. Forsl. New Zealand C2 S. europhylla Harms Ecuador Dl S. euthytricha A.C. Smith Fiji C2 S. fragrans Cuatr. Colombia DI S. gabriellae Baill. New Caledonia D1 S. hypoleucoides Harms var. tomentosa Grushv. & N. Skvorts. North Vietnam C3 S. mildbraedii Harms EastAfrica D2 S. nono BailI. New Caiedonia Dl S. stuhlmannii Harms Tanzania D2 S. vitiensis (A. Gray) Seem. Fiji C2 S. yurumanguinis Cuatr. Colombia DI Subsecl. Octophyllae Tseng & Hoo S. hypoleuca (Kurz) Harms North Vietnam Cl S. octophylla (LoUf.) Harms North Vietnam B S. pacoensis Grushv. & N. Skvorts. North Vietnam Cl S. palmiformis Grushv. & N. Skvorts. North Vietnam B S. tonkinensis
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