IAWA Journal, Vol. 16 (2),1995: 159-190

WOOD ANATOMY OF AND RELATED TAXA () 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, , 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. from , 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

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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 Position (1909) species in type of styles (series) present study

secr. Euschejjlera Ib secr.Agalma A or united subsecr. Agalma spikes into column Ia, III secr. Schejjlera B, Cl 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 (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.

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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 . 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 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

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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 Vig. North Vietnam B S. pesavis Vig. North Vietnam B Subsecl. Heptapleurum (Gaertn.) Tseng & Hoo S. globulifera Grushv. & N. Skvorts. North Vietnam B S. glomerulata Li North Vietnam B S. leucantha Vig. North Vietnam B S. petelotii Merr. North Vietnam B S. tomentosa (Blume) Harms East India B S. trungii Grushv. & N. Skvorts. North Vietnam B Sect. Brassaia (End!.) Tseng & Hoo Subsecl. Actinophyllne Tseng & Hoo S. actinophylla (End!.) Harms Queensland B Subsecl. Cephaloschefflera (Harms) Tseng & Hoo S. chinensis (Dunn) Li Yunnan C3 S. volkensii Harms East Africa D2 Scheffleropsis Ridley S. hemiepiphytica Grushv. & N. Skvorts. North Vietnam B Tupidanthus Hook. f. & Thoms. T. calyptratus Hook. f. & Thoms. North Vietnam B

Downloaded from Brill.com10/07/2021 09:20:01PM via free access Oskolski - Wood anatomy of Schefflera s.l. 163 sampies are designated in the Tables 3-5 as Grushv.). Some sampies were collected by the author from the cultivated in the Botanical Garden of the V. L. Komarov Botanical Institute, St. Petersburg. A list of species studied is given in Table 2 [Hoo & Tseng's (1965) arrangement is used in this table with a small modification: subsection Digitatae Tseng & Hoo is called Schefflera following Grushvitzky et al. (1985)]. Wood sampies examined in this study are taken mostly from mature sterns with a secondary xylem radius of more than 10 mm; it is at this distance from the pith that the vessel element length becomes stable as recorded by Baas (1976) in some Araliaceae. A few sampies from young sterns were also studied. The sampies and their stern diameters are listed in Tables 3-6. Standard procedures for wood anatomy were used to prepare seetions and macerations for light microseopie studies. Sampies for scanning electronic microscopy were pre­ pared according to Exley et al. (1977). Descriptive terminology and measurements follow Carlquist (1988) and the IAWA Committee (1989).

RESULTS

The Schefflera, Tupidanthus, Scheffleropsis, and Didymopanax species under study are diverse in their wood anatomy, and four groups of more or less similar species can be distinguished. Besides wood anatomy, geographical distribution of the species and type of pith (septate or non-septate) were taken into account. Groups A and Bare well-defined while Groups C and D can be subdivided into more closely knit, presum­ ably natural assemblages: Cl, C2, C3, Dl and D2. The species composition of the groups is given in Tables 2 and 3; these groups and their diagnostic features are de­ scribed below (and summarised in Table 7). The wood anatomical diversity patterns will be discussed with respect to their significance in systematics and ecological trends in wood anatomy of the group. Quantitative data are summarised in Tables 3-6.

GroupA (Fig. 1-7; Table 3)

Diagnostic characters: helical thickenings on vessel and vascular tracheid walls, mar­ ginal parenchyma present, and vascular tracheids abundant; vessels narrow (diameter less than 70 /-illl) and numerous (more than 120 per sq. mm); multiseriate rays with sheath cells. Pith septate. Species from East Asia.

Material studied: Schefflera delavayi: Vietnam, Hoang Lien Son (Lao Cai), Sa Pa, LEw 3786; Vietnam, Ha Son Binh, Pa Co, Grushvitzky & N. Skvortsova 108 (1969). - S. fasciculifolio-lata: Vietnam, Ha Tuyen, Ha Giang, mont. Phobang, Grushvitzky & N. Skvortsova 41 (1966), type specimen. - S. laxiuscula: Vietnam, Hoang Lien Son (Lao Cai), Sa Pa, LEw 3146. - S. vietnamensis: Vietnam, Hoang Lien Son (Lao Cai), Sa Pa, Grushvitzky & N. Skvortsova 325 (1963), type specimen.

Growth rings distinct, marked by uniseriate marginal bands ofaxial parenchyma and arrangement of vessels. Vessels are angular, rarely rounded in outline, rather narrow (tangential diameter is less than 64 mm) and numerous (up to 257 vessels and vascular tracheids per sq. mm),

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Figures 1-7

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Table 3. Wood anatomical features of Group A.

2 3 4 5 6 7 8 9 10 11 12 13 14

Schefflera delavayi LEw 3786 TL 1500 25/10 + 805 35 12/8 195 010-15 1062 4.0/2-6 0.8/2.6 1.4 3.8 Grushv. 108 (1969) TR 1100 15/6 + 755 33 26/5 201 010-11 1107 3.912-6 0.7/1.2 2.1 3.2 S. fasciculifolio· lala Grushv.41 (1966) TL 2000 20/5 + 902 31 32/6 124 010-11 1097 3.6/2-6 1.113.0 2.9 3.1 S. laxiuscula LEw 3146 TL 2000 17/7 + 768 33 4/15 178 010-5 983 4.4/3-6 0.912.2 1.3 3.5 S. vietnamensis Grushv.325 (1963) TL 1500 25/10 + 820 44 16/16 168 010-4 1144 3.712-6 1.112.6 1.9 5.0

1: Habit (TR = , TL = treelet) - 2: Altitude of habitat (m) - 3: Radius of wood sample/distance from pith to bark (mm) - 4: Growth rings (+ = distinct, +1- = indistinct, 0 = absent) - 5: Average length of vessel elements ünn) - 6: Average tangential diameter of vessels ().1111) - 7: Percentage of solitary vessels/the greatest number of vessels in avessei gronp - 8: Vessel and vascular tracheid frequency (per sq.mm) - 9: Usuallmin.-max. nnmber of bars per perforation plate - 10: Average 1ength of libriform fibres ().1111) - 11: Width of multiseriate rays (average min.-max., cells) - 12: Height of multiseriate rays (average/max., mm) - 13: Number of uniseriate rays per mm - 14: Number of multiseriate rays per mm. solitary and also in radial multiples of2-5 (up to 16 in S. vietnamensis). Vessel walls 2-5 /JIll thick. Tyloses observed in vessels in S. delavayi. Average vessel element length 755-902 /JIll (total range 388-1100 /JIll). Perforations mostly scalariform with few (up to 15) bars in S. delavayi, S. fasciculifoliolata, and S. laxiuscula, or mostly simple, rarely with 1 or 2 (up to 5) bars in S. vietnamensis, in lateral and oblique end walls. Intervessel pits opposite, rarely transitional, 8-21 /JIll in horizontal diameter, rounded or oval, with oval and lens-like apertures. Vessel-ray and vessel-axial paren­ chyma pits half-bordered or with much reduced borders, similar to intervessel pits in size and shape, sometimes unilaterally compound. The vessel walls as well as vascular tracheid walls have helical thickenings. Vascular tracheids are numerous in all the species of the group. Libriform fibres very thin- to thin-walled (walls 3-8 /JIll thick), septate, with sim­ ple to minutely bordered pits with slit-like apertures.

Fig. 1-7. Group A species. - 1, 2. SchejjZera laxiuscula, TS, TSL. - 1: Growth ring boundaries marked by arrangement of vessels. - 2: High multiseriate rays with marginal rows and sheaths of upright and square cells. - 3,4. S. vietnamensis, TS, SEM. - 3: Distinct growth ring bound­ aries marked by marginal bands ofaxial parenchyma. - 4: Helical thickenings on the vessel wall. - 5-7. S. delavayi, SEM. - 5: Scalariform and simple perforation plates. - 6: Vessel-axial parenchyma pits, helical thickenings. - 7: Opposite intervessel pitting. - Scale bar of Fig. 1-3 = 100 f..U11; ofFig. 4, 6, 7 = 10 f..U11; ofFig. 5 = 20 f..U11.

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't1.J lIiOJ\ . ftT1. .:",,","' ~ I ,..~ ~ I.L' ~ rR'I '1!I ..... ~ ~ '"""I ,...... !!I : e ~ [ I • . ~~ ~ ~ 'r! ,. ~ Ilo,~

~,...~F:: ~~ ~ r1 ~ ~ .....L ~ .- . ~ ~ ~ t~ ~ -I: ~ ~ ~ : 8.~ ~L.._ • ~~ • ~lo

Fig. 8-11. Group B species. - 8, 9. SchejJ7eropsis hemiepiphytica, TS, TLS. - 8: Growth rings absent. - 9: Multiseriate rays with incomplete sheaths of upright and square cells, radial canal (arrowed). - 10. SchejJ7era actinophylla, TS, indistinct growth ring boundary marked by radially flattened fibres. - 11. S. trungii, TS, indistinct growth ring boundary marked by arrangement of vessels. - Scale bar of Fig. 9 & Il = 100 J.li11; magnification of Fig. 8 & 10 as in Fig. 11.

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Axial parenchyma in uniseriate marginal bands and scanty paratracheal (in all spe­ eies of Group A except S.fasciculifoliolata) in solitary strands; in strands of 3-6 (total range 2-9) cells. Rays 3-1O/mm, uniseriate and multiseriate, 2-6 cells wide. Multiseriate rays com­ monly more than I mm high (up to 3 mm high in S. fasciculifoliolata), body ray cells procumbent, usually with 2-4 (up to 10 in S. fasciculifoliolata) marginal rows and complete or incomplete sheaths of upright and square cells. Uniseriate rays mostly with upright and square cells. Radial canals absent. Crystals absent. Note: Schefflera taiwaniana (Nakai) Kanehira (= Heptapleurum racemosum Bedd.) can also be included in Group A according to Kanehira's (1921) data.

Group B (Fig. 8-23; Table 4)

Diagnostic characters: vessel-ray and vessel-axial parenchyma pitting composed of solitary large pits with numerous smaller pits in the same cross field, radial canals present in multiseriate rays; helical thickenings absent; vessels usually narrow (diam­ eter less than 100 J..lI1l) and rather numerous (more than 40 per sq.mm); multiseriate rays usually with few sheath cells. Pith non-septate. Species from East Asia and Aus­ tralia. Material studied: : Australia, North Queensland, Atherton, USw 40193; St. Petersburg, Bot. Gard., cult., LEw 17623. - S. globulifera: Vietnam, Ha Nam Ninh, Cuc Phuong, Grushvitzky & N. Skvortsova 64 (1969) and 67 (1969). - S. glome­ rulata: Vietnam, Ha Nam Ninh, Cuc Phuong, LEw 3764. - S. leucantha: Vietnam, Son La, Moc Chau, LEw 3784. - S. octophylla: Vietnam, Vinh Phu, Tarn Dao, LEw 3790; Vietnam, Son La, road between Hoa Binh and Moc Chau, Grushvitzky & N. Skvortsova 139 (1969); Vietnam, Ha Bac, Hung Son, Grushvitzky & N. Skvortsova 184 (1969). - S. palmiformis: Vietnam, Son La, Chieng Yen, Grushvitzky & N. Skvortsova 91 (1969), type specimen. - S. pesavis: Vietnam, Ha Nam Ninh, Cuc Phuong, calcareous rocks, LEw 3768; Vietnam, Ha Son Binh, calcareous rocks, Grushvitzky & N. Skvortsova 149 (1969). - S. petelotii: Vietnam, Ha San Binh, Vu Ban, calcareous rocks, LEw 3769. - S. tomentosa: EIndia, LEw 10168. - S. tonkinensis: Vietnam, Ha Nam Ninh, LEw 3794. - S. trungii: Vietnam, Ha Giang, Grushvitzky & N. Skvortsova 139 (1963), paratype. - Scheffleropsis hemiepiphytica: Vietnam, Ha Son Binh, Tu Ly, Mt Mien, Grushvitzky & N. Skvortsova 178 (1969), type. - Tupidanthus calyptratus: Vietnam, Lai Chau, Takhtajan 8423, LEw 17621; St. Petersburg, Bot. Gard., cu1t., LEw 17622; Unknown locality, SFCw, s.n. Growth rings absent or indistinct, marked by 1-3 rows of radially flattened fibres or by vessel arrangement (S. trungii). Vessels rounded, rarely angular in outline, ranging from narrow (tangential diam­ eter less than 60 J..lI1l in S. tonkinensis) to rather wide (up to 120 J..lI1l and more in S. actinophylla, S. globulifera and T. calyptratus), solitary or in radial multiples of 3-5 (up to 16 in S. octophylla). Vessel frequency varies from 22 per sq.mm in S. actino­ phylla to 183 per sq.mm in S. tonkinensis. Vessel wal1s 2-9 J..lI1l thick. Tyloses ob­ served in vessels of S. actinophylla and Scheffleropsis hemiepiphytica. Average vessel element length 682-1117 J..lI1l (total range 200-1440 J..lI1l), the shortest vessel elements were found in S. tonkinensis, S. glomerulata, S. leucantha, and S. trungii, the longest

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Fig. 12-16. Perforation plates of Group B species, SEM. - 12-14. Scalariform perforation plates with numerous bars. - 12: Schejjlera pesavis. - 13: S. octophylla. - 14: Schejjleropsis hemiepiphytica. - 15. Schejjlera pesavis, perforation plate with few bars. - 16. S. trungii, simple perforation p1ate. - Fig. 17-19. Intervesscl pitting in Group B species, SEM. - 17: Tupidanthus calyptratus, scalariform pitting. -18: Schejjleropsis hemiepiphytica, transitiona1 pitting. - 19: Schejjlera octophylla, alternate pitting. - Scale bar of Fig. 12 & 17 = 20 fJI11: that ofFig. 13 = 100 fJI11: and that ofFig. 14, 16, 18, 19 = 10 fJI11: magnification ofFig. 15 as in Fig.14.

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Table 4. Wood anatomical features of Group B.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 Schejj1era actinophylla USw 40193 TR 1000 20/- +/- 1000 84 54/3 22 0/0-29 1216 4.4/2-7 0.6/1.3 0.3 2.6 LEw 17623 TL 8/3 767 65 42/6 68 0/0-13 968 3.4/2-5 0.5/1.0 0.4 3.0 S. xlobulifera Grushv.64 (1969) HE 350 18/4 +/- 827 57 43/5 62 0/0-22 1106 4.8/3-7 1.1/2.1 0.3 3.1 Grushv.67 (1969) HE 350 14/1 +/- 984 82 19/6 53 0/0-24 1310 3.2/2-5 1.3/2.8 0.9 3.7 S. xlomerulata LEw 3764 HE 200 7/4 724 68 36/5 45 0/0-19 828 2.6/2-4 0.5/1.1 0.4 3.6 S. leucantha LEw 3784 EP 1200 9/2 +/- 739 52 35/5 31 0/0-30 984 3.3/2-5 0.6/1.3 0.5 3.0 S. octophylla LEw 3790 TL 950 15/5 +/- 927 61 10/8 63 5/0-36 1159 3.9/2-7 0.6/1.3 0.8 4.1 Grushv. 139 (1969) TL 1100 17/4 - 1067 89 21/5 33 6/0-30 1263 3.6/2-5 0.7/1.7 0.9 3.2 Grushv. 184 (1969) TR 200 15/3 923 45 5/16 112 3/0-18 1076 4.1/2-6 0.6/0.9 1.5 3.9 S. palmiformis Grushv.91 (1969) TR 1000 16/4 - 932 61 18/9 52 6/0-42 1097 3.4/2-5 0.9/3.0 1.6 4.3 S. pesavis LEw 3768 SH 500 20/2 +/- 1117 51 31/4 48 5/0-33 1300 3.7/2-5 0.6/1.1 1.2 3.4 Grushv.149 (1969) SH 500 18/2 +/- 962 61 19/8 46 4/0-30 1235 4.0/2-6 0.6/0.8 1.3 3.2 S. petelotii LEw 3769 TR 200 20/2 933 38 34/5 43 0/0-42 1097 4.2/2-6 0.9/1.9 1.1 3.4 S. tomentosa LEw 10168 TL? - 10/4 +/- 820 47 28/7 93 0/0-28 1008 3.2/2-5 0.8/2.6 1.0 3.3 S. tonkinensis LEw 3794 TL 200 6/3 +/- 682 38 12/9 18 0/0-19 848 3.112-4 0.8/3.2 0.7 5.1 S. trungii Grusbv. 139 (1963) EP 200 8/3 +/- 778 52 44/4 82 0/0-5 921 3.5/2-6 0.7/1.3 0.9 4.4 Schejj1eropsis hemiepiphytica Grusbv. 178 (1969) HE 16/7 +/- 948 58 23/6 66 0/0-25 1136 3.6/2-5 0.7/1.8 0.3 3.7 Tupidanthus calyptratus LEw 17621 CL 1000 11/5 +/- 823 74 14/9 49 0/0-22 974 5.012-7 0.8/1.6 0.3 3.6 LEw 17622 CL 9/4 +/- 1009 58 23/7 58 4/0-30 1225 3.712-5 0.5/0.9 0.4 3.0 SFCw s.n. TL? 20/- +/- 1014 80 19/6 28 2/1-21 1393 5.9/2-10 1.3/2.7 0.1 2.9

1: Habit (TR = tree, TL = treeIet. SH = sbrub, EP = epipbyte, HE = bemi~epipbyte, CL = c\imber); für furtber 1egcnds, see Tab1e 3. in S. pesavis, S. octophylla, S. actinophylla, and T. calyptratus. Perforations usually scalariform with numerous (up to 42 in S. palmiformis) or few bars, occasionally sim­ ple (usually simple in S. trungii) and reticulate, in lateral and oblique end walls. Intervessel pits mostly scalariform to transitional (in S. octophylla, S. palmiformis, and S. trungii mainly alternate), 8-15 ~ in horizontal diameter, usually angular (some-

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" ...... ~ ,.~ .. r,~1I! ~ " ~ ~ ,..~ I ~ , ~ ~ 1ooi1 f, =~= ~~ ... ~ ~ =~'- X , ." t'\. ~ • ~ ... ~ t ) ~ ... t ... ~ ~24( ~jl ~'" -- .....Til.'1

Figures 20-25

Downloaded from Brill.com10/07/2021 09:20:01PM via free access Oskolski - Wood anatomy of Schefflera s.l. 171 times rounded or oval), with slit-like and lens-like apertures. Vessel-ray and vessel-axial parenchyma pitting composed of solitary large narrowly bordered pits and numerous smaller distinct1y bordered pits in the same cross-field. These small pits are similar to intervessel pits in size and shape, sometimes unilaterally compound. Helical thicken­ ings absent. Vascular tracheids are rare in S. globulifera, S. octophylla, and S. trungii, and absent in the other species. Libriform fibres very thin-walled in S. actinophylla, S. glomerulata, S. leucantha, S. palmiformis, S. tomentosa, S. hemiepiphytica, T. calyptratus, and thin- to thick-walled in others (walls 2-5 )lill thick, up to 8)lill in S. pesavis), septate, with simple to minute­ ly bordered pits with slit-like apertures. Axial parenchyma scanty paratracheal and narrowly vasicentric, forming narrow incomplete to complete sheaths; in strands of 5-10 (total range 2-17) cells. Rays 2-8/mm (most numerous in S. palmiformis, S. tonkinensis, and S. trungii), uniseriate and multiseriate, 2-5 (up to 7 in S. actinophylla, S. globulifera, S. octophylla, and T. calyptratus) cells wide. Ray height commonly over 1 mm in S. globulifera and S. palmiformis, in other species rays usually less high. Body ray cells of multiseriate rays procumbent; upright and square cells in marginal rows (usually 2-5 rows, but in T. calyptratus up to 10 rows), and in complete (more usually in S. octophylla and S. tonkinensis) or incomplete sheaths. Uniseriate rays composed of upright, square, and procumbent cells. Radial canals present in all Group B species except S. tonkinen­ sis. Crystals absent.

Notes: 1. SchejJlera blancoi Merr.(= CephaloschejJlera blancoi (Merr.) Harms) can also be included in Group B according to Rodriguez's (1957) data. 2. The infraspecific vessel frequency variation is very considerable in Group B spe­ eies, as is shown by its range in S. octophylla (from 33 to 112 per sq.mm). 3. Rodriguez (1957) reported a eonsiderably lower average vessel element length in S. octophylla (622 )lill) than was measured in the examined sampies of that spe­ eies (923 -1067 )lill). 4. The short vessel elements in S. glomerulata and S. tonkinensis as weIl as the absence of radial eanals in S. tonkinensis are probably caused by the juvenile nature of the examined sampies of these speeies (their stern diameter was less than 1 em). 5. Rodriguez (1957: 245-247) described the peculiar vessel-parenchyma pitting pattern in S. octophylla and S. blancoi.

Fig. 20-23. Vessel-axial parenehyma (20-22) and vessel-ray (23) pitting formed by solitary large pits and with numerous smaller pits in Group B speeies, SEM. - 20, 23: Tupidanthus calyptratus. - 21: Schejjlera octophylla. - 22: S. pesavis. - Fig. 24, 25. TS of Group C spe­ eies. - 24: S. vitiensis, growth rings absent, vessels solitary. - 25: S. pacoensis, growth ring boundaries distinet, marked by marginal parenehyma bands and also by differenees in vessel diameter between latewood and earlywood, vessels mainly in radial multiples. - Seale bar of Fig. 21 & 23 = 10 1l111; of Fig. 24 = 100 1l111; magnifieation of Fig. 20 & 22 as in Fig. 23; that of Fig. 25 as in Fig. 24.

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Figures 26-29

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Table 5. Wood anatomical features of Group C.

2 3 4 5 6 7 8 9 10 11 12 13 14 Group Cl S. hypoleuca Grushv.45 (1966) TL 1500 40/6 + 1195 47 8/21 193 5/1-17 1033 4.9/3-8 0.9/1.5 2.1 3.6 Grushv.46 (1966) TL 1500 23/14 + 926 41 10/8 122 4/0-18 1080 3.912-7 0.9/2.1 1.7 4.4 LEw 3763 TR 1150 20/7 + 1046 45 15/7 127 5/0-11 1140 3.8/2-6 0.9/1.7 1.2 4.2 S. pacoensis Grushv. 106 (1969) TR 1100 28/3 + 768 33 23/5 168 2/0-14 1018 4.312-7 0.7/1.4 1.8 4.3 Grushv. 124 (1969) TR 1500 19/6 + 920 33 6/8 234 2/0-6 1186 3.2/2-4 0.9/1.7 1.6 4.5 Grushv. 125 (1969) TR 1500 18/4 + 844 37 27/6 157 5/0-16 1174 3.9/2-6 0.8/1.6 1.4 3.9

Group C2 S. digitata USw 33492 TL 20/- + 402 44 6/12 98 0 644 12.113-19 1.7/3.7 - 1.9 S. euthytricha USw 13760 TR 16/3 874 63 28/6 36 20/0-38 1137 5.8/3-8 2.3/5.9 0.3 3.3 S. vitiensis USw 13610 TR 15/3 963 58 30/4 24 25/0-42 1263 5.7/2-9 2.1/4.4 0.2 3.7

Group C3 S. chinensis LEw 17670 TR 60/15 + 848 76 27110 48 3/0-27 1132 5.0/2-7 0.8/1.2 1.1 5.1 S. hypoleucoides Grushv.50 (1966) TR 1120 20/10 + 1093 53 5/14 119 912-25 1229 3.8/2-6 1.0/3.0 2.0 3.9

For legends, see Table 3.

Group C (Fig. 24-37; Table 5)

This group comprises Schefflera species from the Pacific region (except New Caledo- nian ones). They lack the most important diagnostic characters of Groups A and B such as helical thickenings on vessel and vascular tracheid walls, septate pith, or di- morphie vessel-parenchyma piuing). The species from Group C usually have rather narrow (diameter less than 100 /lffi) and numerous (more than 50 per sq.mm) vessels, and multiseriate rays with more or less numerous sheath cells; these characters, how-

Fig. 26 & 27. TS of Group espeeies, growth ring boundaries distinet. - 26: Schejj1era chinen­ sis, growth ring boundary marked by arrangement of vessels. - 27: S. digitata, note eonsiderable ditferenees in vessel diameter between latewood and earlywood. - Fig. 28 & 29. TLS. - 28: S. chinensis, multiseriate rays witb incomplete sheatbs of upright and square eells. - 29: S. di­ gitata, very wide multiseriate rays with numerous sheath cells. - Seale bar ofFig. 29 = 100/Jlll; magnification ofFig. 26-28 as in Fig. 29.

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Figures 30-37

Downloaded from Brill.com10/07/2021 09:20:01PM via free access Osko1ski - Wood anatomy of Schefflera s.l. 175 ever, are insufficient to clearly distinguish this assemblage. That is why Group C is subdivided into three small well-defined groups Cl, C2 and C3, which are described below.

Group Cl (Fig. 25, 35-37; Table 5)

Diagnostic characters: marginal parenchyma present, vessels narrow (diameter less than 60 /Jl11) and numerous (more than 100 per sq.mm), multiseriate rays with numer­ ous sheath cells. Species from East Asia.

Material studied: Schejj1era hypoleuca: Vietnam, Ha Tuyen, Pho Bang, Grushvitzky & N. Skvortsova 45 (1966) and 46 (1966); Vietnam, Ha Son Binh, Pa Co, LEw 3763. - S. pacoensis: Vietnam, Ha Son Binh, Pa Co, Grushvitzky & N. Skvortsova 106 (1969), holotype specimen; Vietnam, Son La, Moc Chau, Grushvitzky & N. Skvortsova 124 (1969) and 125 (1969), paratype specimens. Growth rings distinct, marked by I or 2 rows of radially flattened fibres, and by in­ complete, uniseriate marginal parenchyma bands, and also by differences in vessel diameter between the latewood and the earlywood (in S. pacoensis). Vessels angular, rarely rounded in outline, narrow (tangential diameter less than 60 /Jl11) and numerous (up to 234 per sq. mm in S. pacoensis). Vessels mostly in radial multiples of2-4 (up to 21 in S. hypoleuca). Vessel walls 2-8 /Jl11 thick. Tyloses present. Average vessel element length 768-1195 /Jl11 (total range 440-1420 /Jl11); vessel ele­ ments in S. pacoensis distinctly shorter than in S. hypoleuca. Perforations scalariform with few bars and simple, in lateral and oblique end walls. Intervessel pits alternate, occasionally opposite, 8-25 /Jl11 in horizontal diameter, rounded or oval, with slit-like and lens-like apertures. Vessel-ray and vessel-parenchyma pits half-bordered or with much reduced borders, similar to intervessel pits in size and shape, sometimes unilat­ erally compound. Helical thickenings absent. Vascular tracheids numerous in S. pacoensis, absent in S. hypoleuca. Libriform fibres very thin- to thin-walled (wall thickness 2-5 /Jl11), septate, with simple to minutely bordered pits with slit-like apertures. Axial parenchyma scanty diffuse and in uniseriate incomplete marginal bands, and also scanty paratracheal in solitary strands (in S. pacoensis); in strands of 5-8 (total range 4-10) cells. Rays 5-7/mm, uniseriate and multiseriate, 3-6 (up to 8 in S. hypoleuca) cells wide and commonly more than I mm (up to 2.1 mm in S. hypoleuca) high. Body cells of

Fig. 30-37. SEM ofGroup C species. - Fig. 30-32. Schejj1era euthytricha. - 30: Scalariform perforation plates with numerous bars. - 31: Scalariform intervessel pitting. - 32: Vessel-axial parenchyma pitting. - Fig. 33 & 34. S. digitata. - 33: Simple perforation plates. - 34: Scalari­ form intervessel pitting. - Fig. 35-37. S. hypoleuca. - 35: Perforation plate with five bars. - 36: Alternate intervessel pitting. - 37: Vessel-ray pitting. - Scale bar of Fig. 30 = 20 1J111; of Fig. 31-34 = 10 1J111; magnification of Fig. 35 as in Fig. 30; that of Fig. 36 as in Fig 31; that of Fig. 37 as in Fig. 32.

Downloaded from Brill.com10/07/2021 09:20:01PM via free access 176 IAWA Journal, Vol. 16 (2),1995 multiseriate rays procumbent; usually 3-5 (up to 8) marginal rows and complete (oc­ casionally incomplete) sheaths of upright and square marginal cells. Uniseriate rays usually composed of upright and square cells. Radial canals absent. Crystals absent.

Group C2 (Fig. 24, 27, 29-34; Table 5)

Diagnostic characters: narrow (diameter less than 90 !lJll) vessels, intervessel pitting exclusively scalariform, multiseriate rays very high (usually more than 2 mm high) with numerous sheath cells. Species from Fiji and New Zealand.

Material studied: Schefflera digitata: New Zealand, USw 33492. - S. euthytricha: Fiji, USw 13760. - S. vitiensis: Fiji, USw W-1361O. Growth rings absent in S. euthytricha and S. vitiensis and distinct in S. digitata; marked by 1 or 2 rows of radially flattened fibres and also by differences in vessel diameter between the latewood and the earlywood. Vessels angular, rarely rounded in outline, rather narrow (tangential diameter less than 90 !lJll). Vessel frequency varies from 24 per sq. mm in S. vitiensis to 98 per sq.mm in S. digitata (together with vascular tracheids). Vessels solitary or in radial multiples of 3-5 (up to 12 in S. digitata). Vessel walls 2-8 !lJll thick. Tyloses not found. Average vessel element length in S. euthytricha and S. vitiensis 874-963 !lJll (total range 420-1310 !lJll); in S. digitata vessel elements distinctly shorter (average 402 !lJll, range 288-572 !lJll). Perforations scalariform with numerous (up to 38 in S. euthytricha and 42 in S. vitiensis) and rarely few bars, or exclusively simple in S. digitata, in lateral and oblique end walls. Intervessel pits exclusively scalariform, 8-25 !lJll in horizontal diameter, rounded, with slit-like apertures. Vessel-ray and vessel-parenchyma pits half-bordered or with much reduced borders, similar to intervessel pits in size and shape. Helical thickenings absent. Vascular tracheids numerous in S. digitata, absent in the other species. Libriform fibres very thin- to thin-walled (wall thickness 2-5 Iilll, up to 9 !lJll in S. vitiensis), septate, with simple to minutely bordered pits with slit-like apertures. Axial parenchyma narrowly vasicentric in S. euthytricha and S. vitiensis, or scanty paratracheal in S. digitata; in strands of 4-10 (total range 2-17) cells. Rays 2-8/mm, uniseriate and multiseriate (in S. digitata multiseriate only), 4-8 (up to 19 in S. digitata) cells wide and commonly more than 1 mm (up to 5.8 mm in S. euthytricha) high. Body cells of multiseriate rays procumbent; usually 2-5 (up to 8) marginal rows and complete sheaths of upright and square marginal cells. Uniseriate rays in S. euthytricha and S. vitiensis mostly composed of upright and square cells. Radial canals present in S. euthytricha and S. vitiensis. Crystals absent.

Note: Meylan and Butterfieid (1978) reported that ray width in S. digitata is up to 25 cells, but such wide rays were not observed in our sampie. As far as I know, this spe­ eies has the widest rays among the Araliaceae. Moreover, this is the only Araliaceae species without uniseriate rays (Oskolski 1994).

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Group C3 (Fig. 26, 28; Table 5)

Diagnostic characters: vessels rather narrow (diameter less than 120 1Jlll) and numer­ ous (more than 40 per sq.mm), intervessel pitting scalariform to alternate, marginal parenchyma absent, multiseriate rays with incomplete sheaths. Species from East Asia.

Material studied: SchejJlera chinensis: China, Yunnan, LEw 17670. - S. hypoleucoides var. tomentosa: Vietnam, Ha Tuyen, Ha Giang, Grushvitzky & N. Skvortsova 50 (1966), isotype specimen. Growth rings distinct, marked by 1 or 2 rows of radially flattened fibres and also by differences in vessel diameter between the latewood and the earlywood. Vessels angular, rarely rounded (the latter more usual in S. chinensis), rather narrow (tangential diameter up to 120 lJlll in S. chinensis and up to 761Jlll in S. hypoleucoides). Vessel frequency varies from 48 in S. chinensis to 119 per sq.mm in S. hypoleucoides. Vessels in radial multiples of 3-6 (up to 14 in S. hypoleucoides), and solitary. Vessel walls 2-81Jlll thick. Tyloses present in S. hypoleucoides. Vessel elements in S. chinensis shorter (average 848 1Jlll, range 410-11 00 1Jlll) than in S. hypoleucoides (average 1093 1Jlll, range 580-1260 1Jlll). Perforations scalariform with few to numerous (up to 27 in S. chinensis) bars, occasionally simple (in S. chinensis) and reticulate (in S. hypo­ leucoides), in lateral and oblique end walls. Intervessel pits mostly scalariform, rarely transitional to alternate, 8-25 lJlll in horizontal diameter, rounded or oval (sometimes angular), with slit-like and lens-like apertures. Vessel-ray and vessel-axial parenchyma pits half-bordered or with much reduced borders, similar to intervessel pils in size and shape, sometimes unilaterally compound. Helical thickenings absent. Vascular tracheids absent. Libriform fibres very thin- to thin-walled (wall thickness 2-5 1Jlll), septate, with simple to minutely bordered pits with slit-like apertures. Axial parenchyma narrowly vasicentric, in strands of 4-6 (total range 3-8) cells. Rays c. 6/mm, uniseriate and multiseriate, 3-6 (up to 7 in S. chinensis) cells wide and commonly c. 1 mm (up to 3 mm in S. hypoleucoides) high. Body cells of multiseri­ ate rays procumbent; usually with 2-5 (up to 8) marginal rows and incomplete sheaths of upright and square cells. Uniseriate rays mostly composed of upright and square cells. Radial canals absent. Crystals absent.

Group D (Fig. 38-51; Table 6)

Group D is composed of the SchejJlera and Didymopanax species from Africa, South America and New Caledonia. These species have usually rather wide vessels (average diameter more than 70 1Jlll) in 10w densities (less than 30 per sq. mm); multiseriate rays with few or without sheath cells; non-septate pith; helical thickenings are absent on vessel and vascular tracheid walls. Group D can be subdivided into two well-defined groups Dl and D2, which are described below.

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jiiiC;,

~~""" ~~~ ...,.; I~ N 'i' ~ --.. ~:, , l

!I; ;.; \' ~ ,

i- ~ . ;.. , ~ l~ :lt, ,. .'

~~~ ... - ~

Fig. 38-41. TS ofGroup D species. - 38: Schefflera stuhlmannii, growth rings absent, wide and numerous vessels. - 39: S. fragrans, growth rings absent, vessels mostly solitary. - 40: Di­ dymopanax angustissimum, growth ring boundary marked by tlattened fibres. - 41: Schejjlera mildbraedii, growth ring boundary indistinct, marked by arrangement of vessels. - Scale bar of Fig. 41 = 100 /ill1; magnification of Fig. 38 -40 as in Fig. 41.

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Table 6. Wood anatomical features of Group D.

1 3 4 5 6 7 8 9 10 11 12 13 14

Group Dl Didymopanax angustissimum USw 25470 TR 50/- +/- 1366 83 24/6 26 2/0--9 1361 3.712-5 0.4/0.7 0.3 2.8 D. macrocarpum USw 35712 TR 16/3 +/- 745 74 27/5 23 0/0--7 948 2.6/2-4 0.6/1.5 0.6 4.4 D. morototoni SFCw S.ll. TR 30/- 961 80 91/4 9 4/0-9 1202 3.0/2-4 0.5/0.8 0.5 4.4 Schefflera europhylla USw 22198 20/- - 1260 70 36/6 16 1/0-17 1487 3.6/2-6 0.5/1.0 0.4 3.2 S·fragrans USw 33035 20/4 944 68 34/6 19 4/0-10 1137 2.5/2-3 0.2/0.5 1.1 4.1 S. gabriellae SFCw s.n. TR 30/- - 1339 85 21/6 20 4/1-20 1386 2.6/2-3 0.4/0.5 0.5 2.7 S. nono SFCw s.n. TL 20/- 961 107 11/7 17 0 1157 3.7/2-5 0.5/1.0 0.4 4.1 S. yurumanguinis USw 33420 20/- +/- 823 76 20/9 17 0/0--1 962 2.7/2-4 0.4/0.8 0.2 2.9

Group D2 Schefflera barteri Bw s.n. 18/5 817 82 63/3 20 1/0-4 955 2.8/2-4 0.3/0.5 0.1 2.4 S. mildbraedii Bw s.n. 24/2 +/- 904 114 11/7 27 0/0-6 1144 3.8/2-5 0.7/1.7 0.2 4.2 S. stuhlmannii Bw S.ll. 21/2 - 1124 158 45/5 23 0/0-3 1289 3.8/2-5 0.9/2.2 0.1 3.6 S. volkensii Bws.n. 20/- 910 110 54/3 16 3/0-7 1178 3.312-5 0.3/0.5 0.1 2.6

Note: Altitudes are unknown for all specimens of Group 0 and therefore column 2 is absent in this table. For further legends. see Table 3.

Group Dl (Fig. 39,40,42,44,45, 50, 51; Table 6)

Diagnostic characters: multiseriate rays low (usually less than 1 mm high, sometimes up to 1.5 mm high), with one or few marginal rows of square and upright cells (Kribs' type Heterogeneous IIb). Species from South America and New Caledonia.

Material studied: Didymopanax angustissimum: Brazil, Santa Catarina, USw 25470. - D. macrocarpum: Brazil, USw 35712. - D. morototoni: South America, SFCw s.n. - ScheJjlera europhylla: Ecuador, Pichincha, Saloya, USw 22198. - S. fragrans: Colombia, USw 33035. - S. gabriellae: New Caledonia, SFCw s.n. - S. nono: New Caledonia, SFCw s. n. - S. yuruman-guinis: Colombia, USw 33420.

Growth rings indistinct in D. angustissimum, D. macrocarpum, and S. yurumanguinis, and absent in the other species; if present, marked by a few rows of radially flattened latewood fibres. Vessels rounded, rarely angular in outline; rather wide (average tangential diameter 68-107 f.ID1, total range 28-148 f.ID1); mostly solitary in D. morototoni, in clusters of

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Figures 42-51

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5-11 in S. nono, in radial multiples of 3-6 (up to 9 in S. yurumanguinis) mixed with solitary vessels in other speeies; not numerous (vessel frequeney ranges from 9 per sq. mm in D. morototoni to 26 per sq. mm in D. angustissimum). Vessel walls 3-8 11111 thiek. Tyloses not found. Average vessel element length 745-1366 ~n (total range 400-1760 11111), the shortest vessel elements oeeur in D. macrocarpum and S. yuru­ manguinis, the longest ones in D. angustissimum and S. gabriellae. Perforations with rather numerous (up to 20 in S. gabriellae) or few bars, or simple (exclusively simple in S. nono, almost so S. yurumanguinis), and reticulate (in D. macrocarpum, D. moro­ totoni, S. fragrans, S. gabriellae); in lateral and oblique end walls. Intervessel pits transition al (mostly scalariform in S. europhylla; and mostly alternate in D. angustis­ simum, S. fragrans, S. gabriellae, S. yurumanguinis) or alternate in D. macrocarpum, D. morototoni, S. nono, 8-2511111 in horizontal diameter, rounded (mostly rounded in D. angustissimum and S. yurumanguinis) or angular, with slit-like and lens-like aper­ tures. Vessel-ray pits half-bordered (borders absent or indistinet in S. fragrans), simi­ lar to intervessel pits in size and shape (these differ from intervessel pits only in S. europhylla by rounded outline, and in S. nono by their scalariform shape), with slit­ like, oval and lens-like apertures. Unilaterally compound pits present in D. macrocar­ pum, S.fragrans, S. nono, and S. yurumanguinis. Vessel-axial parenchyma pits simllar to rounded and oval vessel-ray pits. Helical thickenings absent. Vascular tracheids absent. Libriform fibres very thin- to thin-walled (to thick-walled in D. angustissimum, and to very thick-walled in D. macrocarpum), walls 2-511111 thick (up to 911111 in D. macro­ carpum), septate (except in S. nano), with simple to minutely bordered pits with slit-like apertures. Axial parenchyma seanty paratracheal or narrowly vasicentric, in strands of 4-8 (total range 3-10) eells. Rays 2-6/mm, uniseriate and multiseriate, 2 or 3 cells wide in S. fragrans and S. gabriellae and wider in the other species (up to 6 cells wide in S. europhylla). Ray height commonly less than 1 mm. Body cells of multiseriate rays proeumbent, usually with 1 or 2 marginal rows of upright and square cells and solitary sheath cells (Kribs' type Heterogeneous IIb). Uniseriate rays mostly composed of proeumbent and square eells. Radial eanals present in D. angustissimum, D. morototoni, and S. europhylla. Crystals absent.

Fig. 42 & 43. TLS of Group D speeies, multiseriate rays without sheath eells. - 42: Didyma­ panax angustissimum, radial eanal (arrowed). - 43: Schefflera stuhlmannii, high multiseriate rays. - Fig. 44-51. SEM ofGroup D species. -Fig. 44 & 45. S. europhylla. -44: Scalariform perforation plate. - 45: Transitional intervessel pitting. - Fig. 46 & 47. S. stuhlmannii. - 46: Transitional intervessel pitting. - 47: Vessel-axial parenchyma pits with short grooves near apertures. - Fig. 48 & 49. S. barteri. - 48: Alternate intervessel pitting, small round pits. - 49: Round vessel-ray pits. - Fig. 50 & 51. S. nano. - 50: Alternate intervessel pitting. - 51: Coarse vessel-ray pits. - Scale bar of Fig. 42 & 44 = 100 /lill; of Fig. 46-48 & 51 = 10 /lill; mag­ nification of Fig. 43 as in Fig. 42; that of Fig. 45, 49, 50 as in Fig. 46.

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Notes: 1. According to data of Benoist (1931), Lindeman et al. (1963), Alves de Pinho (1966), Van der Slooten et al. (1970), and Mainieri and Chimelo (1989) Didymopanax calvum Decne. & Planch., D. navarroi A. Samp., D. vinosum Mart., D. pittieri March as weIl as Schefflera decaphylla (Seern.) Harms (= S. paraensis Huber) can also be included in Group D 1. 2. Indistinct growth rings were found also in S. decaphylla by Benoist (1931). 3. A low vessel frequency was noted also in D. pittieri (4-8 per sq. mm) by Van der Slooten et al. (1970), and in S. decaphylla (8-13 per sq.mm) by Benoist (1931).

Group D2 (Fig. 38,41,43,46-49; Table 6) Diagnostic characters: multiseriate rays sometimes more than 1.5 mm high, usually without marginal rows of square and upright cells (Kribs' type Homogeneous I). Species from Africa. Material studied: Schefflera barteri: Cameroon, Yaunde, Zenker 762, Bw s. n. - S. mildbraedii: East Africa, forest Rugege, Mildbraed 1010 (1907), Bw s. n. - S. stuhlman­ nii: Tanzania, East Usambara, Bw s. n. - S. volkensii: East Africa, Schlieben 272, Bw s. n. Growth rings absent or indistinct (in S. mildbraedii), marked by the vessel arrange­ ment. Vessels rounded, rarely angular, very wide in S. stuhlmannii (average tangential diameter 158 /lffi), narrower in the other species (82-114 /lffi), solitary and in radial multiples of 3-5 (up to 7 in S. mildbraedii); not numerous (vessel frequency ranges from 16 per sq. mm in S. volkensii to 27 per sq. mm in S. mildbraedii). Vessel walls 3-8 /lffi thick. Tyloses noted in S. stuhlmannii. Average vessel element length 817- 1124/lffi (total range 400-1760 /lffi), the longest vessel elements occur in S. stuhlmannii, the shortest ones in S. barteri. Perforations usually simple (S. mildbraedii, S. stuhl­ mannii) or scalariform with few bars (S. barteri, S. volkensii), in lateral or oblique end walls. Intervessel pits transitional (mostly scalariform in S. stuhlmannii and S. volkensii, mostly alternate in S. mildbraedii), and exc1usively alternate in S. barteri, 8-25 /lffi in horizontal diameter (6-9 /lffi in S. barteri), rounded or angular, with slit-like and lens-like apertures. Vessel-ray pits half-bordered (borders absent or indistinct in S. barteri), similar to intervessel pits in size and shape, with slit-like, oval and lens-like apertures (in S. stuhlmannii and rarely in S. mildbraedii slit-like apertures lie in short grooves by which they are sometimes coalescent). Unilaterally compound pits present. Vessel-axial parenchyma pits similar to the rounded and oval vessel-ray pits (in S. volkensii they differ from the latter through more reduced borders). HelicaI thicken­ ings absent. Vascular tracheids absent. Libriform fibres very thin- to thin-walled in S. mildbraedii, to thick-walled in other species; walls 2-5 /lffi (up to 7 /lffi) thick, septate, with simple to minutely bordered pits with slit-like apertures. Axial parenchyma narrowly vasicentric, in strands of 4-8 (total range 3-9) cells. Rays 2-5/mm, uniseriate and multiseriate, up to 5 cells wide. Ray height com­ monlyexceeds 1 mm in S. mildbraedii and S. stuhlmannii, in other species the rays are

Downloaded from Brill.com10/07/2021 09:20:01PM via free access Oskolski - Wood anatomy of Schefflera s.1. 183 lower. All cells of the multiseriate rays usually procumbent; square and upright cells usually absent (Kribs' type Romogeneous I); rarely there are 1 or 2 marginal rows of square and upright cells and solitary sheath cells. Uniseriate rays are mostly compos­ ed of procumbent cells. Radial canals present in S. stuhlmannii. Crystals absent.

DISCUSSION

Taxonomie and phylogenetie eonsiderations The results presented here enable the distinction of seven more or less well-defined groups of species. These groups can be subjected to systematic interpretation. There are not enough wood anatomical data, however, to precisely define the relationships of the whole set of studied Sehefflera species. As for the genera other than Sehefflera, Didymopanax is wood anatomically very similar to the South American and New Caledonian Sehefflera species (Group Dl), while Tupidanthus and Seheffleropsis are almost identical in their wood structure to the East Asian and Australian species of Group B. The taxonomie and phylogenetic interpretations of the species groups distin­ guished by their wood anatomy will be diseussed below. Group A coincides with section Agalma Tseng & Roo (but not with subsection Agalma from section Eusehefflera Rarms). The group is weIl defined by the presence of helieal vessel wall thickenings, vascular tracheids, marginal parenchyma (Table 7), opposite intervessel piUing, and also by a septate pith. Moreover, some species of this section, such as S. faseieulifoliolata, S. laxiuseula, S. vietnamensis and others inc1ud­ ed into subsection Glummea Grushv. & N. Skvorts., have very peculiar fasciculate­ compound leaves (Grushvitzky & Skvortsova 1969). This suggests an isolated status of section Agalma Tseng & Roo among the other infrageneric groups in Sehefflera and in a way supports Miquel's point of view that Agalma merits generic status (Miquel 1855; Rutehinson 1967). Rowever, Agalma can be conneeted with other Sehefflera species through such transitional forms as S. hypoleuea and S. paeoensis (see below). Group B consists of species belonging to seetion Brassaia Tseng & Roo, sub sec­ tions Oetophyllae Tseng & Roo and Heptapleurum Tseng & Roo (Table 1), as weIl as to the genera Seheffleropsis and Tupidanthus. This group is distinguished by its dimor­ phic vessel-ray and vessel-axial parenchyma pitting and the presence of radial canals (Table 7). There are considerable differences in the primitiveness of the wood struc­ ture in this group: the most primitive features (scalariform perforations with numerous bars, scalariform intervessel pitting) are found in the speeies of subsection Oetophyllae (S. petelotii, S. pesavis, etc.) and in Seheffleropsis hemiepiphytiea and Tupidanthus ealyptratus; the most advanced of these oceur in subsection Heptapleurum (S. trungii). It should be noted that, in spite of the differenees in fIower and infIorescence mor­ phology (Roo & Tseng 1965; Eyde & Tseng 1971; Grushvitzky & Skvortsova 1973) between Tupidanthus, Seheffleropsis and the above-mentioned Sehefflera groups (as weIl as between Sehefflera seetions Brassaia, Oetophyllae, and Heptapleurum), no wood anatomical boundaries were found between them. In my opinion this means that Group B, therefore, presumably eomprises related species. This interpretation does not agree with the views of the authors who consider Tupidanthus as the most primi-

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Table 7. Wood anatomical diversity in Schefflera and related taxa. More or less striking character associations are emboxed; + = character present; - = character absent; +/- = character variable and/or intermediate between two contrasting character states.

2 3 4 5 6 7 8 9 10 11 12 13 GroupA Schejflera delavayi + + + - - - S. jasciculifoliolata + + + + + S. laxiuscula + + + + + S. vietnamensis + + + + + + GroupB rn m S. actinophylla +/- +/- +/- + +/- + S. globulifera +/- + +/- + +/- +/- + S. glomerula ta +/- +/- +/- + +/- + S. leucantha + +/- + +/- + S. octophylla +/- +/- + + + S. palmiformis + +/- + +/- +/- + S. pesavis + +/- +/- + +/- + S. petelotii + +/- +/- + +/- + S. tomentosa + + +/- + +/- +/- + S. tonkinensis + + +/- + + + S. trungii + + + + +/- + Schlejfleropsis hemiepiphytica +/- + +/- + +/- + Tupidanthus calyptratus +/- +/- +/- + +/- +/- + Group Cl S. pacoensis - -- - S. hypoleuca ITJJ++ + ~+/- GJ Group C2 S. vitiensis - - - - - + S. euthytricha + - + S. digitata lJ-+ + + [TI + ITJ [TI Group C3 S. hypoleucoides + +/- - - S. chinensis QJ-+/- +/- +/- +/- GroupDl ~ S. europhylla +/- +/- + +/- + S.jragrans + S. yurumanguinis + + Didymopanax angustissimum + +/- + D. macrocarpum +/- D. morototoni + + S. gabriellae +/- + S. nono + +/- GroupD2 S. barteri - + - - S. volkensii -- +/- + S. stuhlmannii +/- + + S. mildbraedii [J + [ Characters: 1: Vessellumina narrow (diam. < 100 llffi) - 2: Vessels abundant (> 50 per sq.mm) - 3: Vascular tracheids abundant - 4: Scalariform intervessel pitting - 5: Vessel-ray and vessel-axial parenchyma pitting compos- ed of solitary large pits and with numerous smaller distinctly bordered pits in the same cross-field - 6: Mostly simple perforations (rarely with few bars) -7: Helical thickenings on the vessel walls - 8: Marginal parenchyrna- 9: Very high rnultiseriate rays (height > 3 mm) - 10: Low rnultiseriate rays (height < I rnrn) -11: Sheath cells in the multiseriate rays - 12: Hornogeneous rnultiseriate rays - 13: Radial canals.

Downloaded from Brill.com10/07/2021 09:20:01PM via free access Oskolski - Wood anatomy of Schefflera s.1. 185 tive genus in the Araliaceae (Harms 1894; Viguier 1906). Moreover, the wood ana­ tomical data support those taxonomists who include Schejjleropsis (Hutchinson 1967; Eyde & Tseng 1971) and Tupidanthus (Lowry et al. 1989) into Schejjlera. Both species of Group Cl, S. hypoleuca and S. pacoensis, belong to subsection Octophyllae. These species, however, sharply differ in their wood structure from other species in this subsection, and instead resemble species of Agalma in features such as numerous and narrow vessels, marginal parenchyma, and abundance of vascular tracheids (in S. pacoensis only). Unlike Agalma, both species are characterised by the absence of helical thickenings (Table 7) and by a non-septate pith. These species also resemble others in Group C (especially S. hypoleucoides) by their narrow and numer­ ous vessels and more or less numerous sheath cells in the multiseriate rays, but these relations are not as strong as with Group A. Thus S. hypoleuca and S. pacoensis could be interpreted as transitional forms connecting section (or genus?) Agalma with the type section of Schejjlera (Groups C2 and C3). Group C2 includes species with a very primitive wood structure (S. vitiensis and S. euthytricha, which possess perforations with numerous bars and scalariform inter­ vessel pitting) as well as those with more advanced wood characters (S. digitata, which has the shortest vessel elements, exclusively simple perforations, vascular tracheids, etc.). In spite of such a considerable difference in xylem specialisation level (in the Baileyan sense), S. digitata is closely connected with S. vitiensis and S. euthytricha by the shared occurrence of scalariform intervessel pitting (Table 7). It confirms Lowry's (1989) idea concerning the close resemblance of S. digitata with S. vitiensis, S. euthytricha and four other species from the southwest Pacific. It would be very inter­ esting to study the wood anatomy of all the members of this assemblage which com­ prise 'Schejjlera in the strictest sense'. The wood structure of S. hypoleucoides and S. chinensis (Group C3) superficially resembles that of Group CI and C2 species, but differs by the presence of transitio­ nal intervessel pitting and lower multiseriate rays with less numerous sheath cells (Table 7). Although S. hypoleucoides and S. chinensis belong to different sections of Hoo and Tseng's (1965) system (to the type subsection and section, and to subsection Cephaloschejjlera, section Brassaia, respectively, Table 2), Frodin (1975) maintains the close relationships of these two species with S. hypoleuca. Wood anatomy can confirm this point of view. All the other species studied (Group D) are characterised by narrow (usually 3-5 cells wide) multiseriate rays with few sheath and marginal cells or without them (Table 7). These species belong to the type subsection of Schejjlera and to the genus Didymopanax; they are widespread in South America, Africa, and New Caledonia. There are not enough wood anatomical data to describe exactly the relationships with­ in Group D. The most primitive wood structure (the longest vessel elements in the genus, scalariform perforations with numerous bars) is found in S. gabriellae from New Caledonia and in S. europhylla from Ecuador; the other species of Group D are more advanced. Distinct boundaries between Didymopanax and South American Schejjlera species (Group DI) were not found. This agrees with Maguire et al. (1984) and Cannon and

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Cannon (1989), who include Didymopanax in Schefflera. Nevertheless, the rank of Didymopanax should be defined more precisely. The African Schefflera species (Group D2) are distinguished rather sharply by the predominance of homocellular rays. These species differ considerably from one an­ other in their perforation type, and intervessel and vessel-parenchyma pitting. Thus wood anatomy cannot support Tennant's (1968) point of view on S. mildbraedii and S. stuhlmannii as synonyms of S. barteri. Among the studied Schefflera species, some are remarkable for their capituliferous and therefore belong to section Cephaloschefflera Harms: S. actinophylla, S. chinensis, S. volkensii, and also the 'intermediate' forms such as S. tomentosa and S. europhylla. Obviously, these species vary in their wood structure (Tablc 7) and do not form a natural group, which is in accordance with Frodin's (1975) view of section Cephaloschefflera Harms. As to Brassaia, regarded by Hoo and Tseng (1965) and Frodin (1975) as a section, the only species studied which belongs to this taxon, S. actinophylla, is closely related to subsections Octophyllae and Heptapleurum; i.e., a separate status of Brassaia can thus not be confirmed by the available wood anatomi­ cal data. Attention should also be paid to presence or absence of radial canals in multiseriate rays in Schefflera and so me other Araliaceae. In the species studied presence of canals tends to be associated with presence of perforation plates with numerous bars, and with scalariform intervessel pitting (Fig. 52); therefore, this character could be inter­ preted as an indicator of a low level of xylem specialisation. Distribution of this char-

% of species 70 A B

60

50

40

30

20 10 llil

O+-~L,~~~~'-~~---.~-,,---.----r~~~-L'-~, o 10 20 30 40 50 se mse ma a opp maximum bar number per perforation intervessel pitting type

Fig. 52. Frequency distribution (pereentages) of species with (blaek) and without (white) radial eanals as related to maximum number of bars per perforation (A), and intervessel pitting types (B). Pitting types: sc =exclusively sealariform; mse =mostly sealariform; ma =mostly alternate; a = exclusively alternate; opp = opposite.

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160 •

140 120 ,• j 100 \..; *l Ei o '6'"' 80 ••~ e· "'" .. ">- . 60 <=I •••O~ '"' o • E o 40 o • o 0 O· 00 o

20

0 0 50 100 150 200 250

vessel nurnber per sq. rnrn

Fig. 53. Vessel frequency plotted against mean vessel diameter. 0= samples collected at altitudes of 1000 m and more; • = sampIes collected at altitudes less than 1000 m, or altitude unknown.

acter within the Araliaceae (Oskolski 1994) as weH as the Anacardiaceae (Dong & Baas 1993) has been interpreted in a similar way. Unlike Anacardiaceae, however, presence of radial canals in the taxa under study is not a constant feature at the generic level, but it can be used for delimitation of some assemblages of species (such as Group B).

Ecological trends in the wood anatomy 0/ Schefflera and related taxa Since almost aH the species studied are distributed in the tropics and subtropics, the wood structural variation in them is more affected by the altitude than by the latitude of provenance. Unfortunately reliable and detailed data on altitude and latitude of prov­ enance for many wood sampies were not available. Therefore our analysis of ecologi­ cal trends cannot be very detailed, and only three broad ecological groups were distin­ guished. The first group comprises tropical species of different mainlands and islands of which the altitude of provenance is less than 1000 m or unknown. Evidently this group

Downloaded from Brill.com10/07/2021 09:20:01PM via free access 188 IAWA Journal, Vol. 16 (2),1995 is rather heterogeneous in relation to ecology, but there are no field data for a further subdivision. The second group includes the mountain species from North Vietnam collected at altitudes above 1000 m (S. delavayi, S. fasciculifoliolata, S. hypoleuca, S. hypoleucoides,S. laxiuscula, S. leucantha, S. pacoensis, S. palmiformis). These habi­ tats are characterised by subtropical (with an altitude up to 1200 m) and temperate (more than 1200 m) climates with cool and dry winters (Thai-van-Trung 1967). SchejjZera octophylla covers a wide altitudinal range: specimens of this species can be assigned either to the first [Grushv. 184 (1969)] or the second group [LEw 3790 and Grushv. 139 (1969)]. The third group consists only of S. digitata which is distrib­ uted in the lowland and the subalpine forests of New Zealand up to an elevation of c. 1100 m (WardIe 1962). Tt seems the harshest habitat where SchejjZera has been found. That is why this species is very interesting from the standpoint of ecological wood anatomy. Distinct growth rings marked by radially flattened fibres and marginal parenchyma are common in mountain species (except in S. leucantha, S. palmiformis, and moun­ tain specimens of S. octophylla and T. calyptratus) but also occur in several species from the lowland provenances (S. chinensis, S. mildbraedii, S. trungi, D. macrocarpum). Indistinct growth rings are found in many tropicallow land species. Perhaps, the growth ring formation in these species depends on endogenously determined rhythmic growth, as occurs in S. heterophylla (Koriba 1958) and Oreopanax sp. (Borchert 1969), where­ as in the mountains distinct growth rings are mostly caused by a seasonal c1imate (Thai-van-Trung 1967). The vessel elements tend to be more numerous and narrow in montane than in low­ land species (Fig. 53). Moreover, numerous vascular tracheids and helical thickenings occur in the mountain species only. This is in line with commonly established ecologi­ cal trends (Van der Graaff & Baas 1974; Carlquist 1988). However, other trends such as the shortening of vessel elements (Van der Graaff & Baas 1974) can be distin­ guished only by the comparison of all tropical species (both lowland and montane) with the temperate S. digitata which has conspicuously short vessel elements with exc1usively simple perforations. Variation of these characters within the tropical spe­ cies seems unrelated with habitat factors as far as can be established with the scanty field data.

ACKNOWLEDGEMENTS

This research was supported by the G. Soros Foundation 'Cultural Initiative' (Biodiversity pro­ ject). The author is grateful to Dr. A.F. Wilkins (SFCw), Dr. M. Hakki (Bw) and the late Dr. R. Eyde (USw) far sending wood sampies, to Dr. Nina T. Skvortsova and the late Prof. LV. Grushvitzky who have collected the very important wood material in North Vietnam, to Miss Mary Gregory and Dr. P. Gasson (Kew), who helped me to receive many papers absent in the Russian libraries, to Prof. Eugenia S. Chavchavadze far her useful consultations, and especially to Prof. P. Baas (Leiden) and to an anonymous referee for their very valuable and helpfu1 comments. I would also like to thank Ludmila A. Kartseva and Nina V. Tchentsova far carrying out the SEM observations, Ilya V. Outekhin, Ilya B. Kutcherov and Nicholas T. L. Harrocks far help with the English version ofthis text, and Tatiana B. Rebrieva, Elena S. Rebrieva and Svetlana B. Volkova far their assistance in the preparation of illustrations.

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