IAWA Journal, Vol. 14 (1), 1993: 87-102

WOOD ANATOMY OF TREES AND SHRUBS FROM CHINA. V. ANACARDIACEAE

by

Zhongmin Dong1 and Pieter Baas Rijksherbarium/Hortus Botanicus, P.O. Box 9514, 2300 RA Leiden, The Netherlands

Summary The wood anatomy of twenty species be­ Xinying 1986; Deng & Baas 1990; Zhang Ion ging to eleven genera of Anacardiaceae Shu-Yin & Baas 1992; Zhong et al. 1992). native to or commonly cultivated in China is The Anacardiaceae are trees, shrubs, woody described in detail and a generic wood ana­ climbers and very rarely herbs, occurring tomical key is given. The wood anatomical mostly in the tropical and subtropical regions diversity pattern partly agrees with the tradi­ throughout the world and a few extending in­ tional classification into the tribes Anacardieae to the temperate zones. The familyis of con­ (Anacardium, Buchanania, and Mangijera), siderable economic value. It produces edible Spondieae (Choerospondias, Dracontomelon, fruits, gums and resins, tannins, dyes, drugs Lannea, and Spondias) and Rhoideae (Coti­ and also several timbers of commercial im­ nus, Pistacia, Rhus, and Toxicodendron). portance. The Chinese lacquer ware is derived Rhus, Cotinus and Pistacia are largely ex­ from Toxicodendron vernicijluum and allied tratropical and share a syndrome of ecologi­ species. Many of the plants of the lacquer cally correlated characters: spiral vessel wall producing group of the family are also poi­ thickenings (at least in the narrow vessels), sonous and cause bad skin allergy. oblique to dendritic latewood vessel distribu­ The classification and delimitation of the tion, ring-porosity, and narrow fibres. family is still a matter of debate. For this Within Toxicodendron two wood anatomi­ study we have adopted the concept of Cron­ cal types are apparent coinciding with a tem­ quist (1981), including Pistacia in the tribe perate (T. vernicijluum) versus tropical (other Rhoideae. species studied) distribution. Yet the generic Wood sampIes of eleven out of about six­ status of Toxicodendron, distinct from Rhus, teen genera of Anacardiaceae occurring in is supported by its frequent coalescent inter­ China (native or commonly cultivated) were vessel pit apertures. The ring-porous T. ver­ available for this study. The remaining five nicijluum moreover lacks the clustering or genera, Dobinea (of doubtful affinity), Dri­ oblique patterning of latewood vessels typical mycarpus, Semecarpus, Terminthia and Pegia of ring-porous species of Rhus. are represented by a single or a few species in Key words: Anacardiaceae, systematic wood China, and we did not succeed in obtaining anatomy, wood identification, ecological wood samp1es of these taxa. wood anatomy, China. Material and Methods Introduction Wood sampies were collected by the first This paper is part of aseries, in wbich the author and obtained from various institutional wood anatomy of trees and shrubs native to wood collections in China as follows: PUw: China is described and discussed family by Departrnent of Biology, Peking University, family. The general outline of this study is Beijing; CAFw: Beijing Institute of Forestry provided in previous papers (Baas & Zhang of Academia Sinica of Forestry; NFIw: North-

1) Present address: Department of Biology, Carleton University, Ottawa, Canada KIS 5B6.

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western Forestry Institute, YangIing; NUw: pairs or in groups of three or four, 20% soli­ Department of Biology, Northwest Univer­ tary, somewhat rounded in outline, tangential sity, Xian. diameter 135 (70-200) )lm, walls 3-4 iJm Herbarium vouchers associated with the thick. Vessel members 400 (250-540) iJm sampies from NUw were identified by Prof. long. Perforation plates simple. Intervessel Hu, Department of Biology, Northwest Uni­ pitting nonvestured, alternate, bordered, poly­ versity, Xian. The sampies from PUw were gonal, 10-12 )lm in diameter with slit-like collected by a group of plant taxonomists and apertures. Vessel-ray pits large and simple, all the herbarium vouchers of them are depos­ round to oval. Vessel-parenchyma pits simple ited in South China Botany Institute (IBC). or with strongly reduced borders in clusters Other wood sampies are unvouchered, or of 2-5. Tyloses abundant. Fibres 800 (650- data referring to the herbarium voucher were 950) iJm long, nonseptate, average diameter not available. In the nomenclature we follow­ 17 )lm, very thin-walled, pits minutely border­ ed Flora Reipublicae Popularis Sinicae, Vol. ed, mainly confined to the radial walls. Paren­ 45 (1) (Ming 1980). chyma vasicentric and weakly confluent, in Microscopic slides and macerations were 3-4-celled strands. Rays mainly uniseriate, made according to standard techniques (Baas occasionally biseriate, 13/mm, 7-10 (2-25) & Zhang 1986). Sampies for scanning elec­ cells or 0.1-0.9 mm tall with 1 row of mar­ tron microscopy were prepared after Exley ginal upright cell (Kribs' heterogeneous type et al.(1977). The recommended terminology III). Radial canals absent. Silica bodies (7-10 and procedures by the IAWA Committee iJm) and a few large prismatic crystals present (1989) were followed in descriptions and de­ in ray cells (Fig. 4). termining quantitative values. Fibre diameter, Note: Septate fibres were reported by many a character showing an interesting variation authors (Hess 1946; Kribs 1968; Mitchell & pattern within the Chinese Anacardiaceae, Mori 1987) in Anacardium, but in our mate­ was measured in macerations; average values rial no septate fibres were detected. are based on 25 measurements. Buchanania Spreng. (Figs. 5-8; Table 1) Generic wood anatomical descriptions Material studied: B. arborescens (Blume) Explanatory note Blume: Taiwan, PUw 852. - B. latifolia In the descriptions the sequence used in Roxb.: Hainan, PUw 986. - B. microphylla the first paper of this series (Baas & Zhang Engl.: Hainan, CAFw 14560. 1986) has been adopted. Besides the features Evergreen (B. arborescens) or deciduous mentioned in that paper, several other features (B.latifolia) trees from tropical forests. are observed and described here: average fibre Growth rings absent or faint. Wood dif­ diameter; radial canals - presence or absence, fuse-porous. Vessels 6-1O/mm2, mostly in shape in tangential section, cell wall thickness radial multiples or clusters, 8-20% solitary, of epithelial cells, number of epithelial cell round in cross section, tangential diameter layers and the diameter of the cavity; silica 115-165 (90-200) )lm, walls 3-4 iJm thick. bodies - presence or absence, shape and size; Vessel members length 400-410 (250-570) pith flecks - groups of large irregularly ar­ iJm in B. microphylla and B.latifolia; 730 ranged parenchyma cells. (630-770) )lm in B. arborescens. Perfora­ tions simple. Intervessel pits nonvestured, Anacardium (L.) Rottboell (Figs. 1-4) alternate, polygonal, or round in B. micro­ phylla, 7-11 )lm in diameter with oval to slit­ Material studied: A. occidentale L.: Hai­ like apertures. Vessel-ray pits and vessel­ nan, PUw 379. parenchyma pits similar, simple to indistinct­ Small trees, introduced and cultivated in ly bordered, round to elongate. Tyloses pres­ tropical regions of China. ent. A few vasicentric tracheids present. Fi­ Growth rings absent. Wood diffuse-por­ bres 1000-1200 (770-1300) )lm long, aver­ ous. Vessels very sparse, 5/mm2, usually in age diameter 19-23 )lm, very thin-walled,

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Table 1. Variation in selected wood anatomical characters of Buchanania.

VD VL IP SF PV SD RW B. arborescens 170 730 P + ± 1-2 B. latifolia 170 400 P + + 1-3 B. microphylla 120 410 R ± + 1-3 VD average vessel diameter (ILm). VL average vessel member length (ILm). IP shape of intervessel pits; P = polygonal; R = rounded. SF septate fibres. PV parenchyma vasicentric to aliform. SD silica bodies. RW= ray width.

with minutely bordered pits mainly confined scanty vasicentric in 3-4-celled strands. to the radial walls. Septate fibres present in Rays 9-12/mm, with 1 row ofmarginal up­ B. arborescens in the vicinity of the vessels, right cells (Kribs' heterogeneous type III), mixed with parenchyma cells. Parenchyma in B. latifolia and B. microphylla 1-3 cells aliform, in 3-4-celled strands in B. micro­ wide, 10-15 (2-30) cells or up to 0.8 mm phylla; or in strands of up to over 8 cells in in height; inB. arborescens, mainly 1-2 cells B. latifolia; in B. arborescens parenchyma (text continued on page 94)

Legends 01 Figures 1-31:

Figs. 1-4. Anacardium occidentale. - 1, 2: TS, TLS, x 40. - 3: TLS showing typical interves­ seI pitting found in the Anacardiaceae; x 250. - 4: RLS showing silica bodies in ray ceIls; SEM, x 1750. - Figs. 5-7. Buchanania arborescens. - 5: TS, x 40. - 6: RLS showing prismatic crys­ tals in ray ceIls; x 250. - 7: TLS, radial canals (arrowed) in rays; x 40. - Fig. 8. Buchanania latifolia; TS, x 40. - Fig. 9. Dracontomelon duperreanum; TLS, x 40.

Figs. 10, 11. Choerospondias axillaris; TS, TLS, radial canal arrowed, x 40. - Figs. 12, 13. Cotinus coggygria var. cinerea; TS, TLS, x 40. - Figs. 14-16. Mangifera sylvatica. - 14,15: TS, TLS, x 40. - 16: Crystals in ray cells; SEM, x 525. - Figs. 17-19. Pistacia. - 17. P. chi­ nensis; TS, x 40. - 18, 19. P. weinmannifolia; TS, x 40, TLS with radial canal, x 100.

Figs. 20-28. Rhus. - 20,23,24,27,28: R. chinensis. - 21,25: R. potaninii. - 22,26: R. punjabensis. - 20-22: TS, x 40. - 23: Spiral thickenings in narrow vessels; SEM, x 900.- 24-26: TLS, x 40. - 27: RLS, druses in subdivided and slightly inflated ray cells, x 250. - 28: Druse in ray ceIl; SEM, x 2000. - Figs. 29-31. Spondias. - 29,30: S. lakonensis; TS and TLS, x 40. - 31: S. pinnata; TLS, x 40.

Figs. 32-40. Toxicodendron. - 32, 33, 36-40: T. succedanea. - 34: T. sylvestris. - 35: T. vernicifluum. - 32-35: TS, x 40, note great difference between the ring-porous T. vernicifluum and the other two species. - 36: RLS showing vessel-ray pitting typical for the Anacardiaceae; x 250. - 37: TLS showing septate fibres, x 250. - 38-40: Coalescent intervessel pits in T. succedanea. - 38: Round pit apertures viewed from the bordered pit chambers; SEM, xI150.- 39: LM view showing sIit-Iike coalescent outer apertures (especially in the vessel wall of which a part has been shaved away at the top of the lower vessel element); round inner apertures also visible; x 250. - 40: Coalescent outer apenures; SEM, x 700.

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wide, a few rays with radial canals 3 cells diameter. Prismatie crystals abundant in en­ wide, 15-20 (2-45) cells or up to 1 mm larged marginal ray cells of PUw 232, absent high. Radial canals in B. latifolia and B. mi­ in NFlw 472. crophylla round in tangential seetion, with 1- 2 (3) layers of partly lignified thin-walled Cotinus (Tourn.) Mill. (Figs. 12, 13) epithelial cells, cavity 35 -40 ).Lm in diameter; in B. arborescens elliptical, with 1-2 layers Material studied: C. coggygria Scop. var. of unlignified very thin-walled epithelial cells, cinerea Engler: Beijing, PUw n.s. cavity 30-40 x 50-110 11m in diameter. Pris­ Deciduous shrubs from temperate regions. matie crystals present in nonchambered ray Growth rings distinct and marked by dif­ cells of B. latifolia and in ray cells, axial pa­ ferences in vessel diameter and fibre wall renchyma cells and septate fibres of B. ar­ thiekness. Wood ring-porous. Earlywood borescens. Round silica bodies, 1O-18).Lm in vessels 36/mm2, 35% solitary, remainder in diameter, one per ray cell present in B. lati­ radial multiples of 2-3, round to oval, tan­ folia andB. microphylla. gential diameter 110 (70-150) 11m; latewood Note: Fundter and Wisse (1977) recorded vessels much more numerous, about 500/ vasieentric, sometimes aliform parenchyma in mm 2, mostly in multiples and clusters in a B. arborescens from New Guinea, but in our radial-oblique to dendritic pattern, angular, material of this species parenchyma is scanty. tangential diameter 25 (15-40) 11m. Vessel walls 3-4).Lm thick. Vessel member length Choerospondias Burtt & Hill (Figs. 10, 11) 330 (260-370) ).Lm. Perforations simple. In­ tervessel pits nonvestured, alternate, round, Material studied: C. axillaris (Roxb.) Burtt 6-9 ).Lm in diameter with oval to slit-like & Hill: Hainan, PUw 232; Yunnan: NFlw apertures. Vessel-ray pits simple, round and 472. small. Spiral thickenings present in all vessel Monotypie genus of deciduous trees in elements. Thin-walled tyloses abundant. Li­ tropical to subtropieal regions. briform fibres 730 (520-81O»).Lm long, aver­ Growth rings distinct and 111arked by dif­ age diameter 11 11m, very thin- to thin-walled, ferences in vessel diameter and fibre wall with minutely bordered pits, confined to ra­ thickness. Wood serni-ring-porous. Vessels dial walls. Parenchyma very scanty para­ 12-15/mm2, mostly in radial multiples of 2 tracheal, in 3-4-celled strands. Rays uniseri­ or 3 or in clusters, 15-20% solitary, round, ate to biseriate, rarely 3-seriate, 15 (2-35) or tangential diameter 150-165 (80-280) 11m, up to 0.7 mm high, with 1-4 rows of square walls 4).Lm thiek. Vessel members 430 (370- to upright marginal cells (Kribs' heterogene­ 520) 11m long. Perforations simple. Interves­ ous type II). Radial canals absent. Prismatic seI pits alternate, polygonal, 11-13 11m in crystals common in marginal and procumbent diameter with slit-like apertures. Vessel-ray ray cells. pits and vessel-parenchyma pits simple or with strongly reduced borders, round. Tyloses Dracontomelon Blume (Fig. 9) present, butrare and small. Fibres 1010 (800- 1190) ).Lm long, average diameter 26-27 11m, Material studied: D. duperreanum Pierre: very thin-walled, with minutely bordered pits Hainan, PUw 446. mainly confined to the radial walls. Septate Evergreen trees from tropical regions. fibres present, mixed with nonseptate fibres. Growth rings absent. Wood diffuse-por­ Parenchyma in narrow marginal bands and ous. Vessels 3-4/mm2, 70% solitary, re­ scanty paratracheal, in 5-1O-celled strands. mainder in short radial multiples, round, tan­ Rays 4-6/mm, 1-4 cells wide, 10-20 (2- gential diameter 160 (120-250) 11m. Vessel 40) cells or up to 0.6 mm high, with 1 row of members 480 (350-600) ).Lm long, the walls marginal upright cell (Kribs' heterogeneous 2.5 ).Lm thick. Perforations simple. Interves­ type III). Radial canals round in tangential sec­ seI pits nonvestured, alternate, polygonal, tion, with 1-2 layers of partly lignified thin­ 8-1O).Lm in diameter with slit-like apertures. walled epithelial cells, cavity 35-60 11m in Vessel-ray pits and vessel-parenchyma pits

Downloaded from Brill.com10/09/2021 03:51:19AM via free access Dong & Baas - Wood anatomy of Anacardiaceae from China 95 simple or with strongly reduced borders, Growth rings faint, sometimes marked by round. Thin-walled tyloses present. Fibres denser bands of fibres and sometimes indi­ 1100 (800-1300) 11m long, average diameter cated by narrow to moderately wide bands of 23 11m, thin-walled, with simple or indistinct­ parenchyma. Woods diffuse-porous. Vessels ly bordered pits mainly confined to the radial 3/mm2 in M. sylvatica and 8/mm2 in M. walls, all septate, the septa extremely fine. Pa­ indica, 15-30% solitary, remainder in radial renchyma vasicentric, in 6-8-celled strands. multiples of 2-3, round, tangential diameter Rays 6/mm, mainly 3-4 cells wide, occa­ 110-140 (80-190) 11m, walls 311m thick. sionally uniseriate or biseriate, 10-20 (2-35) Vessel members 420-480 (300-560) 11m cells or up to 0.8 mm high, with 1 row of long. Perforations simple. Intervessel pits marginal upright cell (Kribs' heterogeneous nonvestured, alternate, polygonal, 7-11 11m type III). Radial canals absent. Prismatic crys­ in diameter. Vessel-ray and vessel-parenchy­ tals in marginal and procumbent ray cells. ma pits large and simple or with strongly re­ duced borders, the apertures rounded to irreg­ Lannea A. Rich. ular in shape. Fibres 790-990 (600-1200) 11m long, nonseptate, average diameter 16-21 Material studied: L. coromandelica (Houtt.) 11m, very thin-walled, with simple to minutely Merr.: Hainan, PUw 509. bordered pits mainly confined to radial walls. Deciduous trees from tropical forests. Parenchyma generally paratracheal, aliform, Growth rings faint. Wood diffuse-porous. occasionally to frequently also confluent, in Vessels 19/mm2, 35% solitary, remainder 3-4-celled strands; also in zonate (marginal?) in radial multiples and clusters, round, tan­ bands, 2-6 ceIIs wide. Rays 8-10/mm, uni­ gential diameter 110 (60-150) 11m, walls 3 seriate or biseriate, 7-15 (2-20) cells or up 11m thick. Vessel members 470 (320-650) to 0.6 mm high, with 1 row of marginal up­ 11m long. Perforations simple. Intervessel right cell (Kribs' heterogeneous type III). pits alternate, polygonal, 7-9 11m in diameter Radial canals absent. Prismatic crystals pres­ with slit-like apertures. Vessel-ray and vessel­ ent in marginal and procumbent ray cells of parenchyma pits simple or with strongly re­ M. sylvatica. duced borders, oval to round. Tyloses pres­ ent. Fibres 1060 (700-1320) 11m long, aver­ Pistacia L. (Fig. 17-19) age diameter 22 11m, very thin-walled, sep­ tate, with simple pits mainly confined to the Material studied: P. chinensis Bunge: Hai­ radial waIIs. Parenchyma in narrow bands nan, PUw 004. - P. weinmannifolia J. Pois­ and scanty para tracheal, in 5-8-celled strands. son ex Franch.: Yunnan, CAFw 16321. Rays 7-8/mm, 1-5 cells wide, 10-20 (2- Evergreen (P. weinmannifolia) or decidu­ 30) cells or up to 0.6 mm high, with 1 row ous (P. chinensis) trees in subtropical to tem­ of upright marginal ceII (Kribs' heterogene­ perate regions. ous type III). Radial canals round in tangen­ Growth rings marked by concentric lines tial section, with one layer of thin-walled ofrelatively large vessels. Wood ring-porous partly lignified epithelial ceIls, cavity 20-30 (P. chinensis) or semi-ring-porous (P. wein­ 11m in diameter. Prismatic crystals infrequent mannifolia). Vessels 150-210/mm2, in P. in procumbent ray cells. Silica bodies (10-15 chinensis of two distinct sizes, earlywood 11m) present in ray ceIIs. vessels solitary with tangential diameter of 145 (100-200) 11m, latewood vessels 30 (18-45) Mangifera L. (Figs. 14-16) 11m in tangential diameter in extensive clusters and forming a diagonal pattern; in P. wein­ Material studied: M. indica L.: Hainan, mannifolia, vessels 50 (30-100) 11m in tan­ PUw 485. - M. sylvatica Roxb.: Hainan, gential diameter in extensive clusters and PUw 284. radial pattern; walls 3-4 11m thick. Vessel Evergreen trees, introduced and cultivated member length 320 (250-390) 11m. Perfora­ (M. indica) or native (M. sylvatica) in tropi­ tions simple. Intervessel pits nonvestured, cal regions. alternate, round or partly polygonal, 7-9 11m

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in diameter with round (P. chinensis) or slit­ Rhus (Tourn.) L. emend. Moench. (Figs. like (P. weinmannifolia) apertures. Vessel-ray 20-28; Table 2) pits with reduced borders, round. Vessel-pa­ renchyma pits of two types: distinctly border­ Material studied: R. chinensis Mill: Hai­ ed ones 5 J.lm in diameter and simple ones nan, PUw 165; Shaanxi, NUw 901; Zhejiang, 2-3 J.lm. Spiral thickenings present in narrow CAFw 6615. - R. potaninii Maxim: Beijing, vessels, absent from wide vessels. Thin-wall­ CAFw 13534; Shaanxi, NFlw 83102. - R. ed tyloses common. Fibres 760 (410-1060) punjabensis Stewart: Shaanxi, NUw 902. J.lm long, nonseptate, average diameter 12-13 Deciduous trees, in tropical to temperate J.lm, thick to very thick-walled, with rninutely regions. bordered pits mainly confined to the radial Growth rings distinct and marked by dif­ walls; pit apertures round in P. chinensis and ferences in vessel diameter and fibre wall slit-like in P. weinmannifolia. Parenchyma thickness. Wood ring-porous to semi-ring­ scanty paratracheal, in 3-4-celled strands. porous. Earlywood vessels 12-22/mm2, Rays 7-lO/mm, with one row of marginal mostly solitary, remainder in radial multiples upright cell (Kribs' heterogeneous type III), of 2-3, round to oval, tangential diameter in P. chinensis of two distinct sizes, uniseri­ 120 (60-200) J.lm; latewood vessels much ate rays 1-10 cells high, 3-4-seriate 15-20 more numerous (over 100/mm2), mostly in (10-30) cells or up to 0.4 mm high; in P. multiples and clusters, often in oblique to weinmannifolia, mainly 3-5-seriate, those dendritic pattern, angular, tangential diameter with radial canals 9-10-seriate, 9-25 cells 15-30 J.lm. Vessel walls 3-5 J.lm thick. Ves­ or up to 0.35 mm high. Radial canals round seI member length 290-410 (200-500) J.lm. in tangential section, in P. chinensis, with Perforations simple in oblique to almost hori­ 1-2 layers of unlignified very thin-walled zontal end walls. Intervessel pits nonvestured, epithelial cells, cavity 40-50 J.lm in diameter; alternate, polygonal, rarely round, 7-10 J.lm in P. weinmannifolia, with 1-2layers of lig­ in diameter with oval to slit-like apertures. nified thin-walled epithelial cells, cavity 35- Vessel-ray and vessel-parenchyma pits sim­ 40 J.lm in diameter. Prismatic crystals present ple, large and round. Spiral thickenings weIl in ray cells. In P. weinmannifolia, a few crys­ developed in narrow vessels, absent from tals noted in ty loses. earlywood vessels of R. chinensis but present

Table 2. Variation in selected wood anatomical characters of Rhus.

VD VL ST FL RT

R. chinensis NUw 901 90 290 +- 500 III CAFw6615 120 340 +- 720 III PUw 165 130 360 +- 730 III R. potaninii CAFw 13534 120 410 ++ 810 II NFlw 83102 140 ++ II R. punjabensis NUw 902 140 330 ++ 670 III

VD average earlywood vessel diameter (J.lm). VL average vessel member length (J.lm). ST spiral thickenings in vessels; +- = present in narrow vessels, absent in wide vessels; ++ = present in all vessel members. FL average fibre length (J.lm). RT ray type (Kribs' heterogeneous types II or III).

Downloaded from Brill.com10/09/2021 03:51:19AM via free access Dong & Baas - Wood anatomy of Anacardiaceae from China 97 in all vessel elements of R. potaninii and R. tangential seetion, with 1-2layers ofunligni­ punjabensis. Thin-walled tyloses often pres­ fied very thin-walled epithelial cells surround­ ent. Libriform fibres 660 (410-990) 11m ed by 1-3 layers lignified thin-walled small long, nonseptate, average diameter 16-21 cells, cavity 50-60 11m in diameter. Prismatic 11m, very thin-walled, with minutely bordered crystals present in ray cells of S. pinnata. pits mainly confined to the radial walls. Pa­ renchyma scanty paratracheal, and in discon­ Toxicodendron (Tourn.) Mil!. (Figs. 32-40; tinuous uniseriate marginal bands, in 5-8- Table 3) celled strands. Rays 5-7 (4-9)/mm, usual­ ly 1-3 cells wide, in R. punjabensis up to 6 Material studied: T. succedaneum (L.) O. cells wide, 7-18 (1-35) cells or up to 0.6 Kuntze: Hainan, PUw 571 & 044. - T. syl­ mm high, with 1 or 2-4 (R. potaninii) rows vestre (Sieb. & Zucc.) O. Kuntze: Hainan, of square to upright marginal cells (Kribs' PUw 762. - T. vernicifluum (Stokes) F.A. heterogeneous type III and II), of two dis­ Bark!.: Shaanxi, NUw 903. tinct sizes in R. punjabensis (1-2-seriate and Deciduous trees in tropical to temperate 4-6-seriate). Radial canals absent. Druses regions. present in ray cells of all sampies. Prismatic Growth rings distinct in T. vernicifluum crystals also present in ray cells. Pith flecks and faint in other species. Wood ring-porous frequent in R. chinensis. in T. vernicifluum; diffuse-porous in the other two species. Vessels 9-18/mm 2, 14-50% Spondias L. (Figs. 29-31) solitary, round, tangential diameter 235 (160- 300) 11m in earlywood, 40 (25-65) 11m in Material studied: S.lakonensis Pierre: Hai­ latewood of T. vernicifluum; 105 (50-160) nan: CAFw 14588. - S. pinnata (L. f.) Kurz: 11m in other sampies, walls 3-7(-9) 11m Yunnan, CAFw 13635. thick. Vessel member length 310-480 (250- Deciduous trees in tropical forests. 590) 11m. Perforations simple. Intervessel pits Growth rings absent. Wood diffuse-por­ nonvestured, alternate, mainly polygonal but ous. Vessels 3-7/mm2, 40-60% solitary, round in T. vernicifluum, 7-10 11m in diam­ round, remainder in short radial multiples, eter, very often with long coalescent slit-like tangential diameter 150-160 (100-200) 11m. inner apertures (Figs. 39, 40) and round (T. Vessel members 460-560 (300-750) 11m vernicifluum) or oval outer apertures (Fig. long walls 2.5 11m thick. Perforations simple. 38). Vessel-ray and vessel-parenchyma pits Intervessel pits nonvestured, alternate, poly­ large and simple. Spiral thickening developed gonal,large, 11-14 11m in diameter with oval only in the narrow vessels ofT. vernicifluum. to slit-like apertures. Vessel-ray and vessel­ Fibres 720-955 (600-1100) pm long, very parenchyma pits similar, large, round and thin to thin-walled, with minutely bordered simple. Thin-walled tyloses common. Fibres pits mainly confined to the radial walls, partly 770 (600-900) 11m (S. lakonensis) or 1250 septate in T. succedaneum and T. sylvestre, (800-1350) 11m (S. pinnata) long, average all nonseptate in T. vernicifluum. Parenchy­ diameter 33-34 11m, very thin-walled, with ma scanty paratracheal in T. vernicifluum and minutely bordered pits confined to radial vasicentric in others, and in narrow marginal walls. Septate fibres mixed with non-septate bands, in 4-8-celled strands. Rays 5-7 (3- fibres. Parenchyma scanty paratracheal in 8)/mm, usually 1-3 (or up to 12) cells wide, 3-4- (S. lakonensis) or 5-8- (S. pinnata) 7-25 (2-35) cells or up to 0.6 mm high, celled strands. Rays with one row of upright with I or 1-3 (T. vernicifluum) rows of up­ marginal cell (Kribs' heterogeneous type III); right marginal cells (Kribs' heterogeneous in S. pinnata, 3-5/mm, 1-6 cells wide, type III or II). In sampie PUw 571 of T. suc­ mainly 3-4 cells wide, 15-25 (1-40) cells or cedaneum, rays are of two distinct sizes; the up to 1 mm high; in S.lakonensis, 5-7 I mm, large ones 10-12 cells wide. Radial canals mainly 1-3 cells wide, a few rays with radial absent. Large prisrnatic crystals present in canals 4-6 cells wide, 9-17 (1-40) cells or marginal and procumbent ray celIs, or only in up to 0.9 mm high. Radial canals round in marginal ray cells (PUw 044).

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Table 3. Distinguishing wood anatomical characters of Toxicodendron. (Note large differences between T. vernicifluum and the other species.)

R D ST VD VL VWT SF PV MP RT T. succedaneum PUw 571 + 100 410 4-6 + + + III PUw 044 + 100 480 7-9 + + + III T. sylvestre + 100 390 7-9 + + + III T. vernicifluum + + 240* 310 3 ± II R ring-porous. D diffuse-porous. ST spiral thickenings in vessels. VD average vessel diameter ()lm); * of earlywood vessel only. VL average vessel member length ()lm). VWT vessel wall thickness. S F septate fibres. PV parenchyma vasicentric. MP marginal parenchyma bands. RT ray type (Kribs' heterogeneous types II or III).

Classification and identification of the wood anatomical features more frequently Anacardiaceae from China than others. In Table 4 the qualitative wood anatomical Presence or absence of radial canals in the features of Anacardiaceae from China are rays is a constant character at the generic summarised. The taxa are arranged according level. Canals occur in all tribes represented in to the treatment ofthis family by Ming (1980) China, but are absent from Anacardium, Dra­ and several other authors. There is a fair contomelon, Mangifera, Rhus, Cotinus, and amount of character association within the Toxicodendron. This observation is support­ tribes, but none of the character combinations ed by the wood anatomical descriptions of is absolutely constant. The Anacardieae all Anacardiaceae from outside China (Kanehira are diffuse-porous, have very thin-walled 1921; Pearson & Brown 1932; Heimsch fibres of intermediate diameter, and lack spi­ 1940, 1942; Metcalfe & Chalk 1950; Ghosh ral thickenings or oblique vessel patterning; & Purkayastha 1963; Young 1974; Baas, in in other features such as septate fibres, pa­ Hou 1978). Because of its constancy, this renchyma distribution, radial canals, and pres­ character serves as a very important criterion ence of crystals and silica bodies, the tribe is for delimitation of genera, but not of higher heterogeneous. The Spondieae are constant or lower categories. The distribution of radial for presence of septate fibres but vary for canals in rays among the tribes and genera most other features (Table 4). The Rhoideae and their common occurrence in the related mostly have ring-porous wood, spiral thick­ family of the Burseraceae suggests that they enings, an oblique (latewood) vessel pattern may be a primitive (plesiomorphic) feature in and marginal parenchyma, but there are ex­ the family wh ich was subsequently lost sev­ ceptions and other features tend to be con­ eral times during evolution. stant for an individual genus or some genera The oblique to dendritic vessel distribution rather than for the tribe, e.g., narrow fibres in the latewood is invariably associated with in Cotinus and Pistacia; druses in Rhus; co­ ring-porosity and presence of spiral vessel alescent apertures in Toxicodendron. Similar wall thickenings. The distribution of this syn­ findings were discussed by Heimsch (1942) drome in the Anacardiaceae from China shows who studied species from outside China and a rather distinct systematic pattern. It is re­ noted that there are only some trends for each stricted to tribe Rhoideae where it occurs in tribe of the Anacardiaceae to show particular all genera except T oxicodendron. In the latter

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Table 4. Wood anatomicaI diversity in Anacardiaceae from China.

~ ~» 2 .t:: ~ u Vl c:: »e :§' .t:: Vl ~ u C ~ ~ Po. ~., .,Vl ~ .,~ ~ Vl § ~ § Vl 0/) ~ § :a .S I j ~ u .~ 3Vl ~ .,c:: i Vl 8 .0 .,Vl ~ :s u i'l ·li «= ~ .0 b' ., > «= :;a 'E j u i :s ., ! u .::1 ] B ~ !3 ~ '" .~ 0- B ] ::::'" g, e ~ ~ ~ 8 r .0 .~ ~ :::: Tribe: Anacardieae > Po. ., ~ '" B &: ...:: CI) 0 CI) ~ > ~ ~ it ~ Vi .Ei Anacardium occidentale D II A + + + Buchanania arborescens D II + A + + latifolia D II A + + + + microphylla D II A + + + Mangifera indica D II A + + sylvatica D II A + + +

Tribe: Spondieae Choerospondias axillaris NFIw 472 R III + A + + axillaris PUw 232 R III + A + + + Dracontomelon duperreanum D II + B + + Lannea coromandelica D II + A + + + + Spondias lakonensis D III + A + pinnata D III + A + +

Tribe: Rhoideae Rhus chinensis R II A ± + potaninii R II + ± A ± + punjabensis R II + ± A ± + m Toxicodendron U vernicifluum R II + A ± + - succedaneum PUw 571 D 1I + B + + + succedaneum PUW 044 D 1I + A + + + sylvestre D II + A + + + m Cotinus n coggygria var. cinerea R + + B ± + Pistacia chinensis R + + C + + weinmannifolia S + + C + +

Legend: + = present; - = absent; ± = intermediate; D = diffuse-porous; S = semi-ring-porous; R = ring-porous; 1=< 13 Jlm; II = 16-23 Jlm; III = 26-34 Jlm; A = very thin; B = thick; C = very thick.

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genus only the temperate species T. vernici­ 9a. Marginal parenchyma present, silica fluum has spiral thickenings and ring-porous bodies absent ...... Mangifera wood but lacks an oblique to dendritic vessel b. Marginal parenchyma absent, silica pattern in the latewood. bodies present ...... Anacardium A dichotomous key to the woods of Chi­ lOa. Marginal parenchyma present, interves­ nese Anacardiaceae used in this study is given sei pits mostly coalescent below. It is possible to key out genera of Toxicodendron p.p. Anacardiaceae woods from China. It has not b. Marginal parenchyma absent, interves­ been attempted to separate individual species, sei pits without coalescent apertures because the material studied is too Iimited and Dracontomelon most genera are fairly homogeneous except lla. Fibre walls thick, average fibre diam- Toxicodendron, where two distinct groups eter< 13)lm ...... Cotinus can be distinguished wood anatomicaIly. For b. Fibre walls very thin, average fibre the separation of species of Buchanania, see diameter> 15)lm ...... 12 Table 1. 12a. Latewood vessels in extensive clusters and usually in an oblique to dendritic Generie wood anatomieal key to the pattern, intervessel pits without coales- Anacardiaceae from China cent apertures ...... R hus b. Latewood vessels solitary or in short 1a. Radial canals present in rays, sometimes radial multiples, intervessel pits mostly very rare...... 2 with coalescent apertures b. Radial canals absent ...... 7 Toxicodendron p.p. 2a. Spiral thickenings present in vessels, fi- bre waIls thick ...... Pistacia The taxonomie status of Toxicodendron b. Spiral thickenings absent, fibre walls verythin ...... 3 Barkley (1937) recognised Toxicodendron 3a. Wood ring-porous '" Choerospondias as a genus, distinct from Rhus, a conclusion b. Wood diffuse-porous ...... 4 which was also supported by Heimsch (1940). 4a. Parenchyma scanty, septate fibres pres- Brizicky (1963), disagreeing with Barkley and ent ...... 5 Heimsch, again relegated Toxicodendron to b. Parenchyma vasicentric to aliform, sep­ a subgenus of Rhus. GiIIis (1971) in a com­ tate fibres absent .... Buchanania p.p. prehensive study of Toxicodendron has 5a. Rays (excluding those containing canals) shown that it should be treated as aseparate uniseriate or biseriate Buchanania p. p. genus. Our study shows, however, that b. The widest rays more than 3-seriate. 6 Toxicodendron is, wood anatomie

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has radial canals in its rays. Radial canals tropieal, and constitute the most widespread have been reported in the rays of some spe­ tri be of the family and especially Rhus, Co­ eies of Toxicodendron (Record 1939), al­ tinus and Pistacia cover wide ranges. In though they were 110t observed in this study. China, these genera are distributed in a con­ Heimsch (1940) and Young (1974) also fail­ tinuous zone from warm temperate (N 40°) to ed to detect radial canals in the rays of T oxico­ tropical latitudes (N 19°). Tallying with the dendron and it is likely that they do not occur distribution, Rhus, Cotinus and Pistacia share in this genus. a syndrome of characteristics: spiral vessel The wood structure of T. vernicifluum wall thickenings, oblique to dendritic late­ superficially resembles that of Rhus. Spiral wood vessel distribution, ring-porous wood thiekenings in the small vessels were observed and narrow fibres. only in T. vernicifluum and T. trichocarpum In Chinese Anacardiaceae, spiral thicken­ (Heimsch 1940). This does not agree with ings are mostly found in temperate plants, Record (1939), who stated that spiral thicken­ Rhus, Cotinus and Pistacia. This is in line ings are present in the small vessels of all the with the commonly established ecological species. This may be due to the different def­ trends (Baas 1986; Baas et a!. 1983; Carlquist initions of the character and probably Record 1988). mistook grooves interconnecting the inner pit Wood of seven of the studied genera are apertures (coa1escent pits) for spiral thicken­ without growth rings, of which Anacardium, ings, because coalescent pits are ubiquitous Buchanania, Lannea, Mangifera, S pondias in Toxicodendron. Considering Toxicoden­ and Dracontomelon are tropical genera. In dron as a whole, the presence of coalescent the genus Toxicodendron, all tropical species intervessel pits, and the absence of cIustering lack growth rings. The other five genera have in the small vessels of the latewood set the distinct growth rings; not surprisingly, most genus apart from Rhus. Wood anatomy can of their species are mainly temperate. Only thus be invoked to support Toxicodendron as Choerospondias (Spondias axillaris Roxb. in a separate genus. Ghosh & Purkayastha 1963) is largely tropi­ ca!. Checking the latitude of provenance of woods in these two groups, it appears that in Ecological trends in the woods of Ana­ the group without growth rings only one sam­ cardiaceae in China pIe is from 25° N, aII others are from 19° N; in We have realised the risk of searching for the group with growth rings, all five genera ecologieal trends in such a lirnited number of have some species from high latitudes. In our species, of which moreover data on the altitu­ material, distinct growth rings are concomi­ dinal range and ecological nie he in the forest tant with deciduousness. were usually not available. However, some Fibre diameter shows an ecological trend qualitative characteristics show an obvious in the Anacardiaceae from China: very wide ecological trend in the present study, and these to medium fibres are found in tropical genera, trends are reinforced when consuiting wood medium to very narrow fibres occur in genera anatomical descriptions of the Anacardiaceae in which there are some species from in other Floras (Metcalfe & Chalk 1950; temperate regions (Rhoideae). Record 1939; Heimsch 1940, 1942; Van der Graaff & Baas 1974; Baas, in Hou 1978; Acknowledgements Mitchell & Mori 1987). Therefore we restrict the analysis of ecological trends in the wood We are gratefuI to Prof. Zhang Xinying, anatomy of Chinese Anacardiaceae to rela­ Peking University and Prof. Liu Peng, Bei­ tionships between the distribution of plants jing Institute of Forestry for providing the and some qualitative wood anatornical char­ wood samples, to Bertie Joan van Hellven, acteristics. Leiden, for her rnicrotechnicaI assistance, and The farnily Anacardiaceae attains its high­ to Ms. Teresa Terrazas, University of North est development in the tropics and subtropics. Carolina, Chapel Hili, and Dr. Ding Hou, The Rhoideae are, however, mostly extra- Leiden, for valuable comments.

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