IAWA Bulletin n.s., Vol. 7 (3),1986 175

THE SYSTEMATIC WOOD ANATOMY OF THE (URTICALES) V. GENERA OF THE TRIBE WITHOUT URTICACEOUS *

by

B.1. H. ter Welle, 1. Koek-Noorman and S. M. C. Topper Institute of Systematic Botany, University of Utrecht, Heidelberglaan 2, 3508 TC Utrecht, The Netherlands

Summary The wood anatomy of the Moreae without based upon these characters, however (Berg, urticaceous stamens is described in detail. Ge­ 1983), is not in accordance with the tribes neric descriptions of the following genera are Moreae and sensu Corner (1962). provided: , , , Ba­ Both Berg's and Corner's subdivisions deviate tocarpus, , , , from older classifications, as given by, for in­ Prainea, , Sparattosyce, and . stance, Bentham and Hooker (1880) and Engler Wood anatomical variation below the (1888). level is very limited, except in the genus Clari­ The Moreae characterised by the absence of sia. Intergeneric variation, however, is much urticaceous stamens comprise the genera Antia­ more evident. Most genera can be recognised ropsis (), Artocarpus (Southeast by the presence or absence of septate fibres, ), Bagassa (Neotropics), (Neo­ and of radial latex tubes, the size of the inter­ tropics), Clarisia (Neotropics), Hullettia (South­ vascular pits, the parenchyma distribution, and east Asia),Parartocarpus (), Poul­ crystal distribution. The diagnostic and taxon­ senia (Neotropics), Prainea (Malesia), Sorocea omic value of several characters is discussed. (Neotropics), Sparattosyce (New Caledonia), Key words: Moraceae, Moreae, systematic wood and Treculia (Tropical ). anatomy. Methods and Materials In troduction The methods employed are those given in This paper is part of a series, in which the the first paper of this series (Koek-Noorman et wood anatomy of the Moraceae is described aI., 1984). All wood samples studied are backed and discussed in relation to the of by herbarium vouchers. Taxa of the African the family. The general outline of this study is and Neotropical genera were identified or cited provided in the first paper (Koek-Noorman et by the authors of the various monographs. De­ aI., 1984). In the fourth paper of the series the tails on individual wood samples are provided wood anatomy of those genera of the tribe at the beginning of each generic description. Moreae (sensu Berg) which are characterised by The genus Hullettia is not represented in this urticaceous stamens was described (Ter Welle study because wood samples were not at our et aI., 1986). In this paper the genera without disposal. this type of stamens are dealt with. In addition The data on the specific gravity are based on to the structure of the stamens, the following the samples cited and complemented with data features can be used to characterise this group on other samples of the Utrecht wood collection of genera in Moreae according to Berg (1983): not further included in the anatomical study. rather simple to complex; macro­ Quantitative data generally refer to average spermous; diaspores dry or mostly in fleshy values, unless specified otherwise. structures, mostly zoochorous; mostly in rain­ forest habitats; taxonomically and geographi­ cally segregated, pantropical, but chiefly in the Generic descriptions Indo-Malesian region. The tribe Moreae sensu Berg (1983) com­ Antiaropsis Schumann (Figs. I, 2) prises all genera of the tribes Artocarpeae and A genus with one or two species (Corner, Moreae sensu Corner (1962), and also a few 1962). The genus is probably restricted to Irian genera assigned to the tribe Olmedieae by Jaya (). Its occurrence in Papua New Corner (1. c.). The subdivision in two groups Guinea is questioned by Corner (1962). The

* This project was made possiple by a grant of BION -ZWO (14.45 -0 1).

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shrubs or small trees, up to II m high, are found 22031 b (Uw 24421). - A. fretissii Teijsm. & in lowland forests up to 1000 m altitude, usual­ Binn. ex Hassk. Indonesia, Irian Jaya: BW 9871 ly in the undergrowth. (Uw 18112), BW 2133 (Uw 24316), BW 2512 Mat e ri a 1st u die d : A. decipiens Schu­ (Uw 24317), BW 2175 (Uw 24318). -A. hor­ mann. Indonesia, Irian Jaya: CSIRO H 5198 ridus Jarrett. Indonesia, Irian Jaya: BW 11593 (Uw 25533). (Uw 20479). - A. integer (Thunb.) Merr. Indo­ Ge n e r al fe at u re s: Growth rings absent; nesia, Irian Jaya: BW 5659 (Uw 24319); Jamai­ heartwood not available, sapwood light brown. ca (cult.): US Nat. Herb. (Uw 8294); South Texture fine, grain straight. Specific gravity c. Africa (cult.): Baijnath s.n. (Uw 25507). - A. 750 N per cubic metre. kemando Miq. Malaya: FMS 28853, ex FHOw Micro s cop i c fe a t u re s: Vessels diffuse, 4536 (Uw 24608); Indonesia: RTIw IND. Col. solitary (50%) and in short radial multiples of 3590 (Uw 24605), Pfeiffer E 755 (Uw 24324). 2-3; 9 per sq.mm, round to slightly oval, diam­ - A. lakoocha Roxb. Pakistan: ex MADw 24487 eter 108 J.l1l1, vessel member length 525 J.l1l1. (Uw 18033); Thailand: Royal For. Dept. s.n. Perforations simple, end walls almost transverse. (Uw 24656). - A. lanceifolius Roxb. Indonesia: Intervascular pits alternate, polygonal, occasion­ RTIw IND. Col. 2200 (Uw 24606); RTlw IND. ally round, 7 -9 J.l1l1. Vessel-ray and vessel-paren­ Col. 4027 (Uw 24606). - A. nitidus Trecul. chyma pits larger, elongated, half-bordered, the Malaysia, Sabah: Sandakan For. Dept. 25510 borders often reduced. Thin-walled tyloses com­ (Uw 24217). - A. sepicanus Diels. Indonesia, mon, often with vitreous silica. Fibres septate, Irian Jaya: Fokkinga 7822 (Uw 24329). - A. with several septa per fibre, with small simple sericicarpus Jarrett. Malaysia, Sabah: Sandakan pits restricted to the radial walls; walls 3-4 /-Lm, For. Dept. 50586 (Uw 24218). - A. teysmannii lumina 3-7 J.l1l1; length 1600 /-Lm; F/V-ratio 3.0. Miq. Indonesia, Irian Jaya: BW 1204 (Uw Rays uniseriate and multiseriate, heterogeneous 24330). type II/III, 7 per mm. Uniseriate rays 25 %, Ge neral fe a tures: Growth rings absent; composed of upright cells only, but variable in heartwood light brown to dark brown, often height, height up to 12-24 cells (480-800 /-Lm). sharply distinct from the pale yellow to light Multiseriate rays composed of procumbent cells brown sapwood. Texture coarse, grain (weakly) except for the uniseriate margins of 1-5, occa­ interlocked. Specific gravity 280-940 N per sionally up to 20 rows of square and/or upright cubic metre. cells; 3-4 cells wide, up to 960-1400 /-Lm high. M i c r 0 s cop i c f eat u res: Vessels diffuse, Occasionally with some sheath cells. Parenchy­ solitary (45-80%) and in short radial multiples ma scanty paratracheal, and in concentric and irregular clusters of 2-4; 2-6 per sq. mm, bands, 3-6 cells wide and 4-5 per mm. Paren­ round and oval, diameter 160-310 jlm, vessel chyma strands of 4 cells. A few latex tubes ob­ member length 350-630 /-Lm. Perforations sim­ served in the rays. ple, end walls almost transverse. Intervascular pits alternate, round, polygonal and occasion­ Artocarpus J. R. & G. Forster (Figs. 3-6) ally slightly oval, 9-13 /-Lm. Vessel-ray and ves­ The genus has been monographed by Jarrett sel-parenchyma pits larger, half-bordered, the (l959c,d,e; 1960a, b). About 50 species are borders sometimes reduced. Thin-walled tyloses currently recognised. The genus is distributed common, but occasionally absent. Fibres non­ from India to southern China, Malesia and the septate, with small simple pits restricted to the Solomon Islands. Two species, A. altilis and A. radial walls; walls 2-4 /-Lm, lumina 7-25 /-Lm, heterophyllus, are cultivated all over the tropics gelatinous fibres present or absent; length for their fruits. Trees to large trees, up to 45 m, 1170-1950 /-Lm; F/V-ratio 2.3-3.9. Rays uni­ in evergreen forests and areas with a mild mon­ seriate and multiseriate, heterogeneous type soon climate, usually from sea level up to 1000 II/III, 3-6 per mm. Uniseriate rays 3-30%, m altitude. composed of upright, square and sometimes Mat e ria 1st u die d: A. anisophyllus Miq. few procumbent cells, height up to 4-10 cells Malaysia, Sabah: Sandakan For. Dept. 50567 (160-480 /-Lm). Multiseriate rays composed of (Uw 24213). - A. blancoi (Elmer) Merr. Philip­ procumbent cells, except for the uniseriate pines, Quezon: 1.P. Rojo 262 (Uw 24214). - margins of 1-2 (5) rows of square and/or up­ A. communis 1. R. & G. Forster. Indonesia, Irian right cells; 3-5 cells wide, up to 600-1200 /-Lm Jaya: BW 2904 (Uw 24602), BW 4219 (Uw high. Occasionally few sheath cells present. Pa­ 24603). - A. dadah Miq. Indonesia: RTIw IND. renchyma aliform with short to long wings, oc­ Col. T889 (Uw 25521), RTIw IND. Col. 14317 casionally confluent, and sporadically in short (Uw 25522). - A. elasticus Reinw. ex Blume. wavy bands. Parenchyma strands of 3-4 cells. Malaysia, Sabah: Sandakan Herb. Inst. 18656 Vitreous silica common, in the fibres or in the (Uw 24215); Indonesia, Java: Koorders 1026c/ vessels, often enveloping the tyloses. Radial

Downloaded from Brill.com10/06/2021 06:03:46AM via free access IAWA Bulletin n.s., Vol. 7 (3), 1986 177 latex tubes present in most samples, axial latex Batocarpus Karsten (Figs. 9 & 10) tubes present or absent (cf. Topper & Koek­ A genus with four species according to the Noonnan, 1980). monograph of Mello Filho & Emmerich (1968). The genus is distributed from Costa Rica to Bagassa Aublet (Figs. 7, 8) , Peru and Bolivia. Trees up to 20-30 m In a treatment for the Flora of , high, in the tropical lowland rainforests. Berg (1975) distinguished two species. Recent Material studied: B. amazonicus (Ducke) studies by the same author revealed that the Fosberg. Brazil, Amazonas: Prance & Maas 15741 genus is mono typic, as B. tiliifolia is now con­ (Uw 19158); Krukoff 6422 (Uw 7714); Peru: sidered as a juvenile fonn of B. guianensis (Berg, Williams 5334 (Uw 18416). - B. orinocensis pers. comm.). The genus is distributed in the Karsten. Ecuador: Berg & Akkermans 1079 (Uw Guianas and adjacent areas of northern Brazil. 27049). The large trees, up to 45 m high, are of rather G e n era I f eat u res: Growth rings absent; infrequent occurrence in the lowland rainforests all samples light greyish or yellowish brown, and savannas, up to 650 m altitude. Williams (1936) described the heartwood of Mat e ria 1st u die d: B. guianensis Aubl. the sample collected by himself (no. 5334) as : A.C. Smith 3542 (Uw 21698); Suri­ 'thin, pale or dark brown'. Texture medium to nam: Lanjouw & Lindeman 2433 (Uw 1742); coarse, grain straight to irregular. Specific gravi­ Brazil: INPAX-6288 (Uw23781), Capucho 356, ty 540-650 N per cubic metre. ex MADw 34591 (Uw 24261). - 'B. tiliifolia M i c r 0 s cop i c f eat u res: Vessels diffuse, Benoist.'. Guyana: F.D. 5827, ex MADw 3198 solitary (40-65%) and in short radial multiples (Uw 24263); Surinam: Stahel 194 (Uw 194); and irregular clusters of 2-4; 2-5 per sq. mm, : BAFOG 1302 (Uw 5790). round to oval, diameter 125-205 jJ.m, vessel G e n era I f eat u res: Growth rings poorly member length 405-440 jJ.m (295 fJ.ffi in B. defined; heartwood dark brown, sharply dis­ amazonicus from Peru). Perforations simple, tinct from the light brown sapwood. Texture end walls almost transverse. Intervascular pits medium to coarse, grain interlocked. Specific alternate, round or polygonal, 7 -10 jJ.m. Vessel­ gravity 880-I 000 N per cubic metre. ray and vessel-parenchyma pits larger, elongated M i c r 0 s cop i c f eat u res: Vessels diffuse, and irregular, half-bordered, the borders in part solitary (20-60%) and in short radial mUltiples reduced. Thin-walled tyloses occasionally pres­ and irregular clusters of 2-4; 3-4 (7) per sq. ent in some samples, always few. Fibres non­ mm, round and oval, diameter 145-240 p.m, septate, with small simple pits restricted to the vessel member length 325-445 jJ.m. Perfora­ radial walls; walls 2-5 jJ.m, lumina 7-20 jJ.m, tions simple, end walls almost transverse. Inter­ partly gelatinous (up to 90% of the fibres gela­ vascular pits alternate, round, polygonal or tinous in B. orinocensis); length 1165-1700 slightly oval, 9-13(-15) fJ.ffi. Vessel-ray and jJ.m; F/V-ratio 3.9-4.4 in the samples of B. vessel-parenchyma pits larger, elongated, half­ amazonicus, and 2.6 in the sample of B. ofino­ bordered, borders sometimes reduced. Thin­ censis. Rays uniseriate and multiseriate, hetero­ walled tyloses common. Fibres nonseptate, geneous type III, 4-6 per mm. Uniseriate rays with small simple pits restricted to the radial 6-18%, composed mainly of upright cells and walls; walls 3-5 jJ.m, lumina 5-13 jJ.m, gelatin­ a few rows of procumbent cells, the up'right ous fibres in variable amounts; length 945-1150 cells variable in height, height up to 3-6(-8) jJ.m; F/V-ratio 2.7-3.0. Rays uniseriate and cells (145-480 jJ.m). Multiseriate rays com­ multiseriate, heterogeneous type III, sometimes posed of procumbent cells except for the uni­ almost homogeneous, 3 -7 per mm. Uniseriate seriate margins of 1-2 rows of square and/or rays 0-14%, composed of upright, square and upright cells; 4-6 cells wide, up to 1200- often procumbent cells, height up to 4-9 cells 1400 jJ.m high in B. amazonicus (Prance & Maas (120-360 fJ.ffi). Multiseriate rays composed of 15741, and Krukoff 6422) and up to 700~800 procumbent cells except for the uniseriate mar­ jJ.m in the other two samples. Occasionally gins of 1-2 rows of square and occasionally some sheath cells. Parenchyma scanty paratra­ upright cells; 2-5 cells wide, up to 600-840 cheal to vasicentric-alifonn with short wings, fJ.ffi high. Sheath cells very scarce. Parenchyma wings sometimes long and confluent. The dom­ scanty paratracheal to vasicentric, alifonn with inating pattern and the vast majority of the short wings and some diffuse parenchyma. Pa­ parenchyma is fonned by irregular wavy bands, renchyma strands of 3-4 cells. Rhombic crys­ 3-10 cells wide and 2-3 bands per mm. Paren­ tals common in the margins of the rays, less chyma strands of 3-4 cells. Vitreous silica frequent in the axial parenchyma. Radial latex scarcely present in some samples, in vessels tubes present in all samples, axial latex tubes and/or fibres. occasionally present. (text continued on page 184)

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Figs. 1~26. Scale-line = 225 Mm, unless indicated otherwise.

Fig. 1. Antiaropsis decipiens, Uw 25533. - Fig. 2. Ibid., septate fibres. - Fig. 3. Artocarpus seri­ cicarpus, Uw 24218. - Fig. 4. Artocarpus lakoocha, Uw 18033.

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Fig. 5. Artacarpus sericicarpus, Uw 24218, radia11atex tubes, group 1. - Fig. 6. Ibid., radia11atex tube. - Fig. 7. Bagassa guianensis, Uw 1742. - Fig. 8. Bagassa tiliae/alia, Uw 5790.

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Fig. 9. Batocarpus amazonicus, Uw 18416. - Fig. 10. Ibid., Uw 19158. - Fig. 11. Clarisia bij7ora, Uw 25091. - Fig. 12. Oarisia racemosa, Uw 14464.

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Fig. 13. Clarisia ilicifolia, Uw 7982. - Fig. 14. Parartocarpus bracteatus, Uw 24435. - Fig. 15. Parartocarpus venosus ssp.forbesii, Uw 25807. - Fig. 16. Ibid.

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Fig. 17. Parartocarpus bracteatus, Uw 24436, radial latex tubes, type 2. - Fig. 18. Parartocarpus venosus ssp. forbesii, Uw 25807, homogeneous rays. - Fig. 19. Poulsenia armata, Uw 24638. - Fig. 20. Ibid., Uw 20696, vitreous silica in the fibres.

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Fig. 21. Prainea papuana, Uw 18124. - Fig. 22. Sorocea guilliminiana, Uw 20941. - Fig. 23. Sparattosyce dioica, Uw 24627. - Fig. 24. Ibid.

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Fig. 25. , Uw 24640. - Fig. 26. Ibid., radial latex tubes, type 2.

Note: In Williams 5334 (cited by Mello Filho Maguire et aI. 56675 (Uw 16474). - C. race­ & Emmerich, 1968, as B. amazonicus) rhombic mosa Ruis & Pavon. Venezuela: Breteler 5054 crystals occur in square and upright cells, and (Uw 12293); Colombia: de Bruijn 1056 (Uw less frequently in the procumbent ray cells and 14464), 1553 (Uw 14496); Ecuador: For. Stat. the (occasionally chambered) axial parenchyma Connocotto s. n. (Uw 23877); Brazil, Amazonas: cells. In all other specimens crystals are lacking. Krukoff 6327 (Uw 7645), 6628 (Uw 7829); Loureiro et aI., INPA X-5567 (Uw 23714). Clarisia Ruiz & Pavon (Figs. 11-13; Table 1) G e n era I f eat u res: Growth rings absent The genus comprises three species, according or very faint; heartwood absent or indistin­ to Berg (1968) who included the genus Acan­ guishable from the light brown sapwood in C thinophyllum monographed by Burger (1962) biflora and C ilicifolia, heartwood dark brown by combining A. ilicifolia and A. spruceana to and sharply defined from the light brown sap­ Clarisia ilicifolia. The genus is distributed in wood in C racemosa. Texture medium (C ilici­ Central America and the northern part of South folia) to coarse (C biflora, C racemosa), grain America. Two species (C racemosa and C bi­ straight, but interlocked in C racemosa. Specif­ flora) are trees up to 40 m high. Clarisia ilicifolia ic gravity 400-500 N per cubic metre in C bi­ is a shrub or tree up to 20 m high. All species flora, and 650-850 N per cubic metre in the occur in tropical lowland forests. other two species. Mat e ria 1st u die d: C biflora Ruiz & Pa­ Microscopic fea tures: Vessels diffuse, von. Colombia, Choco: Cuatrecasas 15487 (Uw solitary (25-51 %) and in short radial multiples 25091); Peru: Ellenberg 2511 (Uw 8712). - and irregular clusters of2-5; 3-13 per sq.mm, C ilicifolia (Sprengel) Lanj. & Rossb. Guyana: round to slightly oval, diameter 75-200 /.Lm, For. Dept. 4039 (Uw 982); Surinam: Lindeman vessel member length 310-485 /.Lm. Perfora­ 6776 (Uw 4587); Daniels & Jonker 1079 (Uw tions simple, end walls almost transverse. Inter­ 8593), 1081 (Uw 8595); van Donselaar 1221 vascular pits alternate, round and polygonal, (Uw 10835); Brazil: Krukoff6849 (Uw 7982); 6-11 /.Lm. Vessel-ray and vessel-parenchyma

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Table I. Some wood anatomical characters of species of Clarisia.

E E s :t ct § E '"...... c= :t Q) .... '" OIl -8 >- ~ E § :t >- ....'" ~ '" .... (j .... :§ .B ~ .g (j ;a'" - 0 '"Q) Q) '"Q) .D ..c= Species > > Ul * > ;.;::: ~ .... C. biflora 180-200 3-4 11-16 465-485 1200-1500 2.6-3.1

C. ilicifolia 75-105 7-13 + 13-22 310-320 960-1315 3.8-4.2 (u, p, a)

C. racemosa 144-200 4-5 + 1-3 400 1285-1300 3.2-3.3 (p, a)

Legend: F/V-ratio: ratio of fibre and vessel member length. - Crystal distribution: u: in square and upright cells; p: in procubent ray cells; a: in axial parenchyma cells.

pits larger, elongated, and irregularly shaped, Parartocarpus Baillon (Figs. 14-18) half-bordered, the borders sometimes reduced. The genus has been monographed by Jarrett Thin-walled tyloses common. Fibres nonsep­ (l960c). Two species and four subspecies were tate, with small simple pits restricted to the ra­ recognised by her. A third species was described dial walls; walls 2-4 j.Lm, lumina 6-20 j.Lm, by Corner (1976). The genus is distributed gelatinous fibres often present; length 960- from Thailand throughout Malesia and the 1500 .urn; F/V-ratio 2.6-4.2. Rays uniseriate Solomon Islands. The medium to large trees, and multiseriate, heterogeneous type II, III and up to 45 m high, inhabit evergreen lowland occasionally almost homogeneous, 4-7 per rainforests, extending up to 1800 m altitude. mm. Uniseriate rays 1-22%, composed ofvari­ Material studied: P. bracteatus (King) able amounts of procumbent, square and up­ Becc. Malaysia: RTIw IND. Col. 24620 (Uw right cells, height up to 4-12 cells (\ 50-360 24435), RTIw IND. Col. 24632 (Uw 24436). - j.Lm). Multiseriate rays composed of procum­ P. venenosus (loll. & Mor.) Becc. ssp. borneen­ bent cells except for the uniseriate margins of sis (Becc.) Jarrett. Indonesia, Borneo: DFP 5008 0-2(-5) cells of square and/or upright cells; (Uw 24413). - P. venenosus (loll. & Mor.) Becc. 3-6 cells wide, up to 600-1560 j.Lm high. ssp. forbesii (King ) Jarrett. Malaysia: F. R.l. Sheath cells always present, but scarce. Paren­ Kepong, ex SJRw 38498 (Uw 24639); Indone­ chyma vasicentric, aliform with short, and oc­ sia, Sumatra: Krukoff 4124a (Uw 25807). - P. casionally long wings, but predominantly band­ venenosus (loll. & Mor.) Becc. ssp. papuanus ed, varying from commonly wavy bands to (Becc.) Jarrett. Indonesia, Irian Jaya: BW 12294 sometimes concentric bands, 2-6(-8) cells (Uw 18185), BW 9422 (Uw 20478), BW 4243 wide and 1-3(-4) per mm. Terminal bands (Uw 24352); Philippines, Bataan: BF Manila, occasionally present. Parenchyma strands of ex SJRw 17584 (Uw 24340). - P. venenosus 2-4 cells. Rhombic crystals present or absent (loll. & Mor.) Becc. ssp. venenosus. Indonesia, in the rays and/or axial parenchyma, usually Java: Koorders 4273w (Uw 24633). scarce if present. Vitreous silica common in the G e n era I f eat u res: Growth rings absent; vessels, but generally scarce. Radial latex tubes heartwood light yellow-brown, indistinguishable observed in some samples of C. ilicifolia. from the sapwood. Texture coarse, grain inter­ Not e: The wood anatomical difference be­ locked. Specific gravity (290-)400-600 N per tween the three species of Clarisia are consider­ cubic metre. able. For some characters this is illustrated in M i c r 0 s cop i c f eat u res: Vessels diffuse, Table I. solitary (21-60%) and in short radial multiples

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or regular clusters of2-4 (5); 1-3 persq.mm, stricted to the radial walls; walls 2-3 J.Lill, lumi­ round to oval, diameter 170-275 J.Lm, vessel na 15-35 J.Lill;length 141O-1415!lm;F/V-ratio member length 465-740 !lm. Perforations sim­ 2.0-2.1. Rays uniseriate and multiseriate, het­ ple, end walls almost transverse. Intervascular erogeneous type III, 3-5 per mm. Uniseriate pits alternate, polygonal and occasionally rays 3-27%, composed of upright and square round, 14-15 !lm. Vessel-ray and vessel-paren­ cells and a few procumbent cells, height up to chyma pits larger, elongated and more irregu­ 4-7 cells (180-310 !lm). Multiseriate rays larly shaped, half-bordered, the borders usually composed of procumbent cells except for the reduced. Thin-walled tyloses very scarce. Fibres uniseriate margins of 1-2 rows of square and/or nonseptate, with small simple pits restricted to upright cells; 4-6 cells wide, up to 960--1400 the radial walls; walls 2-4 !lm, lumina 15-25 !lm high. Occasionally with some sheath cells. J.Lill;length 1370-1810 J.Lill; F/V-ratio 2.2-3.8. Parenchyma vasicentric-aliform with short Rays uniseriate and multiseriate, homogeneous, wings, occasionally confluent. Parenchyma 4-6 per mm. Uniseriate rays usually 1-8 %, in strands of 2-8 cells. The parenchyma distribu­ some samples up to 30%, composed of pro­ tion is not very clear in the transverse sections, cumbent cells only, height up to 4-14 cells because the lumen diameter/wall thickness ratio (140-420 J.Lill). Multiseriate rays composed of of the fibres is almost equal to that of the pa­ procumbent cells only; 2-3 (4) cells wide, up renchyma cells. Vitreous silica abundant in the to 600-1200 J.Lill high. Parenchyma rather fibres, and occasionally also in the axial paren­ abundant, vasicentric-aliform with short and/or chyma and the vessels. Radial latex tubes com­ long wings to confluent, occasionally in short mon, the diameter of the tubes in part equal to wavy bands. In all samples a tendency towards the diameter of the ray cells, partly larger (as unilateral abaxial parenchyma is evident. Paren­ seen in the tangential section). chyma strands of 3-4 cells. Radial latex tubes Not e s: Going from Mexico, Costa Rica, Pa­ common and with a large diameter as compared nama, to Colombia, the diameter of the inter­ to the surrounding ray cells; as a result, the vascular pits increases from 6-7 to 9 !lm. At latex tubes are very conspicuous. Rhombic the same time, the specific gravity decreases crystals common, but sometimes scarce, in the from 450 to 250 N per cubic metre. axial parenchyma. In about 50% of the samples a tendency to­ wards an obliq ue vessel distribution was noticed. The genus shows a high accumulation of vitreous Paulsenia Eggers (Figs. 19, 20) silica, which may exceed 7% (of dry weight) A mono typic genus distributed from Mexico according to Van Siooten (1968). to Ecuador and Bolivia. Trees up to 40 m high in tropical rainforests within the area. Prainea King ex Hook. f. (Fig. 21) Material studied: P. armata (Miq.) Stand­ The genus comprises four species according ley. Mexico: Williams 8426, ex SlRw 34588 to the most recent monograph (Jarrett, 1959b). (Uw 24624); Trigos 309 (Uw 25689); Costa Rica: The species are confined to Malesia: 3 species IlCA CCO 29 (Uw 20696); Panama: Cooper & in Sumatra, Malaya and Borneo, and I in the Slater 132, ex SlRw 10312 (Uw 24628), Cooper Moluccas and Irian laya / Papua New Guinea. & Slater 88, ex SlRw 10269 (Uw 24629); Co­ The trees up to 35 m high are found in ever­ lombia: Cuatrecasas 14342 (Uw 24638). green rainforests up to 1200 m altitude. General features: Growth rings faint or Mat e ria 1st u die d: P. frutescens Becc., absent; heartwood yellow to creamish and in­ Malaysia, Sarawak: CSIRO 32639 (Uw 24363). distinguishable from the sapwood. Texture me­ - P. !impata (Miq.) Beumee ex Heyne. Malesia dium to coarse, grain interlocked. Specific grav­ (?): RTIw IND. Col. 20013 (Uw 24437), RTIw ity 250-450 N per cubic metre. IND. Col. 14316 (Uw 24438). - P. papuana M i c r 0 s cop i c f eat u res: Vessels diffuse, Becc. Indonesia, Irian laya: BW 11785 (Uw solitary (35-90%) and in short radial multiples 18124), BW 1559 Fokkinga (Uw 24356), BW or irregular clusters of 2-4; 4-10 per sq.mm, 1664 Fokkinga (Uw 24357). round to slightly oval, diameter 105-210 !lm, G e n era I f eat u res: Growth rings absent; vessel member length 670-695 !lm. Perfora­ heartwood dark-brown, sharply defined from tions simple, end walls almost transverse. Inter­ the light-brown sapwood. Texture medium to vascular pits alternate, round and lor polygonal, coarse, grain slightly interlocked. Specific gravi­ 6-9 !lm. Vessel-ray and vessel-parenchyma pits ty 600-700 N per cubic metre (in P. !impata larger, elongated, more irregular-shaped, half­ 780-980 N per cubic metre). bordered, the borders reduced. Thin-walled M i c r 0 s cop i c f eat u res: Vessels diffuse, tyloses sometimes present, but always scarce. solitary (50-84%) and in short radial multiples Fibres nonseptate, with small simple pits re- and irregular clusters of 2-4; 2-4 per sq.mm,

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round to slightly oval, diameter 190-220 !Lm (Uw 20913); Surinam: Oldenburger et al. 1402 (130 !.!ill in P. !impato, Uw 24438), vessel member (Uw 17968). - S. trophoides Burger ssp. rhodo­ length 350-450 !.!ill. Perforations simple, end rachis Cuatrec. Colombia, Choco: Cuatrecasas walls almost transverse. Intervascular pits alter­ 14891 (Uw 25362), 15572, type (Uw 25436). nate, round and polygonal, 9-12 !Lm. Vessel­ G e n era I f eat u res: Growth rings absent ray and vessel-parenchyma pits larger, elongated, or faint; yellowish to light brown, no differ­ more irregular-shaped, half-bordered, the bor­ ence between heartwood and sapwood. Tex­ ders often reduced. Thin-walled tyloses com­ ture fine, grain straight, sometimes slightly in­ mon, varying from abundant to scarce. Fibres terlocked. Specific gravity 500-1000 N per all or in part septate, several septa per fibre, cubic metre. with small simple pits restricted to the radial M i c r 0 s cop i c f eat u res: Vessels diffuse, walls; walls 2-5 J!.m, lumina 7-14 J!.m, partly solitary (20-63%) and in short radial mUltiples gelatinous in P. frutescens; length 1230-1420 and irregular clusters of 2-4 (5); 8-21 per mm, !.!ill; F/V-ratio 3.1-3.5. Rays uniseriate and round to slightly oval, diameter 68-130 J!.m, multiseriate, heterogeneous type III, 4-7 per vessel member length 325-475 !.!ill. Perfora­ mm. Uniseriate rays 13-20%, composed of up­ tion simple, end walls almost transverse. Inter­ right and square cells, only occasionally some vascular pits alternate, round and sometimes procumbent cells, height up to 4-7 cells (240- polygonal, 6-12 J!.m. Vessel-ray and vessel­ 420 J!.m). Multiseriate rays composed of pro­ parenchyma pits larger, elongated, half-border­ cumbent cells except for the uniseriate margins ed, the borders often reduced. Thin-walled ty­ of 1-3 rows of square and/or upright cells; loses common. Fibres nonseptate, with small 3-5 cells wide, up to 720-1200 J!.m (inP. fru­ simple pits restricted to the radial cell walls; tescens up to 480 !.!ill) high. Occasionally with walls 2-4 J!.m, lumina 4-12 !.!ill, gelatinous some sheath cells. Parenchyma vasicentric-ali­ fibres common; length 1080-1460 !.!ill; F/V­ form with short to long wings, occasionally ratio 2.5-3.9. Rays uniseriate and multiseriate, confluent or in short (wavy) bands. Parenchy­ heterogeneous type II/III, 6-9 per mm. Uni­ ma strands of 4 cells. Vitreous silica common seriate rays 7-40%, composed of upright and in the vessels, enveloping the tyloses; occasion­ square and occasionally few procumbent cells, ally also in the fibres and sporadically in the height up to 6-19 cells (180-600 J!.m). Multi­ axial parenchyma. seriate rays composed of procumbent cells ex­ cept for the uniseriate margins of 1-3 (10) Sorocea St. Hil. (Fig. 22) rows of square and/or upright cells; 2-5 cells The genus has recently been monographed wide, up to 660-1200 !.!ill high. Parenchyma by Berg and Akkermans (1985), who recognised scanty paratracheal, but predominantly as con­ 16 species. The genus Paraciarisia was already centric, sometimes more or less wavy bands en­ included in Sorocea by Burger et al. (1962). closing the vessel in part, sometimes also termi­ Sorocea is distributed in the Neotropics, from nal bands present, 2-9 cells wide and 2.7-5 Guatemala to Paraguay and Argentina, and in bands per mm. Parenchyma strands of 3-4 Curacao. The shrubs or trees are up to 6-20 m cells. Rhombic crystals in the square and up­ high, and are generally components of the un­ right cells from few to abundant, and less fre­ derstory in moist, non-flooded forests, from quent to occasionally absent in the axial paren­ sea level up to 2000 m altitude. chyma cells. Radial latex tubes observed in one Mat e ria 1st u die d: S. bonplandii (Bail­ sample. Ion) W. Burger, Lanjouw & W. Boer. Brazil, Not e s. The wood anatomy of the genus is Santa Catharina: Reitz 15026 (Uw 6409), Para­ very homogeneous. One species, S. muriculata, na: Lindeman & H. de Haas 1988 (Uw 13437), can be identified using the average vessel diam­ 2842 (Uw 13928). - S. duckei W. Burger. Brazil, eter, being 68-70 J!.m in this species, and 90- Amazonas: F. Mello, INPA X-3745 (Uw 23664). 130 J!.m in the other species. - S. faustiniana Cuatrec. Colombia, Choco: Cuatrecasas 15204 (Uw 24928). - S. guillimi­ Sparattosyce Bureau (Figs. 23, 24) niana Gaudich. Brazil, Amazonas: Prance et al. A genus comprising two species, restricted to 18248 (TJw 20902), 19825 (Uw 20941), Matto New Caledonia and occurring in moderately Grosso: Krukoff 1349 (Uw 19305), 7039 (Uw high forest as trees of up to 8 m high. 8126),6143, type (Uw 7513), 6223 (Uw 7568). Mat e ria 1st u die d : S. dioica Bureau. New - S. hirtella Mildbread. Brazil, Acre: Krukoff Caledonia: Sarlin G 153, ex C.T.F.T. 6201 (Uw 5084 (Uw 19745), 5785 (Uw 20188); Peru: 24627), s. n., ex SJR w 14688 (Uw 18425), s. n., Schunke V. 4827 (Uw 21086). - S. muriculata ex SJRw 14301 (Uw 18424). Miq. Brazil, Amazonas: Krukoff 6988 (Uw General features: Growth rings absent; 8082), 8520 (Uw 16200); Prance & Berg 18468 light yellow brown, unknown whether dealing

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with heartwood or sapwood. Texture medium and irregular clusters of 2-5; 3-5 (10) per mm, to coarse, grain straight. Specific gravity 500- round to slightly oval, diameter 145-205 J.Lm, 540 N per cubic metre. vessel member length 400-520 J.Lm. Perfora­ M i c ro s cop i c fe a t u re s: Vessels diffuse, tions simple, end walls almost transverse. Inter­ solitary (25-50%) and in short radial mUltiples vascular pits alternate, round and/or polygonal, and occasionally irregular clusters of 2-5; 4-9 7-9 J.Lm. Vessel-ray and vessel-parenchyma pits per sq. mm, round to slightly oval, diameter larger, elongated, half-bordered, the borders in 130-160 J.Lm, vessel member length 425-530 part reduced. Thin-walled tyloses common, but J.LID. Perforations simple, end walls almost trans­ scarce. Fibres nonseptate, with many small sim­ verse. Intervascular pits alternate, round and lor ple pits in the radial walls, occasionally also in polygonal, 9-11 J.Lm. Vessel-ray and vessel-paren­ the tangential walls; walls 3-5 J.Lm, lumina 9- chyma pits larger, elongated, half-bordered, the 20 J.Lm, gelatinous fibres occasionally present; borders often reduced. Very few thin-walled length 1245-1575 J.Lm; F/V-ratio 2.7-3.7. tyloses occasionally present. Fibres septate, Rays uniseriate and multiseriate, heterogeneous several septa per fibre, but also nonseptate in type II/III, 4-6 per mm. Uniseriate rays 5-20 the same sample, with small simple pits restrict­ %, composed of upright and square cells, height ed to the radial walls; walls 3-4 J.Lm, lumina 7- up to 3-5 cells (180-360 J.Lm). Multiseriate 20 J.Lm, occasionally gelatinous; length 1080- rays composed of procumbent cells except for 1255 J.LID; F/V-ratio 2.1-2.6. Rays uniseriate the uniseriate margins of 1-4 rows of upright and multiseriate, heterogeneous type II/III, and/or square cells, 3-5 cells wide, up to 600- 6-7 per mm. Uniseriate rays 8-23%, composed 660 J.Lm high. Occasionally with some sheath of square, upright and some procumbent cells, cells. Parenchyma aliform with long narrow height up to 6-13 cells (180-600 J.Lm). Multi­ wings, confluent and sometimes in short wavy seriate rays composed of procumbent cells ex­ bands. Parenchyma strands of 2 cells. Rhombic cept for the uniseriate margins of 1-2 (in Uw crystals common in the square and upright 18425 of 1-6) rows of square and lor upright marginal ray cells, few, and very few in the two cells; 3-5 cells wide, up to 720-1450 J.Lm high. samples from Zaire. Radial latex tubes com­ Occasi9nally with a few sheath cells. Parenchy­ mon, the diameter of the tubes large compared ma in concentric bands, partly enclosing the with the diameter of the ray cells (tangential vessels, 4-14 cells wide and 1.4-2 per mm, sections). and some scanty paratracheal strands. Paren­ chyma strands of 2-4 cells. Rhombic crystals abundant in the axial parenchyma cells, and Discussion few in the ray cells. Vitreous silica in the fibres and the axial parenchyma of one sample (Uw Wood anatomical variation within genera 18425). Most studied genera are very homogeneous in their wood anatomical characters. This holds Treculia Decaisne (Figs. 25, 26) in particular for Artocarpus, Bagassa, Pararto­ A recent monograph of the genus by Berg carpus, Poulsenia, Sparattosyce and Treculia. (1977b) comprises three species. The genus is Within Batocarpus, slightly more variation is distributed as trees or shrubs from Senegal to found in the amount and distribution of the Sudan, Angola, Mo~ambique, and Madagascar. parenchyma (Figs. 9, 10). Species available for this study are trees up to In Prainea, the sample of P. !impato deviates 30( -50) m high, from humid to dry primary by its narrow vessels and high specific gravity. forests, ,swamp forest or secondary vegetation, The sample of P. frutescens shows relatively from sea level to 1300 m altitude. low multiseriate rays. These few exceptions do Mat e ria 1st u die d: T. africana Decaisne. not disturb the impression of Prainea as a natu­ Ivory Coast: Detienne 117 (Uw 24640); Came­ ral, well defined genus. roon: Foury 131 (Uw 25687); Sao Tome: Museu The wood anatomy of Sorocea is also homo­ Ultramar no. 30 (Uw 26978); Zaire: Louis 2507 geneous. The fact, that the vessel diameter of (Uw 24242), Inst. Nat. Agron. 3144, ex MADw S. muriculata is lower than those of the other 17257 (Uw 26977). species (see genus description), is not due to a Ge n e r al fe a t ure s: Growth rings absent; dry habitat. Only one of the samples studied light brown, no differences observed between by us was collected on a dry slope in the Sip ali­ sapwood and heartwood. Texture medium to wini Savanna in southern Surinam showing that coarse, grain straight. Specific gravity 480-670 the species can endure water stress in this area N per cubic metre. with its long dry season and low annual rainfall. M i c r 0 s cop i c f eat u res: Vessels diffuse, However, according to Burger et al. (1962), solitary (32-50%) and in short radial multiples this species occurs in 'moist forests and the

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mata de terra firma and the Varzeas.' In other right ray cells only. Crystals are lacking in Anti­ aspects, S. muriculata also completely fits in aropsis, Artocarpus, Batocarpus (mostly), Poul­ with the samples of other species, at least two senia and Prainea. In Clarisia, crystals vary not of which are known from an area with an an­ only quantitatively, but also in distribution. nual rainfall of more than 7000 mm (i.e., Rhombic crystals are absent in C. biflora. Few Choco, near Bajo Calima, Colombia): S. fausti­ to very few rhombic crystals occur in C. race­ niana and S. trophoides ssp. rhodorachis. mosa: they are absent in three samples; in one The genus Clarisia is much less homogeneous. sample very few rhombic crystals were observ­ Three species were recognised by Berg (1968). ed in the ray cells; in three samples from Vene­ The wood anatomical characters can be used to zuela and Colombia few to very few crystals identify the individual species quite easily (see occur in the axial parenchyma cells. The occur­ Table 1 on page 185). In 1936 the genera Clari­ rence of the crystals in C. ilicifolia follows a sia and Acanthinophyllum were revised by Lan­ comparable pattern: three samples lack crystals; jouw. He concluded that they are congeneric. in one sample very few crystals occur in the ray A monograph including these two genera was cells; in the three other samples many to few published in 1962 by Burger. He separated the crystals occur in the ray cells and in the axial species with capitate pistillate inflorescences as parenchyma cells. a distinct genus, Acanthinophyllum, from Cla­ risia, leaving only C. biflora and C. racemosa in Vitreous silica Clarisia. In Acanthinophyllum two species, viz. The quantity of vitreous silica is always A. ilicifolium and A. spruceanum, were recog­ (very) variable. It may surround the tyloses, nised. The most recent taxonomic discussion but in some samples it completely fills up the on this problem is by Berg (1968). In his opin­ smaller vessels. Fibres and axial parenchyma ion, A. spruceanum, only known from the type and occasionally also some of the ray cells may collection, cannot be distinguished from the be completely or partly filled. very variable A. ilicifolium. As a result Berg in­ As can be seen in Table 2, silica is always cluded A. spruceanum in A. ilicifolium. Berg present in Artocarpus, Poulsenia and Prainea; also re-evaluated the characters used by Burger it was never found in Antiaropsis, Bagassa, Par­ (1962) to separate A. ilicifolium from Clarisia. artocarpus, Sorocea and Treculia. His conclusion is that Acanthinophyllum does not merit generic rank. Consequently, A. ilici­ Axial and radial latex tubes folium was transferred again to Clarisia. The A study of the occurrence of axial and radial wood anatomical differences as observed be­ latex tubes in Artocarpus was carried out by tween the three species thus accommodated in Topper and Koek-Noorman (1980). They con­ Clarisia are substantial (Table 1). Based on cluded that the value of axial latex tubes in these differences one could defend a genus taxonomy seems to be restricted. Although rank for C. ilicifolia as suggested by Burger radial latex tubes were recorded for all species (1962). Taking into account the variation range of Artocarpus studied, in three species bpth within other, larger genera, however, we decided specimens with and specimens without radial to' follow the genus concept of Berg (1968). latex tubes were encountered. The radial latex tubes can be classified into Crystals two types, differing in relative size: Rhombic crystals are rather common in the genera studied. The crystals occur in the ray 1. radial latex tubes with a diameter more or cells, and less frequently in the axial paren­ less equal to the surrounding ray cells as seen chyma cells. The number of crystals is general­ in tangential sections; ly known to be variable from sample to sample. 2. radial latex tubes with a diameter (much) Also in the material used for this study, the larger than the surrounding ray cells. amount of crystals varies considerably. How­ Tubes of type 2 are easy to detect, even with­ ever, presence and distribution of crystals, out a special staining. However, tubes of type 1 regardless of quantity, distinguishes (groups of) can only be observed if the contents of the genera (Table 2). tubes have a colour distinct from that of the Bagassa and Sorocea are similar in their crys­ ray cells and are retained in the sections, or tal distribution (predominantly in square and when special staining with Sudan III is applied. upright ray cells but also few in procumbent The results presented here are partly based on ray cells and axial parenchyma). Parartocarpus observations of unstained sections. Therefore, and Sparattosyce show crystals in axial paren­ further studies, using specific staining tech­ chyma (Sparattosyce also occasionally in ray niques, may eventually reveal the occurrence of cells). Treculia shows crystals in square and up- radial latex tubes of type I in more taxa.

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Table 2. Wood anatomical characters of the genera of the Moreae without urticaceous stamens.

Vessels Fibres Rays ,. ~ ~ ::J. E 3 ~'-' c;;.. ~ ...'" ::J. -5 '" Q) OJ) ] [ ~ t::; -.5 ::J. 3 ~ '" ;:., ... >'" () t::; ... 3 .9 Q) Q) ;:., t::; Q) '-' ....'" Q) -:;; p.. ;:., ~ Q) Genus :a'" E :=E Downloaded fromBrill.com10/06/2021 06:03:46AM Poulsenia 105-210 670-695 35-90 4-10 6-9 1410-1415 2.0-2.1 III > ttl Prainea 130-220 350-450 50-84 2-4 9-12 1230-1420 3.1-3.5 E. + III [ Sorocea 68-130 325-475 20-63 8-21 6-12 1080-1460 2.5-3.9 II/III 5· ;::l

Sparattosyce 130-160 425-530 25-50 4-9 9-11 + 1080-1255 2.1-2.6 II/III ~ Treculia 145-205 400-520 32-50 3-5 (10) 7-9 1245-1575 2.7-3.7 II/III <: ?- -.J W via freeaccess 'D 00 0\ > Table 2 (continued) ::E! > tl:I Ray s Parenchyma E. < a (D ~ s '" .... s· a ~ - >-~ .... ,n '" ~ "'~ ( ~ < >- ~

Downloaded fromBrill.com10/06/2021 06:03:46AM Sorocea 660-1200 2-5 6-9 (+) (+) + 2-9 2.7-5 u,p,a Sparattosyce 720-1450 3-5 6-7 (+) + 4-14 1.4-2 a (p) (f, p) Treculia 600-660 3-5 4-6 + (+) + (2) u

Legend: +: present; -: absent; between brackets: occasionally present. - Rays: II, III: heterogeneous types II, III, according to Kribs (1968); ho: homogeneous, i.e., composed of procumbent cells only. - Parenchyma distribution: d: diffuse; sp: scanty paratracheal; v: vasicentric; as: alifonn; al: alifonn with long wings; c: confluent; wb: wavy bands; cb: concentric bands. - Radial latex tubes: I: diameter of the tubes more or less equal to the diameter of the ray cells (in tangential view); 2: diameter of the tubes (much) larger than the diameter of the ray cells (tangential view). - Crystal distribution: u: in square and upright cells; p: in procumbent ray cells; a: in axial parenchyma cells. - Vitreous silica: f: in fibres; p: in parenchyma; v: in vessels; vt: in tyloses in the vessels. via freeaccess \C 192 IAWA Bulletin n.s., Vol. 7 (3),1986

As far as we know, radial latex tubes are vitreous silica, and latex tubes, parenchyma dis­ present in all samples of: tribution and the occurrence of septate fibres Antiaropsis : type 2, radial, diameter approx. are characters sometimes useful in classification. 28 }.Lm, few; Three genera are considered to be closely al­ Artocarpus : type I, radial and axial (in part), lied by all taxonomists: Artocarpus, Pararto­ 10-20 }.Lm, relatively abundant; carpus and Treculia. Jarrett (1959a) mentions Bagassa : type I, radial and occasionally Prainea as a close ally of Artocarpus with Parar­ axial, 12-20 }.Lm, few; to carpus and Treculia in a less closely allied Parartocarpus : type 2, radial, diameter 20-48 series. We found no particular wood anatomical Jlm, usually abundant; features to support a close relationship between Poulsenia : type 2, radial, diameter (10) 16- these four genera, despite their possession of 24 Jlm, few; vasicentric-aliform (to confluent) parenchyma. Treculia : type 2, radial, diameter 24-48 This pattern is also found, however, in Bagassa Jlm, few to abundant. and Poulsenia, which were never placed in the The genus Clarisia is problematic with respect same tribe with Artocarpus and allies before. to the presence of radial latex tubes. They are The genera with concentric parenchyma absent in C. racemosa and C. biflora. In C. ilici­ bands were formerly placed in two different folia three of the four specimens from Surinam groups: Clarisia and Sorocea in subtribe Euarto­ show some radial latex tubes of type I (diam­ carpeae (Engler, 1888) and Moreae (Corner, eter approx. 12 Jlm). 1962); Antiaropsis and Sparattosyce (the latter One latex tube of size type 2 was observed in Ficeae according to Engler, 1888) in Olme­ in Sorocea hirtella (Uw 19745). As none of the dieae (Corner, 1962). The Olmedieae (= Castil­ other Sorocea samples studied showed any leae sensu Berg, 1977 a) are characterised by the latex tubes, the genus Sorocea is regarded es­ presence of septate fibres (Mennega & Lanzing­ sentially as a taxon without latex tubes. Finally, Vinkenborg, 1977). Septate fibres also occur in no latex tubes were found in the genera Bato­ Antiaropsis and Sparattosyce (this paper) and carpus, Prainea, and Sparattosyce. Olmedia (Ter Welle et aI., 1986). Within the Based on these data, the occurrence of radial sensu Berg the parenchyma pattern latex tubes may be regarded as a useful charac­ varies from aliform-confluent to banded (Koek­ ter for generic diagnosis, even if we keep in Noorman et aI., 1984), and we found no wood mind the possibility that narrow latex tubes anatomical arguments to exclude Antiaropsis, (type I) may have been overlooked in some Sparattosyce and Olmedia from the Olmedieae. taxa until now. Prainea is the only genus with septate fibres, which has never assigned to the Olmedieae. Identification of the individual genera All individual genera studied are (highly) It will be evident from the foregoing (see also homogeneous, except Clarisia, where several Ter Welle et aI., 1986), that wood anatomical wood anatomical characters vary from species characters within the Moreae can be used to to species, or even within the species. identify (groups of) genera, but that comments In contrast, the wood anatomy on the inter­ on the delimitation of this tribe can only be generic level is heterogeneous. Most individual made when the family as a whole is taken into genera may be identified using the following consideration. This will be the subject of a sub­ features: the occurrence of septate fibres (Anti­ sequent paper. aropsis, Prainea, Sparattosyce); ofrhombiccrys­ tals (Bagassa, C/arisia, Parartocarpus, Sorocea, Acknowledgements SparattosY.ce, Treculia), homogeneous rays (Cla­ Several persons, in one way or another, con­ risia, Parartocarpus), and the parenchyma pat­ tributed in the preparation and presentation of tern: bands in contrast to a vasicentric or ali­ this series of five publications on the wood form-confluent distribution (see Table 2). The anatomo/ of the Moraceae. only problem is found in C/arisia, as the three In the first place we wish to thank the cura­ species appear to be variable in all these features tors of the wood collections all over the world. with the exception of absence of septate fibres. Owing to their help and willingness in sending us duplicates and answering our questions con­ Taxonomic affinities cerning collectors and localities, we were able The II genera studied, now united in the to study almost all genera of the Moraceae, group of Moreae without urticaceous stamens using exclusively well-vouchered material. Spe­ (Berg, 1983), have been assigned to several tribes cial thanks are due to the curators of the wood and subtribes in the past (see Introduction). collections in Madison (U. S.A.), Hamburg and Next to the presence or absence of crystals, Berlin (F.R.G.), Tervuren (Belgium), Leiden

Downloaded from Brill.com10/06/2021 06:03:46AM via free access IAWA Bulletin n.s., Vol. 7 (3),1986 193 and Amsterdam (the Netherlands), Nogent-sur­ - 1959b. Ibid. II. A revision of Prainea. J. Marne (France), and Kew and Oxford (U.K.), Arn. Arbor. 40: 30-37. who together provided about 85 % of the taxa - 1959c. Ibid. III. A revision of Artocarpus not represented in the Utrecht wood collection. subgenus Artocarpus. J. Arn. Arbor. 40: Drs. L. Y. Th. Westra and G. Schatz kindly cor­ 113-155. rected the English text of the manuscripts. Mr. - 1959d. Ibid. III. Ibid. (continued). J. Arn. E. Elsendoorn, Mr. M.H. Rypkema, and Mr. T. Arbor. 40: 298-326. Schipper prepared the photographs and plates. - 195ge. Ibid. III. Ibid. (continued). J. Am. The discussions with Dr. A.M.W. Mennega and Arbor. 40: 327-368. Dr. C. C. Berg (Utrecht), and Dr. P. Baas (Leiden) - 1960a. Ibid. IV. A revision of Artocarpus have done much to improve the manuscripts. subgenus Pseudojaca. J. Arn. Arbor. 41: 73-109. References - 1960b. Ibid. IV. Ibid. (continued). J. Am. Bentham, G. & J.D. Hooker. 1880. Genera Plan­ Arbor. 41: 111-140. tarum 3 (I). : 341-395. London. - 1960c. Ibid. V. A revision of Parartocarpus Berg, C. C. 1968. Taxonomic and morphological and Hullettia. J. Arn. Arbor. 41: 320-340. notes on Clarisia (Moraceae).Acta Bot.Neerl. Koek-Noorman, J., S.M.C. Topper & B.J.H. ter 17: 309-312. Welle. 1984. Systematic wood anatomy of - 1975. Flora of Suriname V (I). Moraceae: Moraceae (Urticales). I. Tribe Castilleae. 173-299. Brill, Leiden. IAWA Bull. n.s. 5: 183-195. - 1977a. The Castilleae, a tribe of the Mora­ Kribs, D. A. 1968. Commercial foreign woods on ceae, renamed and redefined due to the ex­ the American market. Dover Pub., New York. clusion of the type Olmedia from the '01- Lanjouw, J. 1936. Studies in Moraceae II. The medieae'. Acta Bot. Neerl. 26: 73-82. genus Clarisia Ruiz et Pavon and its syn­ - 1977b. Revisions of African Moraceae (excl. onyms with a discussion of the generic , , Musanga and Myrianthus). name. Rec. Trav. Bot. Neerl. 33: 254-276. Bull. Jard. Bot. Nat. Belg. 47: 267-407. Mello Filho, L. E. & M. Emmerich. 1968. Revi­ - 1983. Dispersal and distribution in the Ur­ sao do genero Batocarpus Krst. (Moraceae­ ticales - an outline. Sonderbiinde Naturw. Euartocarpeae). Bol. Mus. Nat. Bot. 37: 3 - 21. Ver. no 7: 219-229. Mennega, A.M.W. & M. Lanzing-Vinkenborg. - & R.W.A.P. Akkermans. 1985. Studies on 1977. On the wood anatomy of the tribe the flora of 14. New taxa and 'Olmedia' (Moraceae) and the position of combinations in Sorocea (Moraceae) and a the genus Olmedia R. & P. Acta Bot. Neerl. key to its species. Proc. Kon. Ned. Akad. 26: 1-27. Wet. ser.C, 88: 381-395. Siooten, H.J. van. 1968. Investigacion y desar­ Burger, W.C. 1962. Studies in the New World rollo de zonas forestales selectas de Costa Moraceae: , Clarisia, Acanthinophyl­ Rica. Inst. Interamer. Cienc. Agricolas, lum. Ann. Missouri Bot. Gard. 49: 1-34. Turrialba, Costa Rica. - , J. Lanjouw & J. G. Wessels Boer. 1962. The Topper, S.M.C. & J. Koek-Noorman. 1980. The genus Sorocea St. Hil. (Moraceae). Acta Bot. occurrence of axial latex tubes in the sec­ Neerl. II: 428-477. ondary xylem of some species of Artocar­ Corner, E.J.H. 1962. The classification ofMo­ pus J.R. & G. Forster (Moraceae). IAWA raceae. Gard. Bull. Sing. 19: 187-252. Bull. n.s. I: 113-119. - 1976. A new species of Parartocarpus (Mo­ Welle, B.J.H. ter, J. Koek-Noorman & S.M.C. raceae). Gard. Bull. Sing. 28: 183-190. Topper. 1986. The systematic wood anato­ Engler, A. 1888. Moraceae. In: Die naturlichen my of the Moraceae (Urticales) IV. Genera Pflanzenfamilien 3 (I) (ed. A. Engler & of the tribe Moreae with urticaceous sta­ K. Prantl): 66-98. Leipzig. mens. IAWA Bull. n.s. 7: 91-128. Jarrett, F.M. 1959a. Studies in Artocarpus and Williams, L. 1936. Woods of Northeastern Peru. allied genera. I. General considerations. J. Botanical Series, Field Museum of Natural Arn. Arbor. 40: 1-29. History, Vol. XV. Chicago.

ERRATUM

In Koek-Noorman, J., S.M.C. Topper & B.J.H. ter Welle (1984), The systematic wood anatomy of the Moraceae (Urticales) II. Tribe Dorstenieae, IAWA Bulletin n. s. 5 (4): 317-329, discrepancy be­ tween the description of Dorstenia and the data of Table 2 has escaped our attention. The table should be corrected in accordance with the text.

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