IAWA Bulletin n.s., Vol. 13 (2),1992: 135-150

WOOD ANATOMY OF THE GROUP (LEGUMINOSAE)

by

R.W. den Outer and W.L.H. van Veenendaal Department of Cytology and Morphology,Agricultural University, Arboretumlaan 4, 6703 BD Wageningen, The Netherlands

Summary The secondary xylem of 5 of the 6 genera The distribution of the three subfarnilies is belonging to the Baphia-group (viz. Airyan­ mainly tropical and subtropical. The Papilion­ tha, Baphiastrum, Leucomphalos, oideae constitute a much !arger group than the andBaphia) and Ormosia (Leguminosae-Pa­ others and is the only one which is common pilionoideae-) was studied, in or­ in temperate regions. It has a worldwide der to aid generic delimitation in the Baphia­ range, with most of the woody representa­ group. tives in the tropics and southern hemisphere. Storied-structure features, vessel-member The herbaceous and shrubby members of the length, structure of rays and axial parenchy­ Papilionoideae usually are found in temperate ma, and presence of crystals were used to regions, with a high concentration in mediter­ discuss generic delimitation. Based on the ranean regions (Heywood 1971). The Mimos­ number of cells per axial parenchyma strand oideae are woody; the Caesalpinioideae are two groups are recognised: Airyantha and primarily woody and rarely herbaceous. Baphiastrum have 2-4 cells, Leucomphalos, The Papilionoideae consist of 10 to 11 Dalhousiea and Baphia have fusiform paren­ tribes into which the genera are grouped on chyma or 2-celled strands. Ormosia is very features of habit, leaf morphology and degree different from the other genera which c10sely of fusion of the stamens (Chant 1979). The resemble each other. A c1ear wood anatomi­ position ofthe tribe Swartzieae, if included in cal separation of the genera belonging to the the Papilionoideae, and not in the Caesalpini­ Baphia-group is not possible. Especially oideae as is often done, is debatable. Also Airyantha and Baphiastrum are almost iden­ Sophoreae s.l. is a tribe of convenience be­ tical wood anatomically and could be com­ tween the Caesalpinioideae and the bulk of bined into a single genus. There are only the Papilionoideae, and not sharply defined minor differences between Leucomphalos, from either subfarnily. The Sophoreae are Dalhousiea and Baphia. mostly trees, shrubs, lianas or, rarely, herbs. Key words: Airyantha, Baphia, Baphiastrum, The leaves are pulvinate, pinnately I-many­ Dalhousiea, Leucomphalos, Ormosia, sys­ foliolate or digitately 3-foliolate, sometimes ternatic wood anatomy. stipellate; stamens free (Polhill 1981). The tribe contains nearly fifty genera, most of which are small and discrete, except for Introduction Baphia, Ormosia and . Sophoreae The Leguminosae constitute a relatively are divided into 8 groups, of which only the natural taxon which can be divided into three Sophora-group extends significantly outside subfarnilies, viz. Caesalpinioideae, Papilion­ the tropics. The Baphia-group, here consid­ oideae and Mimosoideae. It is a matter of ered, consists of 6 genera. Five genera (Airy­ opinion whether these three groups should be antha, Baphiastrum, , Dalhousiea treated as subfarnilies or as separate farnilies: and Leucomphalos) only include a few spe­ Caesalpiniaceae. Papilionaceae and Mimosa­ eies (2, 2, 4, 3 and 1 respectively), whereas ceae (Hutchinson 1964). Baphia includes c. 60 species (Polhill1981).

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Several plant taxonomists feel that the mated in cross sections; the remaining part is number of genera in the Baphia-group has to occupied by both vessels and ray tissue. The be reduced. The present investigation was amount ofaxial parenchyma is given as a undertaken to find wood anatomical argu­ percentage of the ground tissue and of the ments for or against such areduction. Sam­ total cross sectional area. pIes of 5 genera of the Baphia-group were We have used the definition of libriform available; of Bowringia no sampies could be fibres given by Reinders (1935) and Jans­ obtained. Because Airyantha might belong sonius (1940), which in this case equals the to the Ormosia-group (with 3 genera, viz. description ofthe IAWA Committee (1989). Ormosia, Pericopsis, Haplormosia) of the Ends cells ofaxial parenchyma strands Sophoreae, some Ormosia species were also seen in tangential seetions may be gable- or investigated and compared with descriptions conical-shaped (Reinders-Gouwentak & Rijs­ given by Reinders-Gouwentak and Rijsdijk dijk 1955). Near the top of gable-ended eells (1968). Wood anatomie al evidence is often the longitudinal walls more or less abruptly useful in matters of generic, tribal and sub­ bend into an oblique plane forming a roof­ familial delimitation. However, differences at like top or gable (see Figs. 6, 16, 19). The the species level generally are less clearly ex­ angle of the gable may be obtuse but often is pressed in the wood than in external features slightly less than 90 degrees; the angle at the (Bareua-Kuipers 1981). abrupt bending is always obtuse. In the top part of conical ended eells the longitudinal walls gradually bend towards each other Materials and Methods forming a cone-shaped cell top (see Fig. 24); With the exception of Ormosia sampies, the angle of the top is always sharp, often this investigation is based on material from mueh less than 90 degrees. When the angle tropical Africa. The species studied are listed of the gable is small (about 30 degrees), the below, arranged alphabetically, viz. Airyan­ angle at the abrupt bending is large (about tha with 1 species and 7 sampies; Baphia 165 degrees) and then the end cells of the with 20 species, 41 sampies; Baphiastrum parenchyma strands are almost conical­ with 1 species, 5 sampies; Dalhousiea with 1 shaped and conical ones are then always also species, 5 sampies; Leucomphalos with 1 spe­ present in the species. The greater the angle eies, 2 sampies; Ormosia with 5 species, 5 of the gable-end, the smaller the abrupt bend­ sampies. In total 65 sampies belonging to 29 ing angle, and the smaller the chance that also species and 6 genera. conical-ended top cells are present in the spe­ Anatomical features were studied in trans­ eies. verse, radial and tangential seetions and in Combination types of rays are according macerations. All seetions were embedded in to Baretta-Kuipers (1973) based on features Kaiser's gelatin-glycerin. Means and ranges described by Reinders-Gouwentak and Rijs­ of the different eharaeters are based on twenty­ dijk (1955) and Reinders-Gouwentak (1955). five or more individual measurements. Tan­ Rays investigated by her and by us are nearly gential vessel diameters are given and vessel always homocellular, composed of procum­ member length was measured including the bent cells, sometimes with one or a few rows tails. of nearly square or upright cells at the mar­ In the wood description quantitative data ginal parts. Rays may be either uniseriate, are given as a range of mean values of the or mostly uniseriate and partly multiseriate, different sampies and/or species usually fol­ or almost entirely multiseriate with short lowed between braekets by extremes of all uniseriate marginal parts. In some speeies the sampies together, or as a single mean these different kinds of rays oceur together, value if there is no range (mean values are in others two of the three kinds are found. equal for the different sampies of a species or The ratio of the three kinds of rays can be for the different species of a genus). used for differentiating between the species. Ground tissue percentages for libriform Baretta-Kuipers (1973) distinguished four fibres and axial parenehyma together are esti- different combinations, viz.:

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- combination I, in which the uniseriates Tw & Uw, Dechamps 1410, Angola, small dominate (more than 80%) and the partly tree, 8 cm; Tw, FHOw 18235, Zambia, or entirely multiseriates constitute a small small tree, 5 cm. - B. cf. descampsii De part of the whole; Wild.: Kw, Jefford & Newbould 2071, Tan­ - combination 11, in which most of the rays zania, shrub, 4 cm. - B. dewevrei De Wild.: are uniseriate (60-80%), but quite a num­ Tw, Dechamps 1334, Congo, shrub, 2 cm.­ ber are partly or whoIly multiseriate; B. gracilipes Harms: RBHw, Jentsch 84, - combination III, in which most of the rays Cameroun, small tree, over 10 cm. - B. lau­ are multiseriate (60-80%), but a consid­ rentii De Wild.: RBHw, Gottwald Z 25, Zaire, erable number are partly or entirely uni­ large tree, 40 cm; Tw, Dechamps 222, Zaire, seriate; small tree, 4 cm. -B. laurifolia BailI.: Tw, - combination IV, in which multiseriates Louis 3619, 7031, Zaire, shrub, both 1.5 dominate (more than 80%) and the partly cm; RBHw, Jentsch 87, Cameroun, shrub, or wholly uniseriates constitute but a smaIl over 1.5 cm. - B. leptobofrys Harms: Tw & part of the total number. CTFw, Floret 559, Cameroun, shrub, 10 cm. -B. marceliana De Wild.: Tw, Dechamps List of studied material 129, Zaire, shrub, c. 10 cm. - B. massaien­ Each name is foIlowed by an abbreviation sis Taub.: Tw, Dechamps 1241, 1276, An­ (according to Stern 1988) of the location of gola, small tree, 5 cm; Uw, Dechamps 1326, wood sampie and herbarium voucher, collec­ Angola, small tree, 12 cm. -B. nitida Lodd.: tor and number, geographical origin, habit, Tw, Chalk, Sierra Leone, small tree, over 5 and stern diameter of the sampie. cm; Tw, Gavage, Gabon, small tree, over 5 CfFw = Division d'anatomie des bois, cm; Tw, Detienne 53, Ivory Coast, small Centre Technique Forestier Tropical, Nogent­ tree, over 5 cm; CTFw, Adam 5758, Guinee, sur-Mame, France; Kw = Economic and Con­ small tree, 7 cm; WAGw, Bos 3390, Camer­ servation Seetion, Herbarium, Royal Botanic oun, small tree, 4 cm; WLw, Versteegh & Gardens, Kew, England; RBHw = Bundes­ Den Outer 47, Ivory Coast, small tree, 7 cm. forschungsanstalt fUr Forst- und Holzwirt­ - B. obovata Schinz: Uw, Seiner 20, Botswa­ schaft, Hamburg, Germany; Tw = Service na, small tree, 10 cm. - B. polyantha Harms: d'anatomie des bois, Musee Royal de l'Afri­ Tw, Donis 2004, Zaire, shrub, 4 cm.­ que Centrale, Tervuren, Belgium; Uw = Sys­ Baphia polygalacea (Hook. f.) Baker: WAGw, tematic Botany, University of Utrecht, The Breteler 1580, Cameroun, liana, 1 cm; Uw, Netherlands; WAGw = Department of Plant Breteler 2991, Cameroun, liana, 2 cm. - B. , Agricultural University, Wage­ pubescens Hook. f.: WLw, Versteegh & Den ningen, The Netherlands; WLw = Depart­ Outer 34, 316, Ivory Coast, small tree, 10 ment of Plant Cytology & Morphology, Agri­ and 20 cm resp.; Tw, Louis 2218, 4032, cult. Univ., Wageningen, The Netherlands. Zaire, small tree, 2.5 and 1 cm resp.; Tw, Airyantha schweinfurthii (Taub.) Brum­ Cellulco 243, Zaire, small tree, c. 6 cm. - mitt [syn. Baphiastrum confusum (Huteh. et B. spathacea Hook. f.: WLw, Versteegh & Dalz.) Pellegr., Baphia schweinfurthii Taub., Jansen 757, Liberia, liana, 2 cm. -B. sylva­ Baphia confusa Huteh. et Dalz.]: WAGw, tica Harms: WAGw, Meyer 15155, Camer­ Leeuwenberg 4232, Ivory Coast, liana, 1.5 oun, liana, 5 cm. -B. verschuerenii De Wild.: cm; Tw, Louis 5997, Zaire, liana, 1 cm; Tw, Donis 2381, 2463, Zaire, shrub, 2.5 WAGw, Breteler 1431 and 1765, Cameroun, and 7 cm resp. - B. zenkeri Taub.: Uw & liana, 3 and 1 cm resp.; WLw, Versteegh & WLw, Breteler 2632, Cameroun, liana, 1 cm. Den Outer 108, 142,551, Ivory Coast, liana, Baphiastrum brachycarpum Harms [syn. 4, 6 and 2 cm, resp. Baphiastrum boonei (De Wild.) Vermoesen, Baphia bancoensis Aubrev.: CTFw & Baphia boonei De Wild.]: Tw, Louis 2274, Tw, Detienne 204, Ivory Coast, small tree, 6062, Zaire, liana, both 1.5 cm; WAGw, over 10 cm. - B. bangweolensis R. E. Fries: Breteler 2457, Cameroun, liana, 3 cm; Uw, Tw, Malaisse 9947, 11087, Zaire, liana to Breteler 2178,2842, Cameroun, liana, 3 and shrub, both 3 cm. -B. bequaertii De Wild.: 2 cmresp.

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Dalhousiea africana S. Moore: WAGw, Vessels solitary and in shon radial multi­ Breteler 6470, Gabon, liana, 2 cm; Tw, ples, usually also present in radial multiples Louis 4134, Zaire, liana, 1 cm; Tw, Wage­ of more than four vessels (in 62% of the mans 1752, Zaire, liana, 2 cm; Tw, Tous­ sampies) and in clusters (in 83%·of the sam­ saint 159, Zaire, liana, 2.5 cm; Tw, Ishikawa pIes); in B. laurifolia (Jentsch 87), B. mar­ et al. 48, Zaire, liana, 1 cm. celiana and B. nitida (Detienne 53) the ves­ Leucomphalos capparideus Bentham ex sels are only solitary; round to oval in cross Planch.: WAGw, Bos ?, Cameroun, a liana­ section; number per square mm 5-42 (3-63) shrub grown in the greenhouse at Wagenin­ [5 in B. laurifolia (Jentsch 87) and B. mar­ gen from seed collected by Bos, 1 cm; celiana to 42 in B. massaiensis (Dechamps WAGw, Louis, Breteler & De Bruyn 1320, 1241)]; tangential diameter 60-140 (15-310) Gabon, liana, 1 cm. 11m (60 11m in B. cf. descampsii and B. de­ Ormosia calavensis Azaola: WLw, for­ wevrei to 140 Jlffi in B. bequaertil); thickness merly Dutch Colonial Forestry Service, BW of walls usually c. 6 11m or less, in B. massai­ 4251, West New Guinea (Irian Jaya, Indo­ ensis 8 11m and very thick-walled (14 11m) in nesia), large tree, over 10 cm. - O. coccinea B. polyantha. Vessel member length 155- Jack.: WLw, Stahe1357, Surinam, large tree, 340 (85-440) Jlffi [155 11m in B. massaiensis over 12 cm. - O. costulata (Miq.) Kleinh.: (Dechamps 1326) to 340 11m in B. gracili­ WLw, Stahel 83, Surinam, large tree, over pes]; perforations exclusively simple, slightly 12 cm. - O. coutinhoi Ducke: WLw, Wel­ oblique to transverse; intervessel pits border­ vaartsfonds WS 1025, Surinam, large tree, ed, vestured, often coalescent, alternate, usu­ over 40 cm. - O.fastigiata Tul.: WLw, Sta­ ally not very crowded, horizontal diameter hel 235, Surinam, large tree, over 12 cm. 4-12 11m (4 11m in B. gracilipes to 12 11m in B. bequaertil). In addition to intervessel con­ Descriptions tact, vessels are usually in contact with axial A general description of the secondary and ray parenchyma, less frequently with xylem of the investigated Baphia sampies, libriform fibres. Vessel-ray pitting half-bor­ subdivided in two habit categories, is given dered, nearly always vestured (observations below; in the description of lianas differences with light microscopy), horizontal diameter with the erect shrubs and trees are emphasised. 4-14 11m (4 11m in B. gracilipes to 14 11m 'in This is followed by shon descriptions of Dal­ B. bequaertii); vessel-parenchyma pits half­ housiea, Leucomphalos, Baphiastrum, Airy­ bordered. Brown contents sometimes present antha and Ormosia, highlighting differences in B. polyantha. with Baphia (see also Table 1). Ground tissue of libriform fibres together with axial parenchyma forming about 80% of Baphia (20 species, 41 sampies, Figs. 1-10) the total cross sectional area; usually also Shrubs and small trees [only B. laurentii some vascular tracheids present in association (Gottwald Z25) is a large tree] (15 species, with clusters. Vessels and rays together 33 sampies) occupy the remaining 20%. Growth ring boundaries generally present Libriform fibres, thin- to thick-walled, (hardly visible in B. laurifolia), distinct (in sometimes very thick-walled, wall thickness 80% of the sampies ) or more or less indistinct 4-9 11m (9 11m in B. obovata), diameter of (in 18% ofthe sampies), marked by terminal lumina 1-4 11m (4 11m in B. verschuerenii); parenchyma and/or smaller radial diameters nonseptate; with simple to minutely bordered of the latewood fibres; wood diffuse-porous. pits with slit-like inner apenures, moderately Storied structure of both rays and axial numerous, common in both radial and tan­ elements (in 25% of the sampies), of only gential walls but usually more frequent in axial elements (in 50% of the samples), or radial ones; intercellular spaces almost always only a very slight tendency of storied struc­ absent; fibres with gelatinous layers present ture ofaxial parenchyma to nonstoried (25% in 38% of the sampies. ofthe sampies); fibres are usually not storied; Parenchyma abundant, forming 25-45% storied rays of adjacent tiers often fused. of the ground tissue [25% in B. gracilipes to

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45% in B.laurifolia (Ientsch 87)], or 20- it is less than 790 J.I.ffi); 4-13 (3-14) per tan­ 36% of the total cross section; diffuse and gential mm (4 in B. bequaertii to 13 in B. lep­ diffuse-in-aggregates in 42% of the sampies; tobotrys). Rays tangentially wider when in long uni- to biseriate tangential bands crossing tangential bands ofaxial parenchy­ (usually terminal) in 56% of the sampies; in ma in B. marceliana; ray parenchyma cells long multiseriate tangential bands 2-7 (1-12) with brown contents in B. bequaertii, B. cells wide (7 cells wide in B. pubescens) in massaiensis and B. pubescens (Louis 2218). all sampies; vasicentric parenchyma or in­ Crystals usually present, absent in B. mas­ complete sheaths around the vessels always saiensis (Dechamps 1241) and B. nitida (Bos present, in B. massaiensis (Dechamps 1241) 3390); prismatic, exclusively solitary; in and B. pubescens paratracheal parenchyma chambered axial parenchyma cells only (61 % more or less aliform. Parenchyma strands of the sampies), in both axial and ray paren­ without crystals and not immediately border­ chyma cells (27% of the sampies), or only in ing upon vessels fusiform or composed of ray parenchyma cells (usually in procumbent two cells, except in B. dewevrei, B. graci­ cells, but also in marginal square or upright lipes and B. obovata where 2-4 cells are cells; 6% ofthe sampies); numberofcompart­ present per strand; end cells of strands ex­ ments per crystalliferous parenchyma strand hibit gable-ends in tangential section or are 4-18 (18 inB.polyantha). almost exclusive1y provided with gable-ends, Pith flecks sometimes present (in 17% of with the exception of B. gracilipes, B. laur­ the sampies). entii (Dechamps 222), B. laurifolia, B. mar­ Tyloses, thick-walled, sometimes present celiana, B. obovata and B. polyantha where in B. pubescens (Louis 2218). gable-ends and conical-ends occur equally frequent; strand length 165-335 Ilm (165 Lianas (5 species, 8 sampies) Ilm inB. obovata to 335 J.I.ffi inB.polyantha). Growth ring boundaries present (hardly Brown cell contents present in B. bequaertii visible in B. zenkeri). and B. massaiensis. Storied structure ofaxial elements (axial Rays both uni- and multiseriate with short parenchyma and vessel elements; fibres usu­ uniseriate marginal parts, but multiseriate ally not storied), but in B. sylvatica of rays ones dominate [71 % of the sampies with com­ and axial elements. bination type IV of Baretta-Kuipers (1973)). Vessels solitary, in short radial multiples, In B. dewevrei, B. laurifolia, B. leptobotrys, in radial multiples of more than four vessels B. pubescens (Louis 2218) and B. verschuer­ and in clusters; in B. bangweolensis mainly enii most rays are multiseriate (combination in clusters; usually of two size classes; num­ type 111), while inB. nitida (V. & O. 47) most ber per square mm 16-135 (10-150) (16 in rays are uniseriate (combination type 11). B. spathacea to 135 in B. bangweolensis); Width 1.5-6 (1-12)-seriate [1.5-seriate in tangential diameter 60-160 (20-240) Ilm B.laurifolia (Ientsch 87) andB. nitida (V. & (60 J.I.ffi in B. bangweolensis and B. sylvatica 0.47) to 6-seriate in B. pubescens (V. & O. to 160 J.I.ffi in B. spathacea); thickness of walls 316)]; composed of predorninantl y procum­ in B. zenkeri 7 Ilm. Vessel member length bent cells with regularly some square and/or 175-370 (140-440) Ilm (175 J.I.ffi in B. bang­ upright cells forrning the uniseriate marginal weolensis to 370 Ilm in B. spathacea). Hori­ parts of the multiseriate rays; in most sampies zontal diameter of intervessel pits and vessel­ 30-50% of the rays have marginal cells which ray pits 6-10 Ilm (61lm in B. sylvatica to 10 are higher than the body cells, less than 30% Ilm in B. bangweolensis and B. poly galacea). in B. dewevrei and B. pubescens (V. & O. Ground tissue of libriform fibres together 316),55% or more inB. gracilipes, B.laur­ with axial parenchyma, forrning about 70% entii, B. laurifolia, B. obovata and B. pubes­ of the total cross sectional area. cens (V. & O. 34); height 180-1100 (35- Libriform fibres with wall thickness of 2100) Ilm (180 Ilm in B. leptobotrys to 1100 4-6 J.1.Ill, diameter of lumina 3 J.LII1; gelatinous J.I.ffi in B. bequaenii; only in B. bequaertii the layers present in half of the samples, average ray height is high, in all other species abundant in B. bangweolensis.

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Parenchyma forming 25-50% of the it is over 1070 1J.ffi); 4-10 (3-12) per tangen· ground tissue (25% in B. bangweolensis to tial mm (4 in B. bangweolensis to 10 in B. 50% in B. spathacea), or 20-40% of the sylvatica); aggregate rays, often with stone total cross section; diffuse and diffuse-in­ cell nests, present in B. bangweolensis. aggregates in B. zenkeri; in short multiseriate Crystals always present, in chambered tangential bands with a width of 2 (1-5) cells axial parenchyma cells only (in one sample), in B. bangweolensis, B. polygalacea and B. in both axial and ray parenchyma cells (in zenkeri, in long multiseriate tangential bands three samples) or only in ray parenchyma 3-5 (1-7) cells wide in B. spathacea andB. cells (in four samples); number of compart­ sylvatica; alifonn parenchyma present in B. ments per crystalliferous parenchyma strand spathacea. Parenchyma strands without crys­ 4-16 (4 in B. zenkeri to 16 in B. spathacea). tals and not immediately bordering upon ves­ Pith flecks and t)rloses absent. sels fusifonn or composed of two cells; strand length 200-310 11m (200 11m in B. bangwe­ Dalhousiea africana (5 samples; Figs. 11, olensis to 310 lJ.ffi in B. spathacea). 12, 14) Rays in an samples of combination type IV of Baretta-Kuipers (1973); in B. polygala­ Growth ring boundaries absent or indistinct. cea and B. zenkeri uniseriate rays almost ab­ Storied structure of only axial elements sent. Width 2-6 (1-15)-seriate (2-seriate in (axial parenchyma and vessel elements; fibres B. sylvatica to 6-seriate in B. bangweolensis are usually not storied) present in all samples. and B. zenkeri); square and upright cells Vessels solitary, in radial multiples of less frequently forming uniseriate marginal parts than four vessels, and in clusters; in two size of the multiseriate rays in B. polygalacea and classes; number per square mm 30-75 (25- B. zenkeri; height 190-2100 (55 up to more 85); tangential diameter 140-160 (15-395) than 10,000) 11m (190 11m in B. sylvatica to 11m; vessel member length 240-270 (200- 2100 11m in B. zenkeri; only in B. sylvatica 320) 11m. Horizontal diameter of intervessel the average height is low, in allother species pits 10 1J.ffi, vessel-ray pits 11 11m.

Legend to figures 1-24. Secondary xylem of some Baphia-group representatives and Ormosia. In TS the position of the cambium is near the top side of the photo graph.

Figs. 1-10. Baphia species. - 1: B. marceliana, shrub, TS, x 20. Growth ring boundaries dis­ tinct, marked by terminal parenchyma; vessels nearly always solitary; within Baphia one of the two samples with the lowest number (5) of vessels per square mm; long multiseriate tangential bands ofaxial parenchyma abundant. - 2: B. polyantha, shrub, TS, x 20. Vessels very thick­ walled (14 1J.ffi), frequently in radial multiples of more than four; vessel diameter 115 (25-310) 1J.ffi, representing the widest range within the investigated Baphia species. - 3: B. laurentii, large tree, TS, x 20. - 4: B. laurentii, small tree, RLS, x 67. Multiseriate ray composed of procum­ bent cells with square and/or upright cells forming the short uniseriate marginal parts as weIl as the contact areas with vessels and axial parenchyma. - 5: B. massaiensis, small tree, TS, x 20. Vessels frequently in radial multiples of more than four. - 6: B. massaiensis, small tree, TLS, x 67. Tendency to storied structure ofaxial parenchyma and vessel elements; parenchyma strands composed of two cells; top of end cells gable-shaped. - 7: B. bequaertii, small tree, TS, x 20. Rather short tangential bands ofaxial parenchyma, somewhat 'festooned' like in Dalhousiea; axial and ray parenchyma often with dark contents. - 8: B. sylvatica, liana, TS, x 20. Within Baphia the species belongs to a group with the lowest (60 11m) average vessel diameter. - 9 & 10: B. sylvatica, liana, RLS and TLS respectively, both x 67. Storied structure ofrays and axial ele­ ments (fibres more or less storied); parenchyma fusifonn or in two celled strands; top of end cells gable-shaped; crystals in chambered axial parenchyma cells, 4 or 8 compartments per strand.

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Ground tissue of librifonn fibres together Vessels solitary, in short radial multiples, with axial parenchyma, forming about 50% in radial multiples of more than four vessels, of the total cross sectional area. rarely in clusters; number per square mm Librifonn fibres with wall thickness of 32-45 (20-55); tangential diameter 50-110 6-8 J.UIl, mean diameter of lumina 1 J.UIl; gela­ (15-220) j.lm, vessel member length 260- tinous layers not obselVed. 270 (110-360) j.lm. Horizontal diameter of Parenchyma forming 40-60 % of the intelVessel pits 11 j.lm, vessel-ray pits 6-7 ground tissue, or 20-30% of the total cross j.lm. section; in long, 3-4 (2-10) cells wide, tan­ Ground tissue of librifonn fibres together gential bands often 'festooned' (arranged in with axial parenchyma, forming about 80% tangential arcs) and in complete or incom­ of the total cross sectional area. plete sheaths around the vessels. Parenchyma Librifonn fibres with wall thickness of 4 strands without crystals and not immediately j.lm, diameter lumina 4 J.UIl, gelatinous layers bordering upon vessels fusifonn or compos­ present. ed of two cells; strand length 240-290 j.lm. Parenchyma fonning 65% of the ground Brown cell contents present. tissue, or 52% of the total cross section; in Rays in all sampies of combination type long, 3-4 (1-9) cells wide, tangential bands IV of Baretta-Kuipers (1973). Width 5-7 and in complete or incomplete sheaths around (l-21)-seriate; marginal cells higher than the vessels. Parenchyma strands without crys­ body cells in 35-50% of the rays; height tals and not immediately bordering upon ves­ 1600-2345 (85-7500) j.lm; 3 (2-4) per sels fusifonn or composed of two cells; strand tangential mm. Brown cell contents present. length 280-320 j.lm. Fibres often bending horizontally in a radial Rays in all sampies of combination type direction into wider rays, between the ray pa­ III of Baretta-Kuipers (1973). Width 2-4 renchyma cells (see Fig. 14). (1-8)-seriate; marginal cells higher than body Crystals always present in rays and cham­ cells in 60% of the rays; height 620-950 bered axial parenchyma cells with 12-20 (85-4800) J.UIl; 3-5 (2-6) per tangential mm. compartrnents per strand. Crystals always present, only in chamber­ Pith flecks and tyloses absent. ed axial parenchyma cells with c. 16 cornpart­ ments per strand. Leucomphalos capparideus (2 sampies; Pith flecks and tyloses absent. Figs.15-17) Baphiastrum brachycarpum (5 sampies; Growth ring boundaries distinct. Figs. 13, 18, 19) Storied structure of only axial elements (axial parenchyma and vessel elements; fibres Growth ring boundaries always present, usually not storied) irregularly present. inconspicuous.

Figs. 11-19. Dalhousiea (11, 12 & 14), Leucomphalos (15-17) and Baphiastrum (13, 18 & 19) species. - 11: D. ajricana, liana, TS, x 20. Growth ring boundaries absent or indistinct; next to vessels with a large diameter, many small ones present; tangential bands ofaxial parenchyma more Qr less 'festooned'. -12: D. ajricana, liana, TLS, x 67. Parenchyma in two-celled strands; top of end cells gable-shaped. - 13: B. brachycarpum, liana, TS, x 20. Within Baphiastrum the sampie with the largest average vessel diameter (185 j.lm). -14: D. ajricana, liana, RLS, x 67. Fibres bending horizontally into wider rays. - 15, 16 & 17: L. capparideus, liana, TS and TLS, x 33, x 67 & x 33 respectively. Storied structure of only axial parenchyma and vessel elements, locally present. - 18 & 19: B. brachycarpum, liana, TLS and RLS respectively, both x 67. Storied structure of only axial parenchyma and vessel elements; parenchyma strands composed of 2-4 cells; top of end cells gable-shaped.

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Storied structure of only axial elements cell nests in Brete1er 2178; ray parenchyma (axial parenchyma and vessel elements; fibres cells with brown contents in Louis 6062. are usually not storied) present in all sampies; Crystals in chambered axial parenchyma in Louis 6062 only with a very slight ten­ cells only (in one sampie), in axial and ray dency towards storying. parenchyma cells (in three sampies), or ab­ Vessels solitary and in short radial multi­ sent (in one sampie); number of compart­ ples, but radial multiples of more than four ments per crystalliferous parenchyma strand vessels and clusters always rarely present; in 8-12. two size classes; number per square mm 17- Pith flecks and tyloses absent. 53 (14-65) (17 in Breteler 2842 to 53 in Louis 6062 differs from other Baphias­ Louis 6062); tangential diameter 70-185 trum-species in many aspects. (15-270) Ilm (70 Ilm in Louis 6062 to 185 Ilm in Breteler 2842); vessel member length Airyantha schweinfurthii (7 sampies; 190-320 (110-360) Ilm; vessel walls very Figs. 20, 21, 23) thick-walled (11 1lID) in Breteler 2842. Hori­ zontal diameter of intervessel pits 10 Ilm, of Growth ring boundaries always present, vessel-ray pits 6-10 1lID. distinct (only in Breteler 1431 more or less Ground tissue of libriform fibres together indistinct). with axial parenchyma, forming about 50% Storied structure of only axial elements of the total cross sectional area. (axial parenchyma and vessel elements; fibres Libriform fibres with mean wall thickness are usually not storied) present in all samples. of 4-6 Ilm, mean diameter of lumina 3 Ilm; Vessels solitary and in radial multiples; ra­ gelatinous layers present in 80% of the sam­ dial multiples of more than four vessels and pIes. clusters also present in six samples; number Parenchyma forming 35-45 % of the per square mm 11-42 (8-55) (11 in V. & o. ground tissue (35% in Louis 6062 to 45% in 142 to 42 in Louis 5997); tangential diameter Breteler 2842), or 18-23% of the total cross 55-160 (15-250) Ilm (55 Ilm in Louis 5997 section; diffuse and diffuse-in-aggregates in to 160 Ilm in Breteler 1431), in two size clas­ one sample; in long uni- to biseriate tangen­ ses; vessel member length 260-290 11m; ves­ tial bands (terminal) in one sampie, in short sels very thick-walled (14 Ilm) in Breteler multiseriate tangential bands with an average 1765. Horizontal diameter of intervessel pits width of 2 (1-4) cells in two samples; in 10 Ilm, vessel-ray pits 6-12Ilm. long multiseriate tangential bands 2-4 (1-7) Ground tissue of libriform fibres together cells wide (4 cells wide in Louis 6062), and with axial parenchyma, forming about 70% with vasicentric parenchyma or incomplete of the total cross sectional area. sheaths around the vessels in all sampies. Libriform fibres with mean wall thickness Parenchyma strands without crystals and not of 4-6 Ilm, diameter of lumina 3 Ilm; fibres immediate1y bordering upon vessels, com­ with gelatinous layers present in 60% of the posed of 2-4 cells; strand length 210-340 sampies. llID (210 llID in Louis 6062 to 340 Ilm in Parenchyma forming 15-45 % of the Breteler 2457 and 2178). Axial parenchyma ground tissue (15% in V. & O. 142 to 45% of Breteler 2457 and 2178 sometimes not lig­ in Breteler 1431 and 1765), or 11-32% of nified. the total cross section; diffuse and diffuse-in­ Rays in all sampies of combination type aggregates in six samples; in long uni- to bi­ IV of Baretta-Kuipers (1973). Width 3-7 seriate tangential bands (terminal) in four (1-20)-seriate (3-seriate in Louis 6062 to 7- samples, in short multiseriate tangential bands seriate in Brete1er 2178 and 2842); marginal with an average width of 3 (1-4) only in cells higher than body cells in 55-65% of V. & O. 142; in long multiseriate tangential the rays; height 380-9000 (40 up to over bands 2-3 (1-6) cells wide in all samples 10,000) Ilm (380 Ilm in Louis 6062 to 9000 (more or less so in V. & O. 142); in Leeuwen­ llID in Breteler 2842); 3-8 (2-10) per tangen­ berg 4232 and V. & O. 142 vasicentric pa­ tial mm (8 in Louis 6062). Rays with stone renchyma more or less aliform. Parenchyma

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21

Figs. 20-24. Airyantha (20, 21 & 23) and Ormosia (22 & 24) species. - 20: A. schweinfurthii, liana, TS, x 20. Growth ring boundaries indistinct. - 21: A. schweinfurthii, liana, TLS, x 67. Parenchyma strands composed of 2-4 cells, top of end cells gable-shaped. - 22: O. coutinhoi, large tree, TS, x 20. Vessels nearly always solitary; aliform to confluent vasicentric parenchyma, abundantly present. - 23: A. schweinfurthii, liana, RLS, x 67. Procumbent ray cells short in radial direction; marginal cells higher than body cells. - 24: O. costulata, large tree, TLS, x 67. Storied structure absent; parenchyma strands composed of 2-4 cells, top of end cells more often conical than gable-shaped.

strands without crystals and not immediately Leeuwenberg 4232 to 1400 ~m in Breteler bordering upon vesseIs, composed of 2-4 1765); 5-9 (4-10) per tangential mm (9 in cells; strand length 280-31 0 ~m . Axial pa­ Louis 5997). Procumbent ray parenchyma renchyma with brown cell contents in an cells short in Leeuwenberg 4232. Ray paren­ samples. chyma cells with brown contents in all sam­ Rays in all samples of combination type N pIes. of Baretta Kuipers (1973), except in Louis Crystals absent in five samples, or only in 5997 of combination type I. Width 1-5 (1- chambered axial parenchyma (Breteler 1432 6)-seriate (uniseriate in Louis 5997 to 5-seri­ and 1765); number of compartments per crys­ ate in Breteler 1431 and 1765), marginal cells talliferous parenchyma strand 6-10; prismat­ higher than body cells in 30-40% of the rays; ic or rhomboidal in shape. height 650-1400 (85-3600) ~ (650 ~ in Pith flecks and tyloses absent.

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Ormosia (5 species, 5 sampIes; Figs. 22,24) Crystals always present in axial paren­ chyma; number of compartrnents per crystal­ Growth ring boundaries always present, liferous parenchyma strand 4-8 (4 in O. ca­ mostly distinct; indistinct in O. coccinea and favensis to 8 in the other samples). O. costufata. Pith flecks sometimes present in O. costu­ Storied structure absent. fata and O.Jastigiata; tyloses absent. Vessels often solitary (80% of the vessels) and in short radial multiples and clusters, in Discussion O. coccinea, O. costufata and O. Jastigiata Species in this study of Dalhousiea, Leuc­ also present in radial multiples of more than omphafos, Baphiastrum and Airyantha are four vessels; number per square mm 2 (1-4); lianas, like 5 species of Baphia. The remain­ tangential diameter 140-250 (50-360) ~m ing 15 Baphia species are shrubs or small (140 ~m in O.jastigiata to 250 ~ in O. ca­ trees. Only the sampIes of the Ormosia-spe­ lavensis); vessel member 1ength 330-450 cies are from large trees. In climbing species (170-600) ~m (330 ~ in O. coccinea to the percentage of surface area occupied by 450 ~m in O. cafavensis). Horizontal diam­ vessels, as seen in transverse section, is eter of intervesse1 pits 8 ~m, vessel-ray pits much higher than in non-climbing species 9 ~. Brown contents often present. (Ter Welle 1985). This is due either to an Ground tissue of libriform fibres together increase in number of vessels per square mm with axial parenchyma, forming about 90% and/or to an increase in vessel diameter. of the total cross sectional area. Lianas belonging to the Connaraceae, for in­ Libriform fibres with mean wall thickness stance, in general have wider vessels and of 4-10 ~ (10 ~ in O. costufata); diameter shorter vesse1 members than species of of lumina 4-18 ~m (4 ~m in O. coccinea to shrubs, treelets or trees of the same family 18 ~ in O. calavensis); ge1atinous layers not (Dickison 1972). In Baphia climbing species observed. possess more vessels per square mm than Parenchyma forming 20-40 % of the shrubs or small trees. Different vessel size ground tissue (20% in O. cafavensis to 40% classes are also more frequently represented in O. coccinea and O. costulata), or 18-36% in climbers and dwarf shrubs of dry sites of the total cross section; diffuse and diffuse­ than in shrubs and trees of mesic sites (Baas in-aggregates scarcely present in three of the 1990). This is also the case in Baphia. In sampIes; long uni- and biseriate tangential climbing species ray height and ray width are bands (terminal) and short multiseriate tan­ also larger than in non-climbing species (Ter gential bands absent; in long multiseriate tan­ Welle 1985). In Baphia lianoid species are gential bands 4-7 (2-12) cells wide (7 cells mainly provided with higher rays than spe­ wide in O. coutinhoi); aliform to confluent cies of shrubs and small trees. vasicentric parenchyma abundantly present. Senn (1943) stated that in Leguminosae Parenchyma strands without crystals and not the vessel arrangement varies considerably. immediately bordering upon vessels, com­ He also assurnes that avesseI arrangement in posed of 2-4 cells; end cells of strands ex­ clusters or long radial multiples, usually com­ hibit conical-ends in tangential section or posed of narrower vessels than the solitary conical-ends are more frequently present than ones and those in short radial multiples, is gable-ends; strand length 400-560 ~m (400 associated with a xeric habitat. Baretta-Kui­ ~m in all sampIes except O. costufata). Brown pers (1981) concluded that variation in vessel cell contents absent. diameter and number per square mm is of Rays in all sampIes of combination type N interest for identification purposes and may ofBaretta-Kuipers (1973). Width 2-3 (1-4)­ be habitat related, but is not of much taxon­ seriate; upright cells almost absent in O. cou­ omic value. Features which are considered tinhoi; marginal cells higher than body cells significant in the distinction of leguminous in 55-75% of the rays; height 300-450 genera according to Senn (1943), Normand (80-960) ~ (450 ~m in O. coutinhoi); 4-6 (1950) and Baretta-Kuipers (1973, 1981, (3-7) per tangential mm. 1982) are those ofray and axial parenchyma.

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Table 1. Variation of selected qualitative and quantitative wood characters of the investigated genera.

! ..., ::;ä ..9 § E -<:'" '" .:!t'" t::..., ~ ~ -S'" .;:;'" .-'" .0.; c c .S! <.> 0<: c §.~ ~ ::t §- ~ !::... Characters \ Genera "tl~ !I"tl~ Q --<" "tl :;;: Cl

Storied structure p + r, p p + r, p p p p p

Vessels member length Uun) 155-340 175-370 240-270 260-270 190-320 260-290 330-450 number per sq.mm 4-42 16-135 30-75 32-45 17-53 11-42 2 tangential diam. Uun) 60-140 60-160 140-160 50-180 70-185 55-160 140-250

Rays height in J.IIIl 180-790 1070-2100 1600-2245 620'-685 380-9000 650-1400 300-450 seriate 1-6 2-6 5-7 2-4 3-7 1-5 2-3 combination type IV, (Ill) IV IV 111 IV IV IV

Axial parenchyma no.ofcellsperstrand 1-2 1-2 1-2 1-2 2-4 2-4 2-4 strands with pred. gable-ends + + + + + + width of long tangen- tial bands (cells) 2-7 3-5 3-4 3-4 2-4 2-3 4-7 aliform and/ or confluent (+) (+) (+) arr. in tangential arcs (+) +

Crystals cp,cp+r,r cp,cp+r,r cp+r cp cp, cp+r cp cp*

Symbols and abbreviations: Quantitative data are given as a range of mean values for different species and/or sampIes of the genus, or as a single mean value. + = and; + = present; - = absent; (+) = character poorly developed; * = often only 1 or 2 cells of a strand chambered and with crystals; cp = chambered axial parenchyma cells; diam. = diameter; no. = number; p = axial parenchyma; pred. = predominant (more than 80% of the end cells of the strands); r = rays or ray parenchyma cells; sq. = square.

From the descriptions and Table 1 it is dear structure features (storied structure and units that there are only minor differences between of the storied structure complex), the shape the genera of the Baphia-group concerning of the marginal ray cells and ray structure. these features. In their macromorphology Two units of the storied structure complex several genera in this group are also almost are especially important, viz. the shape of the identical (polhill 1981; Breteler, personal end cells of the parenchyma strands in tan­ communication). gential section and the low number of cells in Using wood anatomy, the three legumin­ the parenchyma strands. U sing the above ous subfamilies are separable according to mentioned criteria, all sampies of Dallwusiea, Reinders-Gouwentak: and Rijsdijk (1955) but Leucomphalos, Baphiastrum and Airyantha the Swartzieae show intergrading features of constitute one group which has to be placed the Papilionoideae and the Caesalpinoideae. in the Papilionoideae. Most Baphia sampies The characteristics in question are storied- were also easily grouped into the Papilion-

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oideae, but for R. gracilipes, B. laurifolia Tbe genera of the Raphia-group can be ar­ (Louis 3619) and R. obovata this is not so ranged in two groups; Airyantha and Raphi­ obvious. Ormosia sampies, however, can be astrUfn with 2-4 cells per axial parenchyma placed in the Papilionoideae but also in the strand, Leucomphalos, Dalhousiea and Ra­ Caesalpinioideae. Tbis does not imply that phia with fusiform parenchyma or 2-celled these species belong to two subfamilies. It strands. In other aspects the genera of the merely indicates that Papilionoideae and Raphia-group resemble each other very Caesalpinioideae not only merge into each much. A dear distinction of the genera based other via the Swartzieae but also via the on wood anatomy, is not very well possible. Sophoreae and that delimitation of the subc Airyanrha andRaphiasrrum especially are al­ families with the help of wood anatomical most identical. This can be used to support criteria is probably illusionary. It is also pos­ a merger of the two genera. Tbere are only sible that features used by Reinders-Gouwen­ minor differences between the genera Leuc­ tak and Rijsdijk (1955) are not always appro­ omphalos, Dalhousiea and Raphia. Tbe priate for African species, since they are wood anatomical diversity of Raphia is rela­ mainly based on material from Java (Indone­ tively large, almost entirely overlapping the sia; Moll & Janssonius 1918) and on material limited structural ranges of the other genera. from Neotropical species. However, storied structure of all elements is Also Baretta-Kuipers (1981) concluded regularly present only in Raphia, and tangen­ that the Sophoreae are intermediate in all tial parenchyma bands are wider. Dalhousiea selected wood anatomical features (percent­ is usually distinguished from Leucomphalos age of heterocellular rays, rays more than and Raphia by the presence ofaxial paren­ 500 J.Illl high, of cells per parenchyma strand, chyma arranged in tangential arcs ('festoon­ storied structure and septate fibres) between ed'); Leucomphalos is the only genus in the two subfamilies Caesalpinioideae and which most of the rays are multiseriate, but a Papilionoideae. considerable number are partly or entirely uni­ From Table 1 it is dear that Ormosia seriate (combination type ill). stands apart from the other genera; storied Acknowledgements structure is absent, end cells of parenchyma We wish to express our appreciation to the strands exhibit conical-ends or conical-ends curators of the following wood collections are more frequently present than gable-ends, for providing valuable research materials: and Ormosia has a large percentage of high Royal Botanic Gardens, Kew; Bundesfor­ marginal ray cells. All this implies that the schungsanstalt für Forst- und Holzwirtschaft, genus is intermediate between the two sub­ Hamburg; Musee Royal de I'Afrique Centrale, families Papilionoideae and Caesalpinioideae Tervuren; Systematische Plantkunde, Utrecht; according to characteristics given by Reinders­ Plantentaxonomie, Wageningen. Gouwentak and Rijsdijk (1955). Also the percentage of cross sectional area occupied References by vessels (somewhat wider but in a small Baas, P. 1990. Ecological trends in the wood number per square mm) is much smaller than anatomy and their biological significance. in the other genera; vessels are also more of­ In: F.H. Schweingruber (ed.), Anatomy of ten solitary (see also Reinders-Gouwentak & European woods: 739-765. Haupt, Bem. Rijsdijk 1968). Tbis, however, may be caus­ Baretta-Kuipers, T. 1973. Some aspects of ed by the fact that Ormosia is represented by wood anatomical research in the genus large trees, the other genera by lianas or in Inga (Mimosaceae) from the Guianas and the case of Raphia also by shrubs and some­ especially Surinam. Acta Bot. Neer!. 22: times small trees. Vessel members in Ormosia 193-205. are longer than in the other genera; rays are Baretta-Kuipers, T. 1981. Wood anatomy of narrow and low; axial parenchyma is often Leguminosae: its relevance to taxonomy. aliform and/or confluent; crystals usually are In: R.M. Polhill & P.H. Raven (eds.), only present in one or two chambered cells of Advances in legume systematics, part 2: an axial parenchyma strand. 677-705. Royal Botanic Gardens, Kew.

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Baretta-Kuipers, T. 1982. Wood structure of Normand, D. 1950. Atlas des bois de la Cöte the genus Erythrina. Allertonia 3: 53- d'Ivoire. I. Les Legumineuses: 98-142. 69. Centre Techn. Forestier Tropical, Nogent­ Chant, S.R. 1979. Leguminosae. In: V.H. sur-Mame. Heywood (ed.), Flowering of the Polhill, R. M. 1981. Papilionoideae. In: R. M. world: 149-152. Oxford Univ. Press, Polhill & P.H. Raven (eds.), Advances in Oxford. legume systematics, part 1: 191-230. Dickison, W.C. 1972. Anatomical studies in Royal Botanie Gardens, Kew. the Connaraceae. 2: Wood anatomy. J. Reinders, E. 1935. Fibre-tracheids, libriform Elisha Mitchell Scientific Soc. 88: 120- wood fibres and systematics in wood anat­ 136. omy. Trop. Woods 44: 30-36. Heywood, V. H. 1971. The Leguminosae - Reinders-Gouwentak, C.A. 1955. The stor­ a systematic purview. In: lB. Harborne, ied-structure-features and the taxonomie D. Boulter & B.L. Turner (eds.), Chemo­ rank of the leguminous taxa. Acta Bot. taxonomy of the Leguminosae: 1-29. Neerl. 4: 460-470. Academic Press, London, New York. Reinders-Gouwentak, C.A. & lF. Rijsdijk Hutehinson, J. 1964. The genera of flower­ 1955. Wood anatomical characterization ing plants. Vol. 1. Clarendon Press, Ox­ of the leguminous taxa. Proc. Kon. Ned. ford. Akad. Wet. Sero C. 58: 41-50. IAWA Committee 1989. IAWA listofmicro­ Reinders-Gouwentak, C.A. & J.F. Rijsdijk scopic features for hardwood identifi­ 1968. Hout van Leguminosae uit Suri­ cation (eds. E.A. Wheeler, P. Baas & name. Veenman & Zn, Wageningen. P.E. Gasson). IAWA Bull. n.s. 10: 219- Senn, H.A. 1943. The relation of anatomy 332. and cytology to the classification of the Janssonius, H.H. 1940. Anatomische Be­ Leguminosae. Chron. Bot. VII: 306-308. stimmungstabelle für Javanischen Hölzer. Stern, W.L. 1988. Index xylariorum. Insti­ Brill, Leiden. tutional wood collections of the world. 3. Moll, J. W. & H.H. Janssonius 1918. Mikro­ IAWA Bull. n.s. 9: 203-252. graphie des Holzes der auf Java vorkom­ Welle, B.J.H. ter. 1985. Differences in menden Baumarten 3: 11-207. Brill, Lei­ wood anatomy of lianas and trees. IAWA den. Bull. n. s. 6: 70.

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