IAWA Bulletin n.s., Vol. 13 (4),1992: 419-453

WOOD ANATOMY OF TREES AND SIffiUBS FROM CHINA. IV. ULMACEAE

by

Y. Zhong!", P. Baas! and E.A. Wheeler 2

Swnmary The wood anatomy of 37 species belong­ ture to be more common in tropical dicotyle­ ing to the eight genera of Ulmaceae native to dons than in temperate ones, storied structure China is described. The wood of Chinese is more common in temperate wne Ulmaceae Ulmaceae is characterised by mostly simple than in tropical Ulmaceae. perforations (sporadic scalariform plates Key words: Ulmaceae, China, wood iden­ occur in Hemiptelea and Zelkova); altemate, tification, systematic wood anatomy, eco­ non-vestured intervessel pits; relatively short logical wood anatomy. vessel elements and fibres; non septate fibres with simple to minutely bordered pits con­ Introduction fined to the radial walls; mainly paratracheal The family Ulmaceae has 15 extant genera parenchyma; rays rarely higher than 1 mm. and about 200 species (Soepadrno 1977) of Tanniniferous tubes are reported for the first deciduous, semi-deciduous or evergreen trees time in Ulmaceae; they are limited to the or shrubs widely distributed in the temperate, genus Pteroceltis. Other, sporadically occur­ subtropical and tropical regions in both the ring features such as perforated ray and axial northern and southern hemispheres, and has parenchyma cells and perforated fibres are a fossil record extending back to the Late also reported for the first time. Cretaceous (Manchester 1989). Many of its Wood structure for the most part supports members are of economic interest for either the placement of genera within the groups their timber (e. g., Ulmus, Zelkova, Celtis), Celtidoideae and Ulmoideae suggested by their bark (e. g. Pteroceltis bark is used for Grudzinskaya (1967). Celtidoideae are char­ Xuan paper for traditional Chinese painting acterised by essentially heterocellular rays, and calligraphy), or as ornamental trees. Tra­ the Ulmoideae by homocellular rays. Wood ditionally, the family has been divided into anatomy supports the traditional placement of two subfamilies, the Ulmoideae and the Cel­ Gironniera cuspidata and G. yunnanensis in tidoideae, and recently many authors favour the genus Gironniera, and not in Aphananthe. treatment of the Celtidoideae as a separate The Ulmaceae, and in particular the Ulmoi­ family, the Celtidaceae, suggesting that the deae, should be considered specialised in their genera in the Celtidoideae are more similar to wood anatomy. the Moraceae than they are to the genera in Ecological trends in the wood anatomy of the Ulmoideae group (Grudzinskaya 1967; the Ulmaceae from China largely follow gen­ Chernik 1975, 1980, 1981, 1982; Oginuma eral trends established for the dicotyledons. et al. 1990; Takahashi 1989; Terabayashi The taxa from temperate provenances tend to 1991; Takaso & Tobe 1990). However, no have shorter vessel elements and fibres than cladistic analysis of these relationships has those from subtropical to tropical provenan­ been done, and there are genera with a mix­ ces. Tropical and subtropical samples are ture of ulmoid and celtoid features, e.g., predominantly diffuse-porous, while temper­ Ampelocera and Aphananthe (Giannasi 1978; ate samples are almost always ring-porous. Zavada 1983; Terabayashi 1991), Gironniera Counter to the general trend for storied struc- (Oginuma et al. 1990; Takaso & Tobe 1990).

1) Rijksherbarium / Hortus Botanicus, P.O. Box 9514, 2300 RA Leiden, The Netherlands. 2) Department of Wood and Paper Science, Box 8005, North Carolina State University, Raleigh, N.C. 27695-8005, U.S.A. *) Permanent address: Department of Forestry, Guangxi Forestry College, Nanning, China.

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Table 1. Numbers of species of the Chinese Ulmaceae.

Subfamily World China Number of species Genus (samples) studied Celtidoideae Aphananthe 8 (4-5) 1 1 (9) Celtis 70 (60) 10-20 10 (36) Gironniera 30 (6) 3 3 (6) Pteroceltis 1 1 1 (5) Trerna 50 (10-15) 6 (5) 4 (10) Ulmoideae Hemiptelea 1 1 1 (1) Ulmus 40 (18) 23 (21) 13 (26) Zelkova 10 (5) 4 4 (10) 1) Number of species for the whole world are based on How (1982) and Mabberley (1987, be­ tween brackets). 2) Number of species for China follow How (1982), the Iconographia Cormophytorum Sini­ corum, Supplement I (Institute & Botany Academia Sinica 1982), and other Chinese litera­ ture sources (Nanjing Forest College 1961 and Cheng et al. 1980).

In this study we use the traditional, broader China. Previous papers in the series were on concept of the Ulmaceae. the Oleaceae (Baas & Zhang 1986), Theaceae Eight genera of Ulmaceae occur in China. (Deng & Baas 1990), and Rosaceae (Zhang The approximate total number of species in & Baas 1992). China and in the world, and the number of species and samples studied here are listed in Material and Methods Table 1. More than 100 wood samples were obtain­ For general surveys of the earlier work ed from several institutional wood collections dealing with wood anatomy of Ulmaceae, see in China (see Acknowledgements). Unfortu­ De Bary (1884), Solereder (1899, 1908) Met­ nately, some samples are not vouchered, or calfe and Chalk (1950) and Sweitzer (1971). data on herbarium vouchers are not available. Sweitzer's study is the most comprehensive For nomenclature we largely follow Icono­ one. Additionally since 1950 wood anatomi­ graphia Cormophytorum Sinicorum (Institute cal data on a limited number of Ulmaceae of Botany, Academia Sinica 1972, 1982). For have appeared in numerous scattered publica­ each sample, the location is listed first, fol­ tions, of which the following are relevant to lowed by the wood collection number preced­ this study: Cheng et al. (1979, 1980, 1985), ed by a code from Stern's Index Xylariorum Chiang (1962, 1964), Desch (1954), Ho (1988), indicating the source of the sample. (1985), Luo (1989), Minaki et al. (1988), Light microscopic studies of sections and Ogata (1975-83), Saiki (1982), Sudo (1959, macerations were carried out in the usual 1963), Tang (1973), Wang (1965, 1966), manner (cf. Baas & Zhang 1986), followed Wheeler et at. (1989), Wu and Wang (1976), by SEM observations of selected samples. Wu et al. (1989), Yang and Huang-Yang Conventions for determining quantitative (1987), and Yao (1988). values and the terminology largely follow pre­ This study surveys the wood anatomy of vious studies (Baas & Zhang 1986; Baas et al. the genera of Ulmaceae native to China and 1988) and recommendations in the IAWA discusses its systematic and ecological signi­ List of Microscopic Features for Hardwood ficance. It is a contribution to the series on Identification (lAWA Committee 1989). the wood anatomy of trees and shrubs from (text continued on page 428)

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Legends to Figures 1-40 (pages 422-427): Figs. 1-4. TS, x 48. - 1: Trema orientalis (CAFw 7267), wood diffuse-porous and growth rings absent. - 2: Trema angustifolia (CAFw 11828), wood diffuse-porous with very vague growth ring boundary. - 3: Trema cannabina (CAFw 19820) wood semi-ring-porous. - 4: Aphananthe aspera (CAFw 17625), wood diffuse-porous; growth rings distinct; parenchyma aliform, vasicentric, and in marginal bands.

Figs. 5-8. TS, x 48. - 5: Ulmus tonkinensis (CAFw 146600), wood diffuse-porous and ves­ sels partly in small clusters. - 6: Celtis vandervoetiana (CAFw 17413), wood diffuse-porous and vessels partly in radial multiples. - 7: Celtis bungeana (CAFw 17072), wood ring-porous, earlywood vessels in 1 or 2 rows, latewood vessels in wavy tangential to diagonal bands, scle­ rotic tyloses present (arrow). - 8: Zelko va serrata (CAFw 16112), wood ring-porous, early­ wood vessels in single rows.

Fig. 9. Zelkova serrata , SEM, RLS, x 1950. Multiple perforation plate with two irregular bars. - Fig. 10. Trema virgata, SEM, RLS, x 1100. Spiral thickenings distinct in very narrow vessel ele­ ments and faint in wider vessel elements, simple perforation present in the narrow vessel element (arrow). - Fig. 1 L Celtis koraiensis, SEM, RLS, x 1650. Tylosis in the libriform fibre. - Fig. 12. Zelkova serrata, SEM, TLS, x 1950. Simple perforations and alternate intervessel pits. - Fig. 13. Celtis koraiensis, SEM, RLS, x 2500. Vessel with distinct spiral thickenings. - Fig. 14. Pteroceltis tatarinowii, TLS, x 1100. Very faint spiral thickenings on vessel wall. - Fig. 15. Celtis koraien­ sis, SEM, TS, x 1000. Latewood vessels intermixed with vascular tracheids and axial parenchy­ ma cells. - Fig. 16. Zelkova serrata, SEM, TLS, x 2500. Slit-like, partly coalescent apertures.­ Fig. 17. Trema virgata, SEM, RLS, x 3600. Unidentified granular deposits on the vessel wall.

Figs. 18 & 19. TLS, x 120. -18: Trema orientalis (CAFw 7267), alternate, polygonal interves­ sel pits. - 19: Pteroceltis tatarinowii (CAFw 13667), minute intervessel pits. - Figs. 20 & 21. RLS, x 300. - 20: Gironniera subaequalis (Guangxi For. ColI. w 8), vessel-ray pits simple to half-bordered, of varying size and shape. - 21: Celtis sinensis (CAFw 13083), vessel lumen filled with sclerotic, partly crystalliferous, tyloses. - Figs. 22 & 23. TLS, x 120. - 22: Celtis philippensis, thin-walled tyloses. - 23: Ulmus pumila (CAFw 13822), vascular tracheids inter­ grading with narrow vessel elements; note also the storied arrangement of narrow vessel elements, vascular tracheids and axial parenchyma, and two distinct ray sizes (narrow rays arrowed).

Figs. 24-26. Ulmus pumila (CAFw 13822), simple perforations in libriform fibres (arrows).- 24: TLS, x 300; 25: RLS, x 614; 26: RLS, x 455. - Fig. 27. Ulmus pumila, TLS, x 300. Simple perforation in axial parenchyma cell (arrow). - Fig. 28. Zelkova serrata (HEFw 001400), RLS, x 614. Perforated ray cell, much smaller than other ray cells. - Fig. 29. Pteroceltis tatarinowii (CAFw 16386), TLS, x 818. Simple perforations in tangential wall ofray cells. The perforated ray cells are as large as other ray cells.- Fig. 30. Celtis koraiensis (CAFw 549) TLS, x 120. Sheath cells (arrow) and storied arrangement of vessel elements, vascular tracheids, axial parenchyma and small rays. - Fig. 31. Ulmus tonkinensis (CAFw 6813), RLS, x 120. Homocellularray and cham­ bered crystalliferous axial parenchyma. - Fig. 32. Celtis wightii var. consimilis (CAFw 6795), RLS, x 182. Rays composed of upright cells only, crystal-containing cells non-chambered.

Fig. 33. Zelko va sinica (CAFw 5503), RLS, x 300. Integumented prismatic crystal. - Fig. 34. Pteroceltis tatarinowii (CAFw 13667), RLS, x 120. Irregularly shaped tanniniferous ray cells. - Fig. 35. Pteroceltis tatariflOwii (CAFw 16386), TLS, x 120. Storied arrangement of vessel ele­ ments, rays and axial parenchyma. - Fig. 36. Gironniera yunnanensis (CAFw 11934), TS, x 120. Vitreous silica in the vessel. - Fig. 37. Gironniera subaequalis (Guangxi For. Coli. w 8), TLS, x 120. Vitreous silica in fibres. - Fig. 38. Gironniera cuspidata (CAFw 12148), RLS, x 120. Unidentified granules in heterocellular ray. - Figs. 39 & 40. Pteroceltis tatarinowii (CAFw 16386), tanniniferous tubes in rays. - 39: TS, x 300; 40: RLS, x 180.

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Survey of wood anatomical character Vessel arrangement and groupings are of states in the Ulmaceae from China substantial diagnostic and systematic value at and above the genus level. Type 1 charac­ Growth rings (Figs. 1-8) terises all diffuse-porous species and semi­ Growth rings vary considerably in their ring-porous species. Type 2 characterises all delimitation from distinct to faint or absent. ring-porous Ulmaceae; this distinct pattern Growth ring boundaries can be of the fol­ is always associated with vascular tracheids lowing types: and narrow elements that intergrade between 1. In diffuse-porous species: tracheids and normal vessel elements, spi­ a) Growth rings absent to faint and mark­ ral vessel wall thickenings, locally inflated ed by differences in fibre wall thickness multiseriate rays, and marginal parenchyma (Figs. 1 & 2). bands. b) Growth rings distinct, marked by radi­ Frequency and element size - Average ally flattened fibres (Fig. 3). vessel frequency ranges from 6 to 50/mm2 c) Marked by marginal parenchyma and ir­ in semi-ring- to diffuse-porous samples. In regular zonate parenchyma bands (then ring-porous species vessels are of very high boundaries faint), with or without ad­ frequency in the latewood and it is difficult to ditional differences in fibre diameter distinguish between narrow vessels and vas­ (Figs. 4-6). cular tracheids. Consequently, we did not II. In ring-porous species: measure frequency or tangential diameter of Marked mainly by differences in vessel latewood vessels of ring-porous woods. diameter, and also by marginal or seem­ Average tangential vessel diameter (including ingly marginal parenchyma, and differ­ the wall) of semi-ring- to diffuse-porous Ul­ ences in radial fibre diameter and wall maceae in China ranges from 70 to 160 ~m; thickness (Figs. 7 & 8), rays often in­ in ring-porous Ulmaceae the average tangen­ flated at the boundaries. tial diameter of early wood vessels ranges from 100 to 250 ~m. Average vessel element Vessels lengths of semi-ring- to diffuse-porous Ulma­ Porosity - As noted above, diffuse-por­ ceae range from 210 to 720 ~m; average ves­ osity, semi-ring-porosity, as well as ring­ sel element lengths ofring-porous Ulmaceae porosity occur in the Ulmaceae. With only a range from 190 to 380 ~m. In some instances, few exceptions, ring-porosity is confined to quantitative characters can be useful in dis­ extra-tropical Ulmaceae, and all ring-porous tinguishing individual genera or infrageneric Chinese Ulmaceae are deciduous. Semi-ring­ taxa. For instance, vessel elements are the to diffuse-porosity occurs in evergreen as longest in Gironniera (on average 700-720 well as in deciduous species. ~m), followed by Trema (on average 400- Although porosity can be of great diagnos­ 510 ~m), while in the other genera the aver­ tic value, semi-ring-porosity is not always a age vessel element length ranges from 190- constant feature at the generic level and some­ 380 ~m. times varies within a species or from ring to Perforations (Figs. 9-10, 12) - Almost ring within a single specimen. all Chinese Ulmaceae have exclusively simple Arrangement and groupings (Figs. 1-8) perforations (Fig. 12). Only in Hemiptelea and - There are two patterns of vessel arrange­ Zelkova scalariform plates with 1 or 2 bares) ment and grouping within the Chinese Ulma­ (Fig. 9) and simple perforations with ceae: Type 1) Vessels mainly or at least partly vestigial bars very rarely occur. Simple per­ solitary, with the remainder mainly in radial forations are also found on side walls of some multiples of 2-4(-9), or in clusters of 3-4 vessel elements (Fig. 10) in some species. (-11) (Figs. 1-6). Type 2) Vessels in the The inclination of the perforation plates latewood forming a more or less wavy tan­ varies from highly oblique to horizontal, large­ gential to diagonal pattern as in Hemiptelea, ly depending on the diameter of the vessels, Zelkova, and ring-porous species of Celtis the narrower vessel elements having more and Ulmus (Figs. 7 & 8). oblique perforations.

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Wall pitting (Figs. 12, 16, 18-20) - In­ Tyloses and deposits (Figs. 17,21,22)­ tervessel pits in the Ulmaceae are non-vestured Tyloses are common to rare or absent in Chi­ and alternate (Fig. 12). Pit shape is polygonal nese Ulmaceae; tyloses were observed in all and/or round (Figs. 12, 18). The apertures genera studied but not as a constant feature. are usually slit-like, frequently to occasionally They are mainly thin-walled (Fig. 22), but in coalescent (Fig. 16) and/or oval to round. some species sclerotic tyloses are common as Intervessel pit size ranges from 3-10 (2-20) well (Fig. 21). Dark deposits (gum-like) are j.Lm. Intervessel pits are minute in Pteroceltis of common occurrence in some species. Irreg­ (3-4 j.Lm, Fig. 19), and minute to small (3-7 ular unidentified deposits on the vessel walls j.Lm) in Aphananthe and diffuse-porous Celtis. were observed in Trema (Fig. 17). All these species with minute to small inter­ Vascular tracheids (Figs. 15,23) vessel pits are diffuse-porous, and mostly The presence of vascular tracheids is a from subtropical to tropical regions. characteristic feature of all ring-porous Ulma­ Vessel-ray and vessel-parenchyma pits ceae with a wavy tangential to diagonal pat­ range from pits with much reduced borders tern. Vascular tracheids are here associated to simple, or half-bordered pits. In Aphanan­ with vessel groups and completely intergrade the, Pteroceltis, Celtis, Hemiptelea, Ulmus, with normal narrow vessel elements (Fig. 15, and Zelkova they are usually similar to or 23). Frequency of vascular tracheids varies smaller than the intervessel pits, rounded (at from abundant to relatively rare; the latter sit­ times elongate in Celtis, Hemiptelea and Zel­ uation prevails in a few specimens where the kova), sometimes unilaterally compound; in wood is sometimes semi-ring-porous and the Gironniera and Trema they are of varying size tangential vessel distribution pattern is not and shape: small to large, round to angular or very pronounced, and in the diffuse-porous elongate, sometimes unilaterally compound species of Ulmus. (Fig. 20). Vascular tracheids are absent in semi-ring­ Wall thickness and sculpturing (Figs. 10, to diffuse-porous Celtis, and of rare occur­ 13, 14) - Vessel walls range from 1-6(-10) rence in diffuse-porous Ulmus. As mentioned j.Lm in thickness for the whole family: 4-7 j.Lffi before, presence of vascular tracheids is ap­ in Aphananthe and Pteroceltis; 2-5 j.Lm in parently closely related to vessel grouping; in Gironniera and Trema; 2-4 j.Lffi in Hemiptelea semi-ring- to diffuse-porous Celtis vessels are and Ulmus; 1-5(-10) j.Lm in Celtis and Zel­ mainly in short radial multiples and vascular kova. tracheids are absent, whilst in diffuse-porous Spiral thickenings are virtually absent in Ulmus some vessels occur in clusters and vas­ Gironniera and Trema, being very faint and cular tracheids are present. only visible with SEM in G. yunnanensis and T. angustifolia. They vary from fine Fibres (Figs. 2, 11, 24-26) (Aphananthe, Pteroceltis) to distinct (Celtis, The ground tissue fibres are typically non­ Hemiptelea, Zelkova, and Ulmus) (Figs. 10, septate libriform fibres, with simple to minute­ 13, 14); and from present in all vessels (Apha­ ly bordered pits (borders smaller than 3 j.Lm) nanthe, Pteroceltis, Celtis austrosinensis, Ul­ mainly confined to the radial walls. Fibre mus changii, and U. lanceaefolia) to confined wall thickness varies from very thin to thin to narrow vessels only. It should be stressed (Aphananthe, Gironniera and Trema), or thin that presence or absence of spiral thickenings to thick, or thick to very thick in other genera. is related to ecological factors. For instance, Average fibre length ranges from 690-1720 they are virtually absent in south subtropical (330-2070) j.Lm (short to moderately long). to tropical diffuse-porous Ulmaceae, but are The systematic value at the genus level of well-developed in extra-tropical Ulmaceae fibre wall thickness or fibre length is limited. with only few exceptions. Ring-porous Ul­ Two peculiar fibre features of aberrant maceae with a wavy tangential to diagonal rather than regular occurrence deserve men­ vessel pattern in the latewood invariably have tioning: Simple perforations in fibre walls well-developed spiral thickenings in latewood (Figs. 24-26) have been observed in almost vessel elements. all genera studied except Aphananthe. These

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perforations are probably the result of 'trans­ cumbent, square and/or upright cells mixed piercing' tip growth of adjoining fibres (Vidal throughout the ray. Gomes, Curitiba, Brazil, unpublished data). Ray frequency ranges from 3-13(-17)/ In a few species of Celtis some fibres have mm, most commonly 4-8 or 8-10(-12)/ ty loses (Fig. 11). mm. In almost all semi-ring- to diffuse-por­ ous Ulmaceae (except C. austrosinensis and Axial parenchyma (Figs. 1-8, 13, 27) C. vandervoetiana) the ranges of ray frequen­ Parenchyma ranges from scanty (in Trema cy are very similar and tend to be higher than and subg. Gironniera) to abundant. The fol­ in ring-porous Ulmaceae. lowing types of parenchyma distribution oc­ Ray width ranges from 1-13 cells, most cur in the Ulmaceae: 1) apotracheally diffuse; commonly 1-3(-5) cells in semi-ring- to dif­ 2) scanty paratracheal; 3) vasicentric; 4) ali­ fuse-porous Ulmaceae or 4-6(-8) cells in form; 5) confluent to confluent banded; 6) ring-porous Ulmaceae. In Pteroceltis, Apha­ wavy tangential to diagonal paratracheal nanthe, Celtis and other ring-porous Ulmaceae bands, intermixed with the vascular tra­ rays tend to be of two more or less distinct cheids; 7) marginal bands; 8) apotracheal zo­ sizes (Figs. 23, 24, 30). Ray height varies nate bands or lines. All genera have at least from 0.03-3.8 mm, most commonly 0.03- two of the above types. Apotracheally diffuse 0.8 mm, and maximum ray height rarely and scanty paratracheal parenchyma is typical exceeds 1 mm (some species of Gironniera, for Trema (Figs. 1-3). In subg. Gironniera Trema, Celtis, Hemiptelea, Ulmus and Zel­ (G. subaequalis) types 1-3 are present. In the kova). other genera including subg. Galumpita of Sheath cells are of common occurrence in Gironniera types 3-7 and sometimes type 8 Celtis, usually well-developed in ring-porous are present (Figs. 4-8). Celtis (Fig. 30), but weakly differentiated to Parenchyma strand length varies from 2-8 absent in semi-ring- to diffuse-porous Celtis. (-24) cells. Gironniera and Trema have nearly In Gironniera and Trema weakly differentiated the same range of parenchyma strand lengths sheath cells sometimes occur. (mostly 3-8, total range 2-12). In Celtis, Perforated ray cells occasionally occur in Ulmus and Zelko va parenchyma strands are Celtis, Hemiptelea, Ulmus andZelkova. Sim­ most commonly 2-4 cells long. ple ray perforations are usually in radial walls Simple perforations in axial parenchyma (Fig. 28), but also occur in tangential walls cell walls have been noted in some species or (Fig. 29). There are two kinds of perforated samples of Celtis, Ulmus and Zelkova (Fig. ray cells in Chinese Ulmaceae; one kind, as 27). occurs in Celtis, is radially enlarged, similar Rays (Figs. 23, 24, 28-32, 38, 40) to those reported in the literature for other Multiseriate rays in Ulmus are homocel­ families; the other kind is unusual in that per­ lular (composed of procumbent cells) only forated ray cells are not enlarged or even (Figs. 23, 31). In Hemiptelea and Zelkova, much smaller (in Zelkova) than normal ray both heterocellular and homocellular rays may cells (Fig. 28). Perforated ray cells have not occur in the same specimen. In the other spe­ been reported before for the family. cies studied only heterocellular rays occur. Disjunctive ray parenchyma cell walls are Heterocellular rays in the Ulmaceae are com­ of common occurrence. posed of procumbent body ray cells and 1 or 2-4(-8) rows of upright and/or square mar­ Storied structure (Figs. 23, 30, 35) ginal cells (Figs. 38, 40). Rays in Gironniera Rays and axial elements are irregularly are markedly heterocellular, rays with more storied in Pteroceltis (Fig. 35). Narrow ves­ than 4 rows of upright/ square marginal cells sel elements, vascular tracheids and axial are common. In Gironniera and Trema some parenchyma are sometimes locally storied multiseriate rays are composed entirely of in Ulmus, Zelko va, Hemiptelea, and ring­ upright and/or square cells. Uniseriate rays porous Celtis (Figs. 23, 30). Storied structure are often composed of only upright and/or is absent in Gironnierra, Trema, and Apha­ square cells (Fig. 32), sometimes with pro- nanthe.

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Tubes (Figs. 39, 40) 6-9 (2-13)/sq.mm, 59-74 (41-88)% soli­ Tanniniferous tubes occur in Pteroce/tis, tary, remainder mainly in radial multiples of extending radially among rays (Figs. 39,40), 2-4, rarely in clusters of 3-4 and oblique and are weakly differentiated to well devel­ pairs, oval or rarely round to weakly angular, oped. They occur together and partly inter­ tangential diameter 120-160 (70-205) ~m, grade with irregularly shaped tanniniferous radial diameter up to 270 ~m, walls 4-7 ~m cells (Fig. 34). This is the first report of tan­ thick. Vessel element length 240-320 (150- niniferous tubes in the Ulmaceae. 400) ~m. Perforations simple in slightly ob­ lique to horizontal end walls. Intervessel pits Mineral inclusions (Figs. 21, 31-33, 36-38) non vestured, alternate, mainly polygonal, 4-6 In our material crystals are absent in Giron­ ~m in diameter, with slit-like, occasionally niera, Hemiptelea and Trema, and are ofvari­ coalescent apertures. Vessel-ray and vessel­ able occurrence in the other genera. Druses axial parenchyma pits half-bordered or with only occur in some samples of some species much reduced borders to simple, similar to of Celtis. When present, prismatic crystals intervessel pits in size, sometimes unilater­ are abundant to infrequent, integumented ally compound. Fine (visible with SEM), (Fig. 33) or not, usually in non-chambered closely spaced spiral thickenings present (sometimes chambered in Celtis) square and/ throughout vessel elements. Thin-walled ty­ or upright ray cells (Fig. 32), less common­ loses rarely present in CAFw 12664. ly in procumbent ray cells, and/or chamber­ Vascular tracheids absent. ed or non-chambered axial parenchyma cells Libriform fibres 1090-1290 (830-1520) (Fig. 31), and occasionally in sclerotic tyloses ~m long, very thin- to thin-walled, with sim­ of C. sinensis (CAFw 13083) (Fig. 21). Crys­ ple to minutely bordered pits (1-2 ~m in diam­ tal-containing cells are enlarged or not, gen­ eter) mainly confined to the radial walls. Ge­ erally with one, sometimes with 2 or more latinous fibres common. crysta1(s) per cell or chamber. Axial parenchyma abundant, mainly ali­ Vitreous silica occurs in Gironniera and form to confluent and confluent-banded, with Ulmus lanceaefolia, mostly in vessels, some­ the bands more common in the latter part of times also in fibres, rarely in axial parenchy­ the growth ring. Also vasicentric (CAFw ma cells (Figs. 36, 37). 17625) and in 2-4-seriate marginal bands, in Unbleachable, unidentified granular depo­ strands of 2-8 cells. sits have been noted only in Gironniera, Rays 7-IO/mm, 1-5(-6) cells wide, in common in both upright cells and procum­ CAFw 17471 rays tend to be of two sizes: bent body ray cells (Fig. 38), occasionally in 1-2-seriate and 4-5(-6)-seriate, 0.1-0.5 axial parenchyma cells. (-0.8) mm high. Multiseriate rays heterocel­ lular, composed of procumbent body cells and Generic wood anatomical descriptions 1 (2-3) row of upright marginal cells, uni­ seriate rays with procumbent, square and up­ Subfamily: Celtidoideae right cells mixed throughout the ray or com­ Aphananthe Planch. (Fig. 4) posed of square and/or upright cells only. Material studied: A. aspera Planch.: Gui­ Disjunctive ray parenchyma cell walls occa­ zhou: CAFw 17625; Jiangxi: CAFw 17471; sionally present. Fujian: CAFw 12664; Guangdong: CAFw Crystals common, prismatic, mostly in up­ 9187; Zhejiang: CAFw 386; Guangxi: CAFw right ray cells, less common in procumbent 15878; Jiangsu: CAFw 6987; Hunan: CAFw body ray cells, absent in axial parenchyma 9418; Anhui: HEFw 000567. cells, integumented or not, usually one (rare­ Deciduous trees from temperate to north ly two) crystal(s) per non-enlarged and non­ tropical regions. chambered ray cell. Silica absent.

Growth rings distinct, marked by marginal Note: Sweitzer (1971) described "uniseri­ parenchyma bands and differences in fibre ate rays homocellular comprised of procum­ wall thickness. Wood diffuse-porous, vessels bent cells" in Japanese A. aspera, but in our

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study, as well as in the studies of Kanehira /lm. Perforations simple in horizontal to ob­ (1921b), Sudo (1959), Cheng et al. (1980), lique end walls. Intervessel pits nonvestured, Yao (1988) etc., homocellular uniseriate rays alternate, mainly polygonal, only round in C. were not observed. philippensis, 5-10 (3-12) /lm in diameter, with slit-like (sometimes round to oval), often Celtis L. (Figs. 6, 7,11,13,15,21,22,30, coalescent apertures. Vessel-ray and vessel­ 32; Table 2) axial parenchyma pits mostly simple or with reduced borders, similar to or smaller than Group I - Ring-porous species of Celtis intervessel pits in size, rounded, and horizon­ Material studied: C. biondii Pamp.: Guang­ tally or at times vertically elongate in some dong: CAFw 16906; Anhui: HEFw 003164; species, sometimes unilaterally compound. Guizhou: CAFw 20264; Jiangxi: CAFw Spiral thickenings present, distinct in narrow 17407; Guangxi: Guangxi Forestry College, vessel elements, faint to absent in wide vessel w 9; Fujian: CAFw 19819. - C. bungeana elements. Both thin-walled and sclerotic Blume: Shaanxi: CAFw 556; Guangdong: tyloses present in most samples (see Table 2). CAFw 17072; Henan: CAFw 5511; Sichuan: Vascular tracheids, intergrading with nar­ CAFw 8724; Hebei: CAFw 5537; Zhejiang: row vessel elements, abundant in late wood CAFw 7047; Jiangxi: CAFw 18124; Guizhou: (only rare in one semi-ring-porous sample of CAFw 20263; Shanxi: HEF 82cw 591. - C. C. biondii, CAFw 20264), associated with koraiensis Nakai: Shaanxi: CAFw 549. - C. the vessel-axial parenchyma groups, with macrocarpa Chun.: Guangxi: Guangxi For­ distinct spiral thickenings. estry College, w 5. - C. philippensis Blanco: Libriform fibres 890-1410 (620-1750) Guangxi: CAFw 16043; Guangxi Forestry /lm long; thin- to thick-walled, or thick- to College, w 10. - C. sinensis Pers.: Fujian: very thick-walled, with simple to minutely CAFw 13083; Jiangshu: CAFw 6663; Shan­ bordered pits confined to the radial walls. dong: CAFw 5709; Zhejiang: CAFw 391; Simple perforations occasionally occur in C. Henan: CAFw 5628; Taiwan: CAFw 7214; bungeana (CAFw 17072) and C. koraiensis. Hunan: CAFw 9415; Anhui: HEFw 000325; Tyloses rarely observed, but present in C. Guangxi: Guangxi Forestry College, w 2; koraiensis and C. sinensis (CAFw 13083). Guangdong: CAFw 16893; Sichuan: CAFw Axial parenchyma abundant, mainly para­ 17552; Jiangxi: CAFw 19157. - C. yunnan­ tracheal: vasicentric, confluent, aliform, and ensis Schneid.: Yunnan: CAFw 11301. in narrow (1-3-seriate) to broad (4-7-seri­ Deciduous trees or rarely shrubs from ate) wavy tangential to diagonal paratracheal temperate to tropical regions. bands, intermixed with the vascular tracheids; Growth rings distinct, marked by differ­ and in marginal (including seemingly mar­ ences in vessel diameter, and mostly by mar­ ginal) 1-2-seriate bands (see Table 2), in ginal or seemingly marginal parenchyma strands of 2-4(-10) cells. Simple perfora­ bands and locally inflated multi seriate rays. tions occasionally occur in tangential walls of Wood ring-porous, semi-ring-porous in some some species. Storied arrangement of narrow rings in certain samples (see Table 2); early­ vessel elements, vascular tracheids and paren­ wood pores in 1-3 continuous rows (only chyma very rarely occurs locally and in but a discontinuous in C. biondii, CAFwI6906), few samples. oval to round, mean tangential diameter of Rays 4-12/mm, usually of two sizes: 1-2 the solitary earlywood vessels 90-200 (60- (-4)-seriate and 5-8(-12)-seriate, 0.06-0.8 280) JUll, radial diameter up to 300 /lm; early­ (0.03-1.3) mm high. Rays mostly heterocel­ wood to latewood transition generally abrupt, lular, composed of procumbent body ray cells but more gradual in C. biondii (CAFw and 1-2(-5) rows of square to upright mar­ 20264), C. bungeana (CAFw 556) and C. ginal cells, also homocellular (all ray cells sinensis (CAFw 5709), latewood vessels pre­ procumbent) in C. koraiensis; some uniseri­ dominantly in clusters, in wavy tangential to ate rays composed of square and/or upright diagonal bands, walls 1-5(-9) /lm thick. cells, generally low (rarely over 10 cells high). Vessel element length 220-380 (160-470) Sheath cells usually present (see Table 2).

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Disjunctive ray parenchyma cell walls present austrosinensis. Spiral thickenings present, in all species. Perforated ray cells occasion­ distinct in narrow vessels and usually faint to ally occur (see Table 2). absent in wide vessels; in C. austrosinensis Prismatic crystals present in all species, thickenings also distinct in wide vessels. integumented or not, abundant to infrequent, Thin-walled and sclerotic tyloses present (see mostly in non-chambered upright and square Table 2). Unbleachable dark materials present ray cells (sometimes chambered in some sam­ in C. vandervoetiana. ples) and as viewed in tangential sections Vascular tracheids absent. crystals often in sheath cells, less common Libriform fibres 1050-1270 (700-1950) in procumbent ray cells, infrequent in non­ J..lm long, thin- to thick-walled (but thick- to chambered axial parenchyma cells in C. kor­ very thick-walled in C. wightii var. consimi­ aiensis, occasionally in sclerotic tyloses (one lis), with simple to minutely bordered pits con­ sample of C. sinensis, CAFw 13083); crystal­ fined to the radial walls. Simple perforations containing cells enlarged or not, generally occasionally present in C. vandervoetiana. with one, sometimes with 2 or 3, crystal(s) Axial parenchyma abundant, mainly para­ per cell. Druses occur in some species or tracheal: vasicentric, confluent, aliform, and samples (see Table 2). Silica absent. slightly wavy tangential paratracheal bands of 4-8 (2-10, occasionally up to 22) cells; and in 1-2-seriate, marginal bands (see Table 2), Group II - Semi-ring-porous to diffuse-por­ in strands of 2-4(-10) cells. Simple perfora­ ous species of Celtis tions present in C. vandervoetiana. Material studied: C. austrosinensis Chun.: Rays 5-8(-12) or 12-17/mm, 1-5(-8) Guangdong: CAFw 16706. - C. vandervoet­ cells wide (see Table 2), usually of two sizes: iana Schneid.: Jiangxi: CAFw 17413; Gui­ 1-2-seriate and 3-5(-8)-seriate, 0.03-0.5 zhou: CAFw 17624. - C. wightii Planch var. (-1.7) mm high. Multiseriate rays heterocel­ consimilis (Blume) Gagnep: Hainan: CAFw lular, composed of procumbent body ray 6795. cells and 1-3(-7) rows of square and/or up­ Deciduous or evergreen trees from sub­ right marginal cells, in C. wightii var. cons i­ tropical to tropical regions. milis, uniseriate rays mostly with procumbent, Growth rings faint to distinct, marked by square and upright cells mixed throughout marginal or seemingly marginal parenchyma the ray. Weakly differentiated sheath cells bands and locally inflated multiseriate rays, rare to absent. Disjunctive ray parenchyma and moderate differences in vessel diameter. cell walls only present in C. austrosinensis. Wood semi-ring-porous to diffuse-porous, Perforated ray cells present (see Table 2). vessels 1O-23/sq.mm, 30-58% solitary, Prismatic crystals integumented, mostly in remainder mainly in radial or oblique pairs or upright and square ray cells, less common in multiples of 3(-9), and in clusters of 3-4 procumbent ray cells, sometimes in axial pa­ (-11), oval to round, tangential diameter 70- renchyma cells. Crystal containing cells un­ 105 (50-200) J..lm, radial diameter up to 270 enlarged and non-chambered, generally with J..lm, only up to 95 J..lm in C. wightii var. con­ 1(-2), sometimes with 3-5 or fragmented similis, walls 1-7 J..lm thick. Vessel element crystal(s) per cell. Druses occur in C. van­ length 290-360 (140-490) J..lm. Perforations dervoetiana. Silica absent. simple in horizontal to oblique end walls. In­ tervessel pits nonvestured, alternate, round in Notes: 1. Celtis philippensis was describ­ C. austrosinensis, mainly polygonal in the ed as diffuse-porous by Kanehira (1921), other two species, 3-10 J..lm in diameter, with Sudo (1963), Sweitzer (1971) and Ho (1985). slit-like, often coalescent apertures. Vessel­ The identity of our material has been confirm­ ray and vessel-axial parenchyma pits simple ed by three collectors, although Tang Yan­ or with reduced borders, rounded or horizon­ cheng, the taxonomist of Chinese Ulmaceae, tally elongate, and unilaterally compound in thought that our material might have been C. austrosinensis and C. vandervoetiana misidentified (personal correspondence). Ac­ (CAFw 17624); partly half-bordered in C. cording to Cheng et al. (1985), C. philippen-

Downloaded from Brill.com10/03/2021 10:07:11PM via free access w~ ~ 1 Table 2. Wood anatomical diversity in Celtis.

Material 2 3 4 5 6 7 8 9 10 11 12 13 Group I

C. biondii CAFw 16906 R 174 VC-BM (D) 4-6 1-9 1-5 + U (P) a CAFw 20264 R (S) 90 240 5-10 1100 VAC-BM 8-13 1-10 1-5 + U (P) a HEFw 003164 R 200 270 5-7 1160 VC-BM 4-7 1-10 1-3 + + UPa

C. bungeana CAFw556 R (S) 169 VMC 5-8 1-8 1-3 + UP (e) a CAFw 5511 R 115 230 5-9 950 VC-BM 6-12 1-6 1-2 + U (P) (e) + CAFw 17072 R 170 290 1300 VC-BM 6-9 1-9 1-4 + + UP

C. koraiensis R 175 220 890 VMC-B 5-7 1-10 1-2 + UP (A + 1- > C. macrocarpa R 190 240 7-12 1170 VMC-B 7-10 1-8 1-4 UP (ec) a >~ tc Downloaded fromBrill.com10/03/2021 10:07:11PM E.. C. philippensis R 190 380 5-9 1410 VMC-B 4-9 1-10 1-5 + + UP (ec) a + (i" g. C. sinensis ? CAFw13083 R 200 280 3-7 1340 VMC-B (A) 3-7 1-8 1-5 + UPS (e) a ~ <: CAFw6663 R 185 VMC-B 8-10 1-9 1-5 + UP (ec) a + ~ CAFw5709 R 150 240 990 VC-BM 6-11 UP (ec) ...... 5-9 1-9 1-6 + w C. yunnanensis R 180 310 5-12 1400 VMC-B 4-8 1-12 1-5 + U (P) (e) ~ + ...... via freeaccess \0 \0 N IN I ::r

Material 1 2 3 4 5 6 7 8 9 10 11 12 13 I~ to Group II I ~ '" C. austrosinensis (S) D 105 290 3-5 1270 AC-BM 5-9 1-5 1-5 (±) + UA(P)a IRo l::r!~ g C. vandervoetiana if CAFwI7413 S (D) 105 330 5-8 1050 VM(CA) 5-12 1-6 1-3 + UPA + I CAFw17624 S 180 5-10 VM(CA) 5-8 1-8 1-4 (±) UPAa + C. wightii [ var. consimilis D 70 360 4-6 1170 C-BAM (V) 12-17 1-5 1-7 UAa §

1. Porosity; S = semi-ring-porous; R = ring-porous; D = diffuse-porous. '<~ 2. Average tangential vessel diameter (Ilm); in ring-porous samples only calculated for early wood pores; in semi-ring-porous and diffuse-porous g, samples calculated for the whole growth ring. s 3 . Average vessel element length (Jlffi). 4. Intervessel pit size (Ilm). 5. Average fibre length (Jlffi). I ::;-> 6. Parenchyma distribution; V = vasicentric; C = confluent; C-B = confluent-banded; A = aliform; M = marginal parenchyma bands; D = diffuse. o 3 Downloaded fromBrill.com10/03/2021 10:07:11PM 7. Ray frequency Umm). Q 8. Ray width (number of cells). S· 9. Number of rows of square to upright marginal cells. I» 10. Sheath cells; ± = weakly differentiated sheath cells present in some rays; (±) = very weakly differentiated and rarely present 11. Perforated ray cells. 12. Prismatic crystals; U = present in upright/square ray cells; P = in procumbent cells; A = in axial parenchyma cells; S = in sclerotic tyloses; a = abundant; e = enlarged; c = chambered. 13. Druses. via freeaccess () = of sporadic occurrence. w VI""" 436 IAWA Bulletin n.s., Vol. 13 (4),1992

sis and Ulmus tonkinensis are ring-porous Growth rings faint, marked by differences when grow-ing in north temperate regions, in fibre wall thickness and diameter, and very but diffuse-porous in Hainan. It is possible limited marginal parenchyma bands. Wood that C. philippensis also becomes ring-porous diffuse-porous, vessels 8-9 (4-13)/sq.mm, in Guangxi (subtropical region). Sweitzer 38-51 % solitary, remainder mainly in radial (1971) earlier found that wood of Planera multiples of 2-4(-7), sometimes in clusters aquatica varied from diffuse-porous to ring­ of 3-4(-5) and oblique pairs, round to oval, porous. Wheeler et al. (1989) also noted a or weakly angular, tangential diameter 130- similar variation in C. laevigata. 140 (85-190) J.lm, radial diameter up to 205 2. According to Tang Yancheng, our C. J.lm, walls 2-5 J.lm thick. Vessel element austrosinensis Chun. may represent an un­ length 700-710 (300-1240) J.lm. Perfora­ published new species in China, and the tions simple in oblique to horizontal end walls. samples we have of C. bungeana (CAFw Intervessel pits nonvestured, alternate, main­ 17072, Guangdong) might be misidentified ly polygonal, 7-15(-20) J.lm in diameter, since this species is not very likely to occur with oval to slit-like, sometimes coalescent in Guangdong. He referred C. wightii Planch. apertures. Vessel-ray and vessel-axial pa­ var. consimilis (Blume) Gagnep to C. collin­ renchyma pits simple to half-bordered, of sae Craib (personal correspondence). varying size and shape: small to large, round 3. Cox (1941), Grumbles (1941), Sweitzer and angular to horizontally elongate, some (1971) and Wheeler et al. (1989) found that with pointed ends, sometimes unilaterally evergreen species of Celtis are diffuse-por­ compound. Spiral thickenings virtually ab­ ous and deciduous species are ring-porous. sent, but in G. yunnanensis very faint spiral However, the deciduous C. vandervoetiana thickenings present throughout some narrow is semi-ring-porous to diffuse-porous. vessel elements. Thin-walled tyloses com­ 4. The wood structure of C. biondii is vari­ mon. able. Most rings in sample CAFw 20264 look Vascular tracheids absent. semi-ring-porous, and vascular tracheids are Libriform fibres 1500-1580 (550-2080) rare. The earlywood vessels are widely spac­ J.lm long, thin- to thick-walled, with simple to ed in CAFw 16906, and crystals occur in minutely bordered pits confined to the radial both procumbent and upright/square ray pa­ walls. Simple perforations occasionally occur renchyma cells ofHEFw 003164. in G. cuspidata and G. yunnanensis. Gelati­ 5. Luo (1989) noted crystals in axial paren­ nous fibres common. chyma cells and tyloses in C. yunnanensis. Axial parenchyma predominantly paratra­ and reported spiral thickenings to be absent cheal (vasicentric, aliform, occasionally con­ in C. wightii. fluent and confluent-banded), in 1-4(-9)­ 6. None of the previous wood anatomical seriate bands or lines, and in very limited descriptions of Celtis in China mentioned I-seriate marginal lines, rarely diffuse (see marginal parenchyma bands. Table 4), in strands of 3-8 (2-13) cells. 7. We did not notice perforations with ves­ Rays 9-12(-13)/mm, 1-4(-5) cells wide, tigial bars in our material as Sweitzer (1971) 0.15-1.0(-1.8) mm high. Heterocellular rays did in the ring-porous species he studied. composed of procumbent body ray cells and 1-4(-7) rows of upright marginal cells, some­ times with procumbent and square cells mix­ Gironniera Gaudich. (Figs. 20, 36, 37, 38; ed throughout the ray; uniseriate rays as high Table 3) as multiseriate rays and composed of upright Evergreen trees or shrubs from south sub­ cells only. Weakly differentiated sheath cells tropical to tropical regions. common (see Table 3). Disjunctive ray pa­ renchyma cell walls present. Subgenus Galumpita Blume Crystals absent. Vitreous silica common in Material studied: G. cuspidata (Blume) vessels and fibres. Unbleachable, unidenti­ Planch. ex Kurz.: Hainan: CAFw 12148. - fied granular contents common in ray cells, G. yunnanensis Hu: Yunnan: CAFw 11934. occasionally in axial parenchyma cells.

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Table 3. Wood anatomical diversity in Gironniera.

Material 2 3 4 5 6 7 8 91011 Subg. Galumpita

G. cuspidata F 8 51 140 710 8~15 1580 BACVSM(D) 7~11 ± 1.8 G. yunnanensis F 9 38 130 700 7~14 1500 BACVSM 9-13 ± 1.8

Subg. Gironniera G. subaequa/is w8 A 10 43 120 720 8~13 1260 SV(D) 8~12 ~ 3.8 w9596 A 10 132 7~12 SV(D) 5~11 - 3.0 1. Growth rings boundaries; A = absent, F = faint. 2. Vessel frequency (per sq.mm). 3. Solitary vessels (%). 4. Average tangential vessel diameter (Jllll). 5. Average vessel member length (11m). 6. Intervessel pit size (11m). 7. Average fibre length (11m). 8. Parenchyma distribution; A = aliform; B = banded; C = confluent; D = diffuse; M = margi­ nal; S = scanty paratracheal; V = vasicentric. 9. Ray frequency (/mm). 10. Sheath cells; ± = weakly differentiated; - = absent. 11. Maximum ray height (mm).

Subgenus Gironniera Planch. (Subgenus Ne­ times unilaterally compound. Spiral thicken­ matostigma) ings absent. Thin-walled tyloses common. Material studied: G. subaequalis Planch.: Vascular tracheids absent. Guangxi: Guangxi Forestry College, w 8; Libriform fibres 1260 (870~1670) 11m Guangdong: CAFw 17005; Hainan: CAFw long, thin- to thick-walled, with simple to 9596; Yunnan: CAFw 11758. minutely bordered pits confined to the radial walls. Perforations absent. Growth rings absent. Wood diffuse-por­ Axial parenchyma less abundant than in ous, vessels 10 (5~20)/sq.mm, 43% soli­ subgenus Gaiumpita, scanty paratracheal and tary, remainder mainly in radial multiples of vasicentric, rarely diffuse, in strands of 4~8 2~4(~9), sometimes in clusters of 3~4(~5) (2~11) cells. and oblique pairs, round to oval, or weakly Rays 5~12(~13)/mm, 1~5 cells wide, angular, tangential diameter l20~132 (85~ 0.15~1.2(~3.8) mm, most frequently over 180) 11m, radial diameter up to 210 11m, walls 1 mm high. Heterocellular rays composed of 2~5 11m thick. Vessel element length 720 procumbent body ray cells and 1~4(~7) rows (300~ 1000) 11m. Perforations simple in ob­ of upright marginal cells, sometimes with pro­ lique to horizontal end walls. Intervessel pits cumbent and square cells mixed throughout nonvestured, alternate, mainly polygonal, 8~ the ray; uniseriate rays composed of upright 13(~20) 11m in diameter, with oval to slit-like, cells only. Weakly differentiated sheath cells sometimes coalescent apertures. Vessel-ray sometimes present (see Table 3). Disjunctive and vessel-axial parenchyma pits simple to ray parenchyma cell walls absent. half-bordered, of varying size and shape: Crystals absent. Vitreous silica common small to large, round and angular to horizon­ in vessels and fibres, occasionally in axial tally elongate, some with pointed ends, some- parenchyma cells. Unbleachable, unidentified

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granular contents common in ray cells, occa­ sometimes in clusters of 3-4(-5) and ob­ sionally in axial parenchyma cells. lique multiples of 2-3, oval to round tan­ diameter 85-110 (70-145) Ilm, ;adial Notes: l. Sweitzer (1971) observed pris­ g~ntial dIameter up to 185 Ilm, wallS,4-7 Ilm thick. matic crystals in ray cells and axial parenchy­ Vessel element length 230-295 (180-390) ma cells of G. cuspidata, and Cheng et al. Ilm. Perforations simple in oblique to hori­ (1979, 1985) noted prismatic crystals in axial zontal end walls. Intervessel pits nonvestured, parenchyma cells of G. cuspidata, G. subae­ al~emate, polygonal, 3-4 Ilm in diameter, qua/is and G. yunnanensis. Only unbleach­ WIth oval to slit-like, often coalescent, aper­ able granular contents were observed in the tures. Vessel-ray pits half-bordered or with species we studied. Absence of crystals in much reduced borders to apparently simple, G. subaequalis was also reported by Cheng small, rounded, sometimes unilaterally com­ et al. (1980). pound. Vessel-axial parenchyma pits half­ 2. Sweitzer (1971) did not observe silica bordered, similar to intervessel pits in shape in the 9 samples (belonging to 5 species, in­ and size. Fine spiral thickenings present cluding G. cuspidata and G. subaequalis) he throughout vessel elements visible with SEM. studied. Cheng et al. (1979, 1985) noted sili­ Both thin-walled and sclerotic (medium thick­ ca in axial parenchyma cells and ray cells of walled) tyloses common in P. tatarinowii G. cuspidata and G. yunnanensis, and in axial var. pubescens only. parenchyma cells, ray cells and fibres of G. Vascular tracheids absent. subaequalis; Cheng et al. (1980) noted silica Libriform fibres 1035-1280 (590-1400) common in ray cells and occasionally in axial Ilm long, thick- to very thick-walled, with parenchyma cells and fibres of G. subaequalis simple to minutely bordered pits confined to (CAFw 9596 etc.). We only observed vitre­ the radia~ walls. Simple perforations fairly ous silica in vessels, fibres and occasionally common m both tangential and radial walls. in axial parenchyma cells in this genus. Axial parenchyma abundant, mainly ali­ 3. Rays are very markedly heterocellular form to confluent and confluent-banded and in this genus with the multiseriate rays hav­ in 2-4(-8)-seriate marginal bands, also ~asi­ ing high upright marginal cells and the uni­ centric and less commonly scanty paratra­ seriate rays having all upright cells. cheal, in strands of 5-8 (2-9) cells. 4. Intercellular canals were reported by Rays 8-12(-13)/mm, of 2 more or less Den Berger and Bianchi in the rays of Giron­ dis~inct sizes: 1(-2)-seriate and 3-5(-6)­ niera (Metcalfe & Chalk 1950), but we did not senate, 0.15-0.75 mm high. Rays hetero­ find such canals in our material. cellular, composed of procumbent body ray cells and 1-3(-8) rows of upright margi­ nal cells, uniseriate rays with procumbent, Pteroceltis Maxim. (Figs. 14, 19,29,34,35, square and upright cells mixed throughout the 39, 40; Table 4) ray; and at times composed of square and/or Material studied: P. tatarinowii Maxim.: upright cells only. Disjunctive ray paren­ Anhui: CAFw13667; Guangdong: CAFw chyma cell walls present. Irregular tannini­ 16386; Henan: CAFw 5493; Shanxi: HEFw ferous cells with pointed outgrowths present. 779. -Po tatarinowii Maxim. var. pubescens Rays and axial elements irregularly storied Hand.-Mazz.: Guangxi: CAFw 1617l. (CAFw 16386) or non-storied (CAFw 16171). Deciduous trees from temperate to sub­ Prismatic crystals present (see Table 4), tropical regions, native to China, also culti­ mostly in upright cells, sometimes in 2-cham­ vated. bered ray cells, less commonly in procumbent Growth rings faint, marked by marginal cells, sporadic or absent in axial parenchyma parenchyma bands, locally inflated multiseri­ cells; cells mostly enlarged, chambered or ate rays and differences in fibre wall thickness. not, usually one integumented crystal per cell Wood diffuse-porous, vessels 19-24 (12- or chamber. Silica absent. Reddish-brown 39)/~q.mm, 73-75 (52-100)% solitary, tannins present in ray cells (and tanniniferous remaInder mainly in radial multiples of 2-4, tubes), occasionally in axial parenchyma cells.

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Table 4: Wood anatomical diversity in Pteroceltis.

Material 2 3 4 5 6 7 P. atarinf)wii CAFw 13667 24 85 260 1280 3-8 nm (s) CAFw 16386 20 110 295 1035 2-8 (c) a als P. tatarinowii var. pubescens 19 85 230 1065 2-4 (c) a (s)

1. Vessel frequency (per sq.mm). 2. Average tangential vessel diameter (J.Lm). 3. Average vessel member length (J.Lffi). 4. Average fibre length (J.Lm). 5. Width (number of cells) of marginal parenchyma. 6. Crystals; n = in non-chambered ray cells; (c) = chambered; a = abundant; m = moderately common. 7. Tanniniferous tubes; a = abundant; 1 = long; s = short.

Tanniniferous tubes common in P. tatari­ talis (L.) Blume: Taiwan: CAFw 7267; Hai­ nowii (CAFw 16386), extending radially nan: CAFW 14601; Yunnan: CAFw 12022; among rays, weakly differentiated to well de­ Guangdong: CAFw 16690; Guangxi: Guang­ veloped, up to 400 J.Lm long or longer (some­ xi Forestry College, w 11. - T. virgata Blume: times difficult to measure), 7-20 J.Lm high, Yunnan: CAFw 11805; Sichuan: CAFw 6215; rare and weakly differentiated in the other Hunan: CAFw 13472. two samples. Evergreen or deciduous small trees or shrubs from south subtropical to tropical re­ Notes: 1. Vessels are mainly in oblique or gions. radial multiples according to Cheng et al. (1985) and Ho (1985), but in our and Sweit­ Growth rings faint to intermediate, and dis­ zer's material (1971) 73-75% of the vessels tinct in T. cannabina and T. virgata (CAFw are solitary. Spiral thickenings in vessels were 13472), marked by differences in fibre wall not reported by the authors mentioned above, thickness and diameter, in T. cannabina and but such thickenings are visible with the SEM. T. virgata (CAFw 13472) also by differences 2. Sweitzer (1971) referred to uniseriate in vessel diameter. Wood mostly diffuse-por­ rays as homocellular with procumbent cells, ous, but semi-ring-porous in T. cannabina but we, as well as other authors, only ob­ and T. virgata (CAFw 13472), vessels 9-26 served heterocellular rays. (6-49)/sq mm, 23-66 % solitary, remain­ 3. Tanniniferous tubes have not been re­ der mainly in radial multiples of 2-3(-7), ported in the Ulmaceae before. To verify tube rarely in clusters of 3-4(-11) and oblique contents, we stained unbleached radial sec­ pairs, oval to round, or weakly angular, tan­ tions of two samples of CAFw 16386 with a gential diameter 90-140 (65-165) J.Lm, radial FeC13.5H20 solution and compared them diameter up to 230 J.Lm, walls 2-6 J.Lm thick. with unstained ones. The tube contents of the Vessel element length 400-510 (190-850) stained sections are obviously darker than J.Lm. Perforations simple in oblique to horizon­ those of the unstained ones. tal end walls. Intervessel pits nonvestured, alternate, polygonal or round, 4-8 (4-14) Trema Lour. (Figs. 1-3, 10, 17, 18; Table 5) J.Lm in diameter, with oval to round or slit­ Material studied: T. angustifolia (Planch.) like, sometimes coalescent apertures. Vessel­ Blume: Yunnan: CAFw 11828. - T. canna­ ray and vessel-axial parenchyma pits with bina Lour.: Fujian: CAFw 19820. - T. orien- much reduced borders to simple, or half-bor-

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Table 5. Wood anatomical diversity in Trema.

Material 2 3 4 5 6 7 8 9 T. angustifolia F D 15 66 130 440 4-11 890 9-13 T. cannabina D S 20 36 90 400 4-10 695 9-15 T. orientalis CAFw7267 F D 14 44 128 8-14 7-13 CAFw 14601 F D 9 65 140 460 7-11 915 7-13 T. virgata CAFw 11805 F D 10 55 90 510 5-11 1035 7-13 CAFw621 F D 26 23 115 410 5-10 1115 9-12 1. Growth rings boundaries; D = distinct; F = faint. 2. Porosity; D = diffuse; S = semi-ring-porous. 3. Vessel frequency (per sq. mm). 4. Solitary vessels (%). 5. Average tangential vessel diameter (!lm). 6. Average vessel element length (!lm). 7. Intervessel pit size (!lm). 8. Average fibre length (!lm). 9. Ray frequency (fmm).

dered (see Table 5), of varying size and shape: Notes: 1. Spiral thickenings have not been small to large, round to angular, or horizon­ reported before for T. angustifolia and T. vir­ tally to vertically elongate, sometimes unilate­ gata (CAFw 6215). In our material, very faint rally compound (T. angustifolia). Very faint spiral thickenings occur in narrow vessel ele­ spiral thickenings present in narrow vessel ments. The occurrence of helical thickenings elements of T. angustifolia and T. virgata in T. angustifolia, a wood without distinct (CAFw 6215). Thin-walled tyloses present, growth rings, is unusual. and sclerotic ones in T. cannabina and T. vir­ 2. Cheng et al. (1980) and Luo (1989) re­ gata (CAFw 6215) (see Table 5). Granular ferred to fibres in T. angustifolia and T. orien­ deposits present inside vessel walls. taUs as fibre-tracheids with distinctly border­ Vascular tracheids absent. ed pits (3.2-4.0!lm in diameter). Yang and Libriform fibres 695-1040 (510-1350) Huang-Yang (1987) also reported fibre-tra­ !lm long, very thin-walled, with simple to cheids in T. orientalis. But we only observed minutely bordered pits mainly confined to the libriforrn fibres with simple to minutely bor­ radial walls. Simple perforations observed in dered pits. T. angustifolia and T. virgata, relatively com­ 3. Yang and Huang-Yang (1987) described mon in tangential section. intervessel pitting as opposite in T. orientalis, Axial parenchyma rare, scanty paratracheal but we, as well as Cheng et al. (1985) and and rarely diffuse, in strands of 3-8 (2-12) Luo (1989) only observed alternate interves­ cells. sel pits. Rays 7-12(-15)/mm, 1-3(-4) cells wide, 4. Sweitzer (1971) referred to uniseriate 0.1-1.0 (0.05-2.0) rnrn high. Rays com­ rays as homocellular with procumbent cells in posed of comparatively few procumbent body T. orientalis. However, in our and others' ray cells and 1-2(-6) rows of upright margi­ studies, only heterocellular rays were ob­ nal cells, sometimes with procumbent and served. upright cells mixed throughout the ray; or 5. Cheng et al. (1980) and Luo (1989) re­ composed of square and/or upright cells only. ported that crystals are occasionally present in Disjunctive ray parenchyma cell walls present. axial parenchyma cells of T. angustifolia and Crystals and silica bodies absent. T. orientalis. Cheng et al. (1985) describ-

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ed that silica sometimes occurs in the ray and 4-9-seriate, 0.05-0.8(-1.5) mm high. cells of these two species. We, as well as Rays sometimes homocellular, but mainly Sweitzer (1971) and Yang and Huang-Yang heterocellular, composed of procumbent body (1987) did not find crystals or silica. ray cells and 1-4 rows of upright to square cells, sometimes with all ray cells square and/ or upright, and at times with procumbent, Subfamily: Ulmoideae square and upright cells mixed throughout the ray. Perforated ray cells infrequently present. H emiptelea Planch. Locally storied arrangement of vessels, Material studied: H. davidii (Hance) vascular tracheids and axial parenchyma Planch.: Liaoning, w 1. sometimes occurs. Deciduous small trees or shrubs from Crystals and silica absent. temperate to subtropical regions. Notes: 1. Sweitzer (1971) observed some Growth rings distinct, marked by differ­ septate fibres and prismatic crystals in ray ences in vessel diameter, marginal parenchy­ parenchyma, however, we did not find these ma bands and locally inflated multiseriate features in our material, nor did Minaki et al. rays. Wood ring-porous and som~times se~i­ (1988) who examined seven specimens of H. ring-porous, earlywood pores In 2-3 dIS­ davidii. continuous rows, round to oval, mean tan­ 2. The specimens of H. davidii examined gential diameters of the solitary earlywood by Minaki et al. (1988) were all ring-porous vessels 70 (30-135) 11m, radial diameter up with an abrupt transition from earlywood to to 150 11m, latewood vessels predominantly latewood and the mean tangential diameter in clusters, in wavy tangential to diagonal (142 11m) and mean radial diameter (174 11m) bands, walls 1.5-4 11m thick. Vessel element of the earlywood vessels in their material were length 210 (150-300) 11m. Perforation~ sim­ larger. ple, rarely scalariform with 1 bar, and SImple with vestigial bars, in oblique to horizontal end walls. Intervessel pits nonvestured, alter­ Ulmus Linn. (Figs. 5,23-27,31; Table 6) nate, polygonal and round, 5-7 11m in diam­ Group I - Ring-porous species of Ulmus. eter, with round to oval and slit-like, some­ Material studied: U. changii Cheng.: Jiang­ times coalescent apertures. Vessel-ray and xi: HEFw 002179. - U. davidiana Planch.: vessel-axial parenchyma pits with much re­ Shanxi: HEFw 82cw844; Shaanxi: CAFw duced borders to simple, rounded, and hori­ 7767; Jiangxi: CAFw 18041. - U. laciniata zontally elongate, smaller than intervessel (Trautv.) Mayr: Hebei: CAFw 7784; Jilin: pits. Spiral thickenings present, distinct in CAFw 18074. - U. lasiophyl/a (Schneid.) narrow vessels, faint to absent in wider ves­ sels. Cheng: Guizhou: CAFw 17623. - U. macro­ carpa Hance: Hebei: CAFw 5321; Shaanxi: Vascular tracheids, intergrading with nar­ CAFw 554. - U. parvifolia Jacq.: Shandong: row vessel elements, associated with llie ves­ CAFw 5708; Guangxi: Guangxi Forestry sel-axial parenchyma groups, with distinct College, w 14; Jiangxi:CAFw 8468; Jiangsu: spiral thickenings. CAFw 6685; Guangdong: CAFw 17066; An­ Libriform fibres 1120 (710-1400) 11m hui: HEFw 3362. - U. propinqua Koidz.: long, thick- to very lliick-walled, with simple Jiangxi: CAFw 17397. - U. pumila Linn.: to minutely bordered pits confined to the ra­ Anhui: HEFw2134; Xinjiang: CAFw 13822. dial walls. uyematsui Hayata: Taiwan: CAFw 5033. Axial parenchyma abundant, vasicentric, - U. - U. wilsoniana Schneid.: Guangxi: CAFw in latewood intermixed willi llie vascular tra­ 15898. cheids and surrounding the vessels, confluent Deciduous trees or rarely shrubs from tem­ to confluent-banded, and in marginal 1-3- perate to subtropical (rarely tropical) regions. seriate bands; in strands of 2-4 cells. Rays 6-8/mm, 1-9 cells wide, ten~ing Growth rings distinct, marked by differ­ to be of two distinct sizes: 1-2(-3)-senate ences in vessel diameter and marginal paren-

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Table 6. Wood anatomical diversity in Ulmus.

Material 2 3 4 5 6 7 8 9

Group I (Ring-porous) U. changii 1 (2) cd 200 245 5-12 1505 4-7 1-5 A (P) (e) U. davidiana HEFw 82cw844 mcd 155 190 5-14 965 6-9 1-6 + (Pe) CAFw7767 1-2 cd 261 270 975 3-7 1-6 U. laciniata CAFw7784 1 (2) c 167 1-7 CAFW 18074 1 (2) c 195 245 6-10 1385 4-7 1-7 (A) U. lasiophylla 1-2 c 240 8-13 5-7 4-5 A U. macrocarpa CAFw 5321 1 (2) c 210 7-12 4-6 1-13 - (A)(P) CAFW554 2-3 c 170 210 7-10 1000 5-9 1-7 (A) U. multinervis CAFw 15928 1-2 d 145 280 8-13 1200 3-6 1-4 A (P) e CAFw20280 1-2 d 150 195 7-13 1245 4-8 1-6 A (P) e U. parvifolia CAFw 570 2-3 c 156 6-10 7-10 1-5 A (P) e U. propinqua 1 dc 145 210 5-7 1115 5-9 1-3 + A (P) (e) U. pumila HEFw 002134 mc 140 210 5-9 1090 4-8 1-7 + CAFw 13822 mc 161 6-10 4-7 1-7 + U. uyematsui 1 d 190 290 8-10 1600 4-6 1-5 U. wilsoniana 1 (2) d 240 235 8-13 1545 4-6 1-5 A (P) ea

Group II (Diffuse-porous) U. lanceaefolia 135 310 7-9 1680 5-9 1-8 + Aa U. tonkinensis CAFw 146600 100 295 2-4 1500 9-12 1-4 A (P) a CAFw 6813 115 290 8-10 1420 8-10 1-4 A (P)

1. Number of rows of earlywood pores; m = many rows; c = continuous; d = discontinuous; cd = continuous to discontinuous. (for ring-porous woods, Group I). 2. Average tangential vessel diameter (11m); in ring-porous samples only calculated for early- wood pores; in diffuse-porous samples calculated for the whole growth ring. 3. Average vessel element length (J.Ul1). 4. Intervessel pit size (J.Lm). 5. Average fibre length (J.Lm). 6. Ray frequency (/mm). 7. Most common ray width (number of cells). 8. Perforated ray cells. 9. Crystals; A = in chambered axial parenchyma cells; P = in procumbent ray cells; e =enlarged; a = abundant.

Downloaded from Brill.com10/03/2021 10:07:11PM via free access Zhong, Baas & Wheeler - Wood anatomy of Ulmaceae from China 443 ------chyma bands and locally inflated multi seriate ied arrangement of narrow vessels, vascular rays. Wood mostly ring-porous, eariywood tracheids and parenchyma sometimes occurs pores mainly in 1-2 continuous or discontin­ locally. uous rows but in 3-10 continuous rows in Rays 4-10 (3-12)/mm, 1-3(-7) cells U. pumila and some samples of U. davidi­ wide and up to 13 cells wide in U. macrocarpa ana, round to oval, mean tangential diameters (CAFw 5321), in U. davidiana (HEFw 82cw of the solitary eariywood vessels 145-240 844) of two more or less distinct sizes: 1-2 J.Lm, radial diameter up to 350 ).tm, earlywood (-3)-seriate (rarely more than 10 cells high) to latewood transition generally abrupt but and 4-6(-13)-seriate, 0.04-0.6 (0.03-1.1) gradual in wide rings of U. davidiana (HEFw mm high. Rays essentially homocellular with 82cw844), U. lasiophylla, U. macrocarpa all cells procumbent, upright marginal cells (CAFw 554), U. multinervis, U. propinqua occur sporadically. Disjunctive ray parenchy­ and U. pumila (HEF w2134), latewood ves­ ma cell walls present in some species. Perfo­ sels predominantly in clusters, in wavy tan­ rated ray cells occasionally occur (see Table gential to diagonal bands, walls 2-4 (1-6) 6). Irregular cells with pointed outgrowths J.Lm thick. Vessel element lengths 190-290 present. (110-350) ).tm. Perforations simple in hori­ Prismatic crystals present or absent (see zontal to oblique end walls, occasionally in Table 6), integumented or not, abundant to side walls. Intervessel pits nonvestured, al­ infrequent, mostly in chambered, occasion­ ternate, round to oval and sometimes poly­ ally non-chambered, axial parenchyma cells, gonal, 5-10(-14) ).tm in diameter, with oval less common to absent in procumbent ray to round and/or slit-like, sometimes coales­ cells. Crystal-containing cells mostly enlarg­ cent apertures. Vessel-ray and vessel-axial ed, generally with one, sometimes with 2 parenchyma pits simple or with reduced bor­ crystal(s) per cell. Bleachable dark contents ders, small, rounded, and at times unilaterally abundant in parenchyma and ray cells, some­ compound. Spiral thickenings present, dis­ times in fibres. Silica bodies absent. tinct in narrow vessel elements, faint to ab­ sent in wide vessel elements, but in U. chan­ Group II - Diffuse-porous species of Ulmus. gii distinct spiral thickenings throughout the Material studied: U. lanceaefolia Roxb.: Si­ body of all vessel elements. Thin-walled ty­ chuan: CAFw 7960. - U. tonkinensis Jacq.: loses present in most samples (see Table 6). Guangxi: CAFw 16146; Hainan: HEFw Unbleachable dark contents present in some 146600, CAFw 6813. samples. Evergreen or deciduous trees from sub­ Vascular tracheids, intergrading with nar­ tropical to tropical regions. row vessel elements, associated with the ves­ sel-axial parenchyma groups, with distinct Growth rings distinct in U. lanceaefolia spiral thickenings. and U. tonkinensis (CAFw 6813), marked Libriform fibres 965-1600 (400-1990) by marginal parenchyma bands and locally J.Lm long; thin- to thick-walled, or thick- to inflated multi seriate rays. Vessels diffuse, very thick-walled, with simple to minutely 36-50 (12-71)/sq.mm, 20 (3-39)% bordered pits confined to the radial walls. solitary, remainder mainly in clusters of 3-4 Simple fibre perforations observed in U. pu­ (-11), and in radial pairs or multiples of mila. Gelatinous fibres common. 3(-9) and oblique pairs, round to oval, tan­ Axial parenchyma mainly paratracheal: gential diameter 100-135 (75-200) ).tm, ra­ vasicentric, scanty paratracheal, and inter­ dial diameter up to 180 ).tm, walls 3-7 ).tm mixed with the vascular tracheids and sur­ thick. Vessel element length 290-310 (170- rounding the vessels; and in marginal 1-2 440) ).tm. Perforations simple in horizontal to (-3)-seriate bands; rarely diffuse (see Table slightly oblique end walls. Intervessel pits 6), in strands of 2-4 cells if not containing nonvestured, alternate, round to oval, and crystals, crystal-containing strands generally polygonal in U. tonkinensis (CAFw 6813), of 2-8(-15) cells. Simple perforations ob­ 7-9(-10) ).tm in diameter (see Table 6), with served in U. lasiophylla and U. pumila. Stor- round to oval and slit-like, often coalescent

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apertures. Vessel-ray and vessel-axial pa­ U. wilsoniana (Guangxi) are questionable renchyma pits simple or with reduced bor­ because there have been no reports of these ders, small, rounded, sometimes horizontally species in Jiangxi / Guangxi. However, Xie elongate and unilaterally compound in U. ton­ Fuhui (personal communication) suggested kinensis (CAFw 6813). Spiral thickenings that the flora of Guangxi is not completely present, distinct in narrow vessels, faint to known and these samples may represent new absent in wide vessel elements, but in U. lan­ and legitimate records. ceaefolia such thickenings present throughout 2. Marginal parenchyma was not reported body of all vessel elements. Thin-walled and in Chinese publications about Ulmus. In sclerotic tyloses present (see Table 6). Un­ agreement with our observations, Sweitzer bleachable dark materials present in U. ton­ (1971) and Wheeler et al. (1989) mentioned kinensis (CAFw 6813). that most species of Ulmus have marginal Vascular tracheids very rare to absent. parenchyma bands. Libriform fibres 1420-1680 (850-2010) 3. Cheng et al. (1980) described druses in ~m long, thick- to very thick-walled, with U. pumila, and noted that druses and pris­ simple to minutely bordered pits confined to matic crystals are only present in axial the radial walls. Simple perforations present parenchyma cells. However, in our two sam­ in all species, especially common in U. ton­ ples neither druses nor prismatic crystals kinensis (CAFw 6813). occur. Axial parenchyma vasicentric, scanty para­ 4. In contrast with our observations, tracheal, and in 1-2-seriate (in U. lanceae­ Sweitzer (1971) described that in U. macro­ folia up to lO-seriate) marginal bands, rarely carpa and U. pumila there are vestigial bars diffuse (see Table 6), in strands of 2-4 cells on the perforation plates of narrow vessels if not containing crystals, crystalliferous and that there are no radial multiples, that in chains of 2-8(-24) chambers. Simple per­ U. macrocarpa tyloses are absent and inter­ forations observed in U. tonkinensis (CAFw vessel pitting is opposite, and in U. uyemat­ 6813). Sometimes local storying of narrow sui crystals are present. vessels, vascular tracheids and parenchyma. Rays 8-10 (5-12)/mm, 1-4(-8) cells Zelkova Spach (Figs. 8, 9, 12, 16, 28, 33; wide, 0.03-0.8 mm high. Rays essentially Table 7) homocellular with all cells procumbent, mar­ Material studied: Z. formosana Hayata: ginal upright cells occur sporadically. Disjunc­ Taiwan: CAFw 7340. - Z. schneideriana tive ray parenchyma cell walls present in U. Hand.-Mazz.: Guangxi: CAFw 16151; Gu­ tonkinensis. Non-enlarged perforated ray cells angdong: CAFw 17157; Jiangshu: CAFw occasionally occur in U. lanceaefolia. Irreg­ 13685; Sichuan: CAFw 412. - Z. serrata ular cells with pointed outgrowths present. Makino: Anhui: HEFw 001400; Guangxi: Prismatic crystals present (see Table 6), CAFw16112; Jiangsu: CAFw 4938; Shan­ integumented or not, abundant, mostly in dong: CAFw 5710. - Z. sinica Schneid.: Ji­ chambered axial parenchyma cells, crystallif­ angsu: CAFw 6686; Henan: CAFw 5503; erous chains of up to 24 chambers, less com­ Shaanxi: CAFw 581; Shanxi: CAFw 684; Si­ mon to absent in procumbent ray cells. Crys­ chuan: CAFw 23. tal containing cells often enlarged, generally Deciduous trees from temperate to tropical with one, sometimes with 2 crystal(s) per regions. cell. Bleachable dark contents abundant in parenchyma and ray cells, sometimes in Growth rings distinct, marked by differ­ fibres. Vitreous silica present in some small ences in vessel diameter, and mostly by vessel elements, rarely in fibres of U. lan­ marginal or seemingly marginal parenchyma ceaefolia. bands and locally inflated multiseriate rays as well. Wood ring-porous, earlywood pores in Notes: 1. According to Fu Liguo (person­ 1-2 discontinuous rows, round to oval, mean al communication), the identifications of our tangential diameter of the solitary earlywood two samples of U. propinqua (Jiangxi) and vessels 170-245 (60-320) ~m, radial diam-

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Table 7. Wood anatomical diversity in Zelkova.

Material 2 3 4 5 6 7 8 9 10 Z. formosana 190 195 5-7 T 1425 5-9 1-6 0.7 - SPA a Z. schneideriana CAFw 13685 195 220 8-13 TS 1715 4-8 1-10 0.8 + SPA a CAFw 16151 245 245 TS 1565 3-6 1-9 0.8 - SAa Z. serrata CAFw 5710 220 200 6-10 1600 3-6 1-8 0.9 - P (SA) CAFwOO1400 210 170 6-8 TS 1620 3-7 1-9 0.5 + P (SA) Z. sinica CAFw6686 180 235 6-9 1300 4-7 1-8 1.1 + SPA a CAFw 581 120 195 6-9 1150 4-7 1-7 0.9 + S (P) CAFw 5503 130 200 5-10 880 5-9 1-6 0.6 - S (P) 1. Average tangential vessel diameter (J.UTI) of earlywood vessels. 2. Average vessel element length (Jlm). 3. Intervessel pit size (Jlm). 4. Tyloses; T = thin-walled tyloses; S = sclerotic tyloses. 5. Average fibre length (Jlm). 6. Ray frequency (lmm). 7. Ray width (number of cells). 8. Maximum ray height (mm). 9. Perforated ray cells. 10. Crystals; S = in square and/or upright marginal parenchyma cells; P = in procumbent body ray cells; A = in axial parenchyma cells; a = abundant; ( ) = rarely present or variable.

eter up to 380 Jlm, earlywood to latewood Vascular tracheids, intergrading with nar­ transition generally abrupt, but more gradual row vessel elements, associated with the ves­ in one sample of Z. sinica (CAFw6686) which sel-axial parenchyma groups, with distinct has very wide growth rings, latewood vessels spiral thickenings. predominantly in clusters, in wavy tangential Libriform fibres 880-1715 (600-2070) to diagonal bands, walls 1-6(-10) Jlm thick. J.l.m long, thin- to thick-walled or thick- to Vessel element length 170-245 (90-300) very thick-walled, with simple to minutely Jlm. Perforations mostly simple, very rarely bordered pits confined to the radial walls. scalariform with 1-2 bars, in oblique to hori­ Simple perforations occasionally occur (see zontal end walls. Intervessel pits nonvestured, Table 7). Gelatinous fibres present in Z.for­ alternate, polygonal or weakly polygonal, in mosana. some species also round, 5-10(-13) Jlm in Axial parenchyma abundant, mainly para­ diameter, with round to oval and/or slit-like, tracheal, vasicentric, confluent, scanty para­ often coalescent apertures. Vessel-ray and tracheal in earlywood, in latewood intermixed vessel-axial parenchyma pits smaller than with the vascular tracheids and surrounding illtervessel pits, but mostly with reduced bor­ the vessels, and in 1-3-seriate marginal bands d~rs, rounded, and horizontally elongate in (see Table 7); in strands of 2-4 cells, simple some species, sometimes unilaterally com­ perforations occasionally occur (see Table 7). pound. Very distinct spiral thickenings pres­ Local storied arrangement of narrow ves­ ent throughout narrow vessel elements. Thin­ sels, vascular tracheids and axial parenchyma walled and sclerotic tyloses present (see Table sometimes present. 7). Unbleachable dark materials present in Z. Rays 3-9/mm, 1-6(-10) cells wide, formosana, Z. schneideriana (CAFw 13685) tending to be of two sizes in some samples: andZ. serrata (HEFw 001400). 1-2(-3)-seriate and 4-6(-1O)-seriate, 0.07-

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Table 8. Selected wood anatomical features of the Ulmaceae from China.

Celtidoideae 1 2 3 4 5 6 7 8 9 10 11 12 13

Trema DS He 400-510 4-14 Vrh (an) - Gironniera D He 700-720 7-15 Vrh (an) +/- Aphananthe D He 240-315 4-6 Shr iw + UP Plerocellis D He 230-295 3-4 Shr iw + RE UP(A) Cellis Group II DS He 290-360 3-10 Sr(h) dnw + USP(A) ± +/- Group I R(S) + + He* 220-380 3-12 Sr dn + VTA USP(A) + +/-

Ulmoideae Hemiplelea RS + + HeHo 210 5-7 Sr dn + VTA + Zelkova R + + HeHo 170-245 5-10 Sr dn + VTA SUPA -/± Ulmus Group I R + + Ho 190-290 5-14 Sr dnw + VTA A(P) -/± Group II D Ho 290-310 7-10 Sr dnw + VTA A(P)

Legends: + = character present, - = character absent, ± or () = character variable and/or intermediate between two contrasting character states. The order indicates relative abundance of a character.

1. Porosity; R = ring-porous, D = diffuse-porous, S = semi-ring-porous. 2. Latewood vessels in a wavy tangential to diagonal pattern. 3. Vascular tracheids. 4. Ray type; He = heterocellular, Ho = homocellular. * Homocellular rays occur in C. korai- ensis. 5. Average vessel element lengths (11m). 6. Intervessel pit size (11m). 7. Vessel-ray and vessel-parenchyma pits; S = similar to the intervessel pits in size and/or smaller than the intervessel pits, V = of varying shape and size, r = with much reduced bor­ ders to simple, h = half-bordered. 8. Spiral thickenings in vessels; a = very faint, i = fine, d = distinct, w = present in all vessels, n = confined to narrow vessels. 9. Marginal parenchyma bands. 10. Storied structure; RE = rays and axial elements irregularly storied, VTA == narrow vessel elements, vascular tracheids and axial parenchyma sometimes locally storied. 11. Crystal location; U == in upright ray cells, S == in square ray cells, P == in procumbent ray cells, A == in axial parenchyma cells. 12. Sheath cells. 13. Druses.

0.8 (0.04-1.1) mm high. Rays both hetero­ sana and Z. sinica. Very small perforated ray cellular and homocellular, heterocellular rays cells occasionally occur (see Table 7). composed of procumbent body ray cells and Crystals present in all species, integument­ 1-2(-3) row(s) of square to upright margi­ ed and prismatic, infrequent to abundant, nal cells; in Z. serrara mostly homocellular usually in non-chambered upright and square with all ray cells procumbent. Disjunctive ray cells, and/or procumbent ray cells, and/ parenchyma cell walls present in Z. formo- or chambered axial parenchyma cells (in

Downloaded from Brill.com10/03/2021 10:07:11PM via free access Zhong, Baas & Wheeler - Wood anatomy of Ulmaceae from China 447 ~----'=--~-~.-~-~~- chains of up to 8 chambers) (see Table 7); and shape. However, the promotion of the crystal-containing cells mostly enlarged, with two subfamilies to family level as suggested one crystal per cell. Silica bodies absent. by Grudzinskaya (1967) and Chernik (1975, 1980, 1981, 1982) is not supported by wood Notes: 1. Sweitzer (1971) noted only fibre­ anatomy, because of the many anatomical tracheids (fibres with bordered pits) in his characters common to both subfamilies. More­ material. We observed only libriform fibres over, there are transitional forms between Cel­ as did Cheng et al. (1985). tidoideae and Ulmoideae: Zelkova and Hemi­ 2. The above description of ray type agrees ptelea have both homocellular and heterocel­ closely with that of Japanese Zelkova by lular rays and resemble ring-porous species Sweitzer (1971), however, not with that of of Celtis; C. koraiensis has some homocellu­ Chinese Zelkova by Cheng et al. (1985). The lar rays. In some non-wood anatomical char­ latter authors refer to ray type as heterocellu­ acters Ampelocera and Aphananthe are inter­ lar ill (with one row of square to upright mar­ mediate between the Celtidoideae and Ulmoi­ ginal cells). Kanehira (1921a, 1921b) referred deae to the ray type as predominantly homocellu­ Our study provides some support for the lar. placement of genera by Grudzinskaya (1967). 3. Yang and Huang-Yang (1987) noted She placed Hemiptelea and Zelkova in the sheath cells and half-bordered vessel-ray pits ulmoid group, whilst Hutchinson (1967) in their material. Metcalfe and Chalk (1950) placed them in the Celtidoideae. There is sim­ also described rays with sheath cells in some ilarity between ring-porous Celtis species and species of Zelkova. Hemiptelea and Zelkova, but homocellular 4. Marginal parenchyma bands have never rays regularly occur in the former two genera been reported in Zelkova before. and their wide rays lack sheath cells, charac­ ters which we suggest ally them with the Ul­ Discussion moideae, rather than with Celtidoideae. The Ulmaceae, and in particular the Ulmoi­ Taxonomic and phylogenetic implications deae, should be considered specialised in Although the Ulmaceae as a family are di­ their wood anatomy. Derived characters, in verse in wood anatomy, there are many char­ the Baileyan sense, include the mostly sim­ acters pervading the entire group. These in­ ple perforation plates, alternate intervessel clude mostly exclusively simple perforations; pits, relatively short vessel elements and alternate, nonvestured intervessel pits; rela­ fibres, rays with a tendency to homogeneity, tively short vessel elements and fibres; non­ axial parenchyma occurring in groups (ali­ septate fibres with simple to minutely border­ form, confluent, confluent-banded, marginal ed pits confined to the radial walls; mainly bands); vessels in a wavy tangential to diago­ paratracheal parenchyma; rays rarely higher nal pattern and associated with vascular tra­ than 1 mm. However, two subfamilies, the cheids in many species; and storied structure Ulmoideae and the Celtidoideae, can be dis­ in some species. Wood anatomy does not tinguished on the basis of differences in wood support Giannassi's (1986) suggestion, based anatomy, mainly in ray structure (Table 8). on flavenoid chemistry, that Ulmoideae are Genera in the Ulmoideae have exclusively primitive relative to Celtidoideae. Gironniera homocellular rays or a mixture of homocellu­ and Trema (Celtidoideae) have the most prim­ lar rays and heterocellular rays, while genera itive wood anatomy of any of the Ulmaceae; in the Celtidoideae have heterocellular rays. their rays are markedly heterocellular, con­ Sheath cells are absent from the Ulmoideae, forming to Kribs heterogeneous type I in but are present in some Celtidoideae, particu­ Gironniera, and they have the longest vessel larly in the wider rays. All Ulmoideae have elements. vessel-ray parenchyma pits similar in size This study supports the generally accepted and shape to or slightly smaller than the in­ placement of the Ulmaceae in the order Urti­ tervessel pits, while some Celtidoideae have cales. The Ulmaceae, especially the Celtidoi­ vessel-ray parenchyma pits of varying size deae, resemble Moraceae in their wood anat-

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omy, the features listed above as characteristic because of the well-known paucity of field of the Ulmaceae also occur in the Moraceae notes on labels accompanying wood samples (Tippo 1938; Metcalfe & Chalk 1950). in institutional wood collections, the scanty The genus Gironniera - Traditionally, the ecological infonnation in most floristic litera­ genus Gironniera contains two subgenera: ture on the Chinese Ulmaceae, and the rela­ 1) Gironniera (Nematostigma Planch.) with tively small number of specimens studied G. subaequnlis and 2) Galumpita Blume with which may be insufficient to eliminate 'noise' G. cuspidata and G. yunnanensis (Bentham variation due to sampling diversity, only two & Hooker 1862; Engler & PrantI1893). How­ different ecological categories were recognis­ ever, G. cuspidata (Blume) Planch. ex Kurz ed in this study for the Ulmaceae from China: and G. yunnanensis H.H. Hu have been 1) Subtropical to tropical (19°-32° N latitude); transferred to the genus Aphananthe as A. this group includes all diffuse-porous (and a cuspidata (Blume) Planch. and A. yunnanen­ few semi-ring-porous) samples and part of sis (H.H. Hu) Phuphat. (Phuphathanaphong the ring-porous samples studied. 2) Temper­ 1972; Soepadmo 1977). Our study supports ate (above 32° N latitude); this group has only the traditional classification because wood ring-porous samples. anatomically subg. Galumpita is more similar In Figures 41-43, frequency diagrams are to subg. Gironniera than to Aphananthe (see given for vessel element length and fibre Table 8). Our material of subg. Galumpita length classes. It appears that in the temperate differs from Aphananthe in vessel element material vessel elements and fibres tend to be length, intervessel pit size, vessel-ray and shorter than in the subtropical to tropical ma­ vessel-parenchyma pitting, spiral thicken­ terial. Such trends can also be retraced within ings, tyloses, fibre length, ray height and individual genera in China such as Celtis, Ul­ composition, and mineral inclusions (crystals mus and Zelkova, or even within individual present in Aphananthe, silica present in subg. species like Z. sinica, and are in full agree­ Galumpita.) Wood anatomically Gironniera ment with the well-established general trends is more similar to Trema than to Aphananthe. (Baas 1986; Carlquist 1975, 1980). The relationships of Gironniera can be re­ Although vessel diameter and vessel fre­ solved only after the study of more material quency are crucial parameters for the hydraulic of Aphananthe, Gironniera, and Trema. Ka­ conductivity of wood, we can hardly analyse ryomorphological and palynological features, their ecological trends here. All diffuse-porous vernation patterns and seed coat morphology (including semi-ring-porous) samples studied all suggest that Gironniera may have a key belong to the subtropical to tropical group. In role in the understanding of the evolutionary ring-porous samples with a wavy tangential relationships between the celtoids and ulmoids to diagonal pattern and vascular tracheids in (Oginuma et al. 1990; Takahashi 1989; Tera­ latewood, it is too difficult, as well as mean­ bayashi 1991; Takaso & Tobe 1990; Zavada ingless, to calculate the average vessel diam­ 1983). Its wood anatomy clearly allies it with eter and vessel frequency of whole growth the Celtidoideae. rings. Figure 43 shows that in both groups the average tangential vessel diameter of Ecological trends in the wood anatomy of the earlywood pores tends to have a similar fre­ Ulmaceae from China quency distribution. The Ulmaceae are widely distributed in the Some qualitative characters tend to differ tropics, subtropics, and temperate regions in between tropical and subtropical material. As China, as far south as 19° and as far north as mentioned above, in the temperate samples, 50°, from sea-level up to 2500 m, varying wood is almost always ring-porous; in the from evergreen to deciduous and from tall tropical samples, ring-porosity is rare. Dif­ trees to small shrubs. The Chinese Ulmaceae fuse-porosity is nearly confined to the tropi­ are therefore an attractive group to test gener­ cal to SUbtropical samples. Some species, al hypotheses on ecological trends in wood such as C. philippensis, can be either diffuse­ structure as proposed by Baas (1986), Carl­ porous (in tropical regions) or ring-porous quist (1975, 1980), and others. However, (in a more seasonal environment). This in-

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Percentage of the samples (%) Percentage of the samples (%) 100 90 45

80 40

70 35

60 30

50 25

40 20

30 15 20 10 10 5

100 200 300 400 500 600 700 SOO 600 800 1000 1200 1400 1600 1800 2000 Vessel element length (~m) Fibre length (~m) Fig. 41. Vessel element length in different Fig. 42. Fibre length in different latitudinal latitudinal zones. Hatched = subtropical to zones. Hatched = subtropical to tropical; black tropical; black = temperate. = temperate.

Percentage of the samples (%) 70 terspecific vanatlon in porosity has been observed in other species, such as Planera 60 aquatica (Ulmaceae, Sweitzer 1971). How­ 50 ever, Aphananthe aspera is diffuse-porous in both temperate and subtropical regions, and 40 Celtis yunnanensis and Zelko va Jormosana are ring-porous even though they grow in 30 southern subtropical regions. Wavy tangen­ 20 tial to diagonal vessel patterns in combination with vascular tracheids are also confined to 10 extratropical latitudes. Spiral vessel wall o I.----r-"''''' thickenings are virtually absent in the tropical 50 100 150 200 250 to south subtropical genera Gironniera and Fig. 43. Earlywood vessel diameter in differ­ Trema, and well-developed in extratropical ent latitudinal zones. Hatched = subtropical to genera, being especially distinct in ring-por­ tropical; black = temperate. ous Ulmaceae. Parenchyma distribution and abundance show a similar trend: marginal parenchyma is rare in Gironniera, absent in Trema, but is always abundant in ring-porous porosity; tangential to diagonal vessel pattern, Ulmaceae. Storied structure is only absent in vascular tracheids, and vasicentric and Gironniera and Trema and diffuse-porous marginal parenchyma; storied structure; and Celtis (also tropical to south subtropical), growth rings) are also traceable within some while it is always present in ring-porous sam­ genera in China such as Celtis, and are in ples. This runs counter to the general trend basic agreement with general trends (e.g., for storied structure to be more common in Baas et al. 1984). tropical dicotyledons than in temperate ones. Gilbert (1940), Tippo (1938), Metcalfe and Related to ring-porosity to some extent, Chalk (1950) and Sweitzer (1971) have sug­ growth rings are less distinct in tropical to gested that the ring-porous condition is a re­ subtropical samples than in the temperate sponse to a temperate (seasonal) environment samples. Some of these ecological trends in and that resulting evolution has given impetus qualitative wood anatomical characters (ring- to development of other so-called 'advanced'

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characters, such as spiral vessel wall thicken­ 6. Spiral thickenings in vessels very fine to ings, relatively short vessel elements and absent, rays markedly heterocellular with fibres. The fossil record supports these sug­ distinctly upright marginal cells, crystals gestions as the incidence of these characters mostly in upright ray cells or absent. 7 is higher in the Late Tertiary than in the Early 6. Spiral thickenings in vessels distinct, rays Tertiary (Wheeler & Baas 1991). Our study heterocellular with square and/or upright of Ulmaceae wood also tends to support this marginal cells or homocellular, crystals contention. In the Chinese Ulmaceae it is es­ mostly in upright and square ray cells or pecially striking that all ring-porous woods axial parenchyma cells...... 10 possess many specialised characters such as 7. Vessel-ray pits similar to intervessel pits a wavy tangential to diagonal vessel pattern, in shape and size, crystals present, mar­ vascular tracheids, spiral vessel wall thicken­ ginal parenchyma bands obvious. .. 8 ings, abundant marginal parenchyma, rela­ 7. Vessel-ray pits of varying shape and tively shorter vessel elements and fibres and size, crystals absent, marginal parenchy­ storied structure, and are always restricted to ma absent or in very limited lines. .. 9 extratropicallatitudes, or seasonal environ­ 8. Growth rings faint, vessels 12-39/mm2, ments. However, ring-porous Ulmaceae have tangential vessel diameter smaller than very infrequent scalariform perforation plates 110 Ilm, rays and axial elements irreg­ with 1-2 bars and simple perforation plates ularly storied, crystal containing cells with vestigial bars in some narrow vessels, mostly enlarged, tanniniferous tubes while diffuse-porous woods invariably have sometimes present ...... Pteraceltis exclusively simple perforations. 8. Growth rings distinct, vessels 2-13/ mm2, tangential vessel diameter larger Generic wood anatomical key to Chinese than 120 Ilm, storied arrangement absent, Ulmaceae crystal containing cells non-enlarged, tan­ niniferous tubes absent . . . Aphananthe 1. Wood ring-porous or semi-ring-porous, 9. Axial parenchyma vasicentric, aliform, latewood vessels in wavy tangential to confluent-banded and in apotracheal diagonal bands...... 2 bands and limited marginal lines, rays 1. Wood diffuse-porous to semi-ring-por­ 1-4(-5)-seriate ...... Gironniera ous, latewood vessels not in wavy tan­ 9. Axial parenchyma scanty paratracheal and gential to diagonal bands ...... 6 diffuse, rays 1-2(-4)-seriate ... Trema 2. Rays both homocellular and heterocellu- 10. Rays heterocellular, sheath cells usually lar in one sample ...... 3 present ...... Celtis (Group II) 2. Rays either heterocellular or homocellu- 10. Rays essentially homocellular, sheath lar ...... 5 cells absent ...... Ulmus (Group II) 3. Sheath cells and druses present, crystals mostly in chambered axial parenchyma cells ...... Celtis koraiensis Acknowledgements 3. Sheath cells and druses absent .... 4 We are very grateful to the following per­ 4. Crystals present (mostly in enlarged non­ sons and institutions for providing wood sam­ chambered upright and square ray cells ples and information: Mr. Liu Peng, Wood and/or procumbent ray cells) . Zelkova Anatomy Division, Institute of Wood Indus­ 4. Crystals absent ...... Hemiptelea try, Chinese Academy of Forestry; Prof. Ke 5. Rays essentially homocellular, sheath Bingfan, Prof. Wei Guangyang, Mrs. Jiang cells absent, crystals mostly in chambered Zehui, Mrs. Wang Yanyan, Institute of For­ axial parenchyma cells or absent, druses est Products, Anhui Agricultural College: absent ...... Ulmus (Group I) Prof. Xie Fuhui, Mr. Xu Feng, Mr. Liang 5. Rays heterocellular, sheath cells usually Wen, Department of Forestry, Guangxi For­ present, crystals mostly in non-chamber­ estry College; Dr. Li Jian, Department of ed upright and square ray cells, druses Forest Industry, Northeast China Forestry sometimes present ... Celtis (Group I) University. We are indebted to Prof. Tang

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Yancheng and Prof. Fu Liguo (Chinese Chernik, V. V. 1975. Arrangement and re­ Academy of Sciences) and Dr. Kuoh Chang duction of perianth and androecium parts Sheng (Chengkung University of Taiwan) in representatives of the Ulmaceae Mirbel for kindly giving taxonomic advice, and to and Celtidaceae Link. Bot. Zhurnal 60: Bertie Joan van Heuven (Rijksherbarium) for 958-962. (In Russian.) her skillful technical assistance. Our thanks Chernik, V. V. 1980. Peculiarities of struc­ are also due to Dr. Shu-Yin Zhang for wide­ ture and development of the pericarp of ranging help. the representatives of the family Ulmaceae and Celtidaceae. Bot. Zhurnal 65: 521- Referenres 531. (In Russian.) Baas, P. 1986. Ecological patterns in xylem Chernik, V.V. 1981. Pseudomonomeric gy­ anatomy. In: T.J. Givnish (ed.), On the noecium of the Ulmaceae and Celtidaceae economy of plant form and function: 327- representatives. Bot. Zhurnal 66: 958- 352. Cambridge Univ. Press, Cambridge. 962. (In Russian.) Baas, P., P.M. Esser & M.E. T. van der Chernik, V. V. 1982. Characteristics of the Westen. 1984. Systematic and ecological structural development of spermoderm in wood anatomy of the Oleaceae, with spe­ some representatives of Ulmaceae and cial reference to dendritic vessel patterns. Celtidaceae. Bot. Zhurnal67: 1216-1220. In: S. Sudo (ed.), Proc. Regional Wood (In Russian.) Anatomy Conference Tsukuba: 153-155. Chiang, F.-C. 1962, 1964. Studies on the FFPRI, Tsukuba. anatomical structure and identification of Baas, P., P.M. Esser, M.E. T. van der the commercial timbers in Taiwan (1), (2). Westen & M. Zandee. 1988. Wood anat­ Bull. Taiwan For. Res. Inst., No. 81: 24 omyoftheOleaceae.IAWA Bull. n.s. 9: pp., 1962; No. 95: 25 pp., 1964. (In 103-182. Chinese & English.) Baas, P. & Xinying Zhang. 1986. Wood anat­ Cox, M.J. 1941. The comparative anatomy omy of trees and shrubs from China. I. of the secondary xylem of five American Oleaceae. IAWA Bull. n.s. 7: 195-220. species of Celtis. Amer. Midland Nat. 25: Bentham, G. & J.D. Hooker. 1862. Genera 348-357. Plantarum. Vol. I. L. Reeve. London. De Bary, A. 1884. Comparative anatomy of Carlquist, S. 1975. Ecological strategies of the vegetative organs of the phanerogams xylem evolution. Univ. Calif. Press, Ber­ and ferns (Transl. by F. O. Bower & D.H. keley, Los Angeles, London. Scott). Clarendon Press, Oxford. Carlquist, S. 1980. Further concepts in eco­ Deng, L. & P. Baas. 1990. Wood anatomy logical wood anatomy, with comments on of trees and shrubs from China II. Thea­ recent work in wood anatomy and evolu­ ceae. IAWA Bull. n.s. 11: 337-378. tion. Aliso 9: 499-553. Desch, H.E. 1954. Manual of Malayan tim­ Cheng, J.Q., N. Li, lJ. Yang & C.Z. Sun. bers II. Malay. For. Rec., No. 15: 329- 1980. Chinese tropical and subtropical 762. Revision of Foxworthy 1921. timbers, their distinction, properties and Engler, A. & K. Prand. 1893. Die Natiirlichen uses. Chinese Science Press, Beijing. (In Pflanzenfamilien. Volume 3. Borntraeger, Chinese.) Berlin. Cheng, J.Q., J.J.Yang & P. Liu. 1979. Mul­ Giannasi, D.E. 1978. Generic relationsips in tiple-entry perforated card key for identi­ the Ulmaceae based on flavenoid chemis­ fication of Chinese hardwoods by micro­ try. Taxon 27: 331-344. scopic features. Chinese Agricult. Press, Giannasi, D.E. 1986. Phytochemical aspects Beijing. (In Chinese.) of phylogeny in Hamamelidae. Ann. Mis­ Cheng, J.Q., J.J. Yang & P. Liu.1985. Prop­ souri Bot. Gard. 73: 417-437. erties and application of mainly commercial Gilbert, S.G. 1940. Evolutionary signifi­ wood. In: J.Q. Cheng (ed.), Wood Sci­ cance of ring porosity in woody angio­ ence. Chinese Forestry Publ. House, Bei­ sperms. Bot. Gaz. 102: 105-120. jing. (In Chinese.)

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