IAWA Bulletin n.s., Vol. 5 (3),1984 183

THE SYSTEMATIC WOOD ANATOMY OF THE (URTICALES) I. TRIBE 1

by

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

Summary The wood anatomy of the tribe Castilleae The main aim of the present research project sensu Berg is described. Similarities and differ­ is to find an answer to the following questions: ences are discussed in relation to his concepts What is the anatomical variability range in taxa of the of the tribe. The wood anato­ of Moraceae as compared to c10sely related mical variation does not enable to distinguish taxa? - To what extent can the anatomical in­ between , and . fonnation be used to contribute to our under­ , , , and standing of patterns of relationship, and aid in can be recognised on the basis of generic and tribai delimitation? slight differences. However, no reasons are A thorough treatment of the wood anatomy found to question the delimitation of the Cas­ of the tribe Olmedieae was published by Men­ tilleae sensu Berg on the basis of their wood nega and Lanzing-Vinkenborg (1977). One of anatomy. their conc1usions was that the exclusion of 01- Key words: Wood anatomy, systematics, media (Berg, 1977a) was justified. They did Moraceae, Castilleae. not discuss the relation of the Olmedieae (later renamed Castilleae) to the other Moraceae. Be­ Introduction sides, they did not study the African genera The present paper is the first one of aseries Antiaris and Mesogyne, also placed in the Cas­ dealing with the systematic wood anatomy of tilleae by Berg (1973). Therefore we decided to Moraceae. It includes wood anatomical descrip­ inc1ude the Castilleae in our survey of the tions of the genera of the tribe Castilleae. This wood anatomy of Moraceae. We intend to pre­ project follows in logical sequence the work of sent aseries of papers, in each of which the C.C. Berg, taxonomist of the family, who had genera of one tribe are described and their experienced problems in delimiting and defining wood anatomical similarities and differences the family as a whole, as weH as the individual are discussed in relation to ideas about delimi­ genera and tribes. The wood and leaf anatomy tation and relationships as brought forward by, of the Cecropiaceae, asegregate of the Mora­ La., Corner (1962, and his unpublished Flora ceae, in relation to the taxonomy of this group Malesiana account of 1972) and Berg (1972, has already been discussed in a paper by Bonsen 1977a, b, 1983). We will follow the most re­ and Ter Welle (1983). Their study on the sys­ cent views of Berg on the tribai arrangement, as tematic anatomy of the Urticaceae, c10sely re­ indicated by hirn in his paper on distribution lated to the Moraceae, is in the press (1984). and dispersal in the Urticales (1983). Most wood anatomical literature on Mora­ We intend to publish separate papers on the ceae is rather difficult to interpret taxonomi­ tribes ofthe Moraceae in thefollowing sequence: cally, due to the often dubious identifications I. Castilleae (Antiaris, Castilla, Helicostylis, and the frequent nomenclatural changes. Fur­ Maquira, Mesogyne, Naucleopsis, Perebea, thermore, descriptions are nearly always based Pseudolmedia) ; on few sam pies and relate only to individual 11. Dorstenieae (Bosqueiopsis, , Dor­ genera or commercially important timbers. stenia, Helianthostylis, , Tryma­ Numerous accounts in the literature were com­ tococcus; of and Vtsetela no pared with our findings. Here we only refer to material is available); literature inc1uding elaborate wood anatomical III. Ficeae (); descriptions based on reliably identified mate­ IV. Moreae p.p., characterised by the presence rial, and not mentioned elsewhere in the text: of 'urticaceous' stamens (, Cal­ Janssonius, 1934; Koek-Noonnan & ter Welle, pidochlamys, Cardiogyne, Chlorophora, 1976; Record & Hess, 1940; Tippo, 1938. s.1. incl. Plecospermum and Cudra-

1 This project was made possible by a grant of BION-ZWO (14.45-01).

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nia, Maillardia, Malaisia, Militia, , Pa­ Finally, the specific gravity was established chytrophe, s.I., s.1. incl. in the usual way. Measuring of the moisture Olmedia ; of Ampalis and Fatouia no mate­ content of the sampies was considered unneces­ rial is available); sary as all sampies were stored in a room with V. Moreae p.p., characterised by the absence central heating and an air humidity of c. 40% of 'urticaceous' stamens (, Arto­ during at least several weeks. It is the general carpus, , , , Parar­ experience that this results in a relative mois­ tocarpus, , Prainea, , Spa­ ture content of approx. 12%. rattosyce, ; of Hullettia no material is available). In a concluding paper we intend to discuss Generic descriptions the taxonomie value of the various wood ana­ tomical features on the tri bai level and to com­ Antiaris Lesch. (Figs. I, 2) pare the individual tribes. From the 17 , described since the was established, Corner (1962) reduced the number to four, but he already pointed out Material and Methods that there were no essential differences between Most wood sam pies studied are backed by these species. Berg (l977b) combined all spe­ herbarium vouchers which were identified by eies in Antiaris toxicaria Lesch., lowering the C. C. Berg, the taxonomist of the family. Most rank of these four species to subspecies. The sampies were taken from the trunk of the tree; genus Antiaris (trees, up to 40-60 m tall, or sometimes, however, the origin was unknown. shrubs) occurs in the Palaeotropics in primary Many wood sam pies were provided by institu­ and secondary forests in wet and dry habitats, tional wood collections all over the world. The from sea level up to 1500 m altitude. Utrecht wood collection served as a first basis M a t e r i als t ud i e d: A. toxicaria Lesch. for this study. In the generic descriptions, only Uganda: B. T. Styles 201 (Uw 19390), For. those sampies are cited of which sections were Dept. 8-1950 (Uw 18387); Zaire: C. Donis 413 made. The general features of the wood are (Uw 24211), J. Louis 11201 (Uw 18780); West often based upon the study of more wood Africa: F.P.R.L. 209A (Uw 18386); EastAfri­ sam pies not cited here. Seetions and macera­ ca: H.E. Desch-F.R.I.C.S. (Uw 24305); Malay­ tions were prepared according to standard tech­ sia: For. Dept. 9-1954 (Uw 18388); Indonesia, niques and embedded in Canada balsam and in Java: Koorders 8316t/12757B (Uw 24417), glycerin respectively. The sections were stained 1127g/22746B (Uw 24418), 1898m/13415B with safranin and the macerations in astrablue. (Uw 24430); Indonesia, Irian Jaya: BW 11519 Besides, unstained sections were also studied. (Uw 20481), BW 11916 (Uw 18169), Fokkinga The terminology proposed by the Commit­ 1573 (Uw 24306); Philippines: Bur. of For., tee on Nomenclature of the IAWA (1964) is Manila P.1. 21092b.f. (Uw 24431). followed. As for the quantitative data: vessel General features: Growth rings faint diameters were measured in tangential direc­ or absent; colour light brown, no difference be­ tion inc1uding the walls and averages are based tween sapwood and heartwood. Texture coarse; on 25 measurements. Vessel frequency is based grain interlocked. Specific gravity 270-530 N on 25 counts. In the descriptions, for both per cubic metre. eharaeters minimum and maximum averages Mi c r 0 s e 0 pie fe a t ure s: Vessels diffuse, found in sam pies of eaeh genus are provided. in some sam pIes tending towards a diagonal ar­ The pereentage ofsolitary vessels was ealculated rangement; solitary (45-65%) and in short ra­ after examining an area with at least 100 spores. dial multiples or irregular clusters of 2-6; 3- Clusters and multiples were regarded as 2, 3, 4, 10 per sq.mm, round to oval, diameter 145- ete. vessels, depending on the number of vessels 230 JJ.m, vessel member length 425-575 JJ.m. per group. Vessel member length and fibre Perforations simple, end walls transverse. Inter­ length are based on at least 100 measurements vaseular pits alternate, round, oval or polygonal, per sampie, and again, minimum and maximum 9-13 JJ.m. Vessel-ray and vessel-parenchyma sampie averages found in the genera are pro­ pits larger and irregularly shaped, half-bordered. vided. Additionally, the averages were used to Thin-walled tyloses occasionally present in ealculate the ratio of fibre length/vessel mem­ sampies from Asia. Fibres septate with small ber length, in the descriptions referred to as simple pits restricted to the radial walls; walls F/V-ratio. For the fibres, maximum wall thiek­ 1-3 JJ.m, lumina 10-30 JJ.m; often gelatinous; ness and maximum lumen diameter are given. length 1030-1390 JJ.m; F/V-ratio 1.8-3.1. The percentage of uniseriate rays is based on Rays uniseriate and multiseriate, 4-7 per mm. counts of at least 100 rays. Uniseriate rays 2-15%, composed of procum-

Downloaded from Brill.com09/26/2021 01:46:11AM via free access IAWA Bulletin n.s., Vol. 5 (3),1984 185 bent and few square or upright cells, height up or partly septate with sm all simple pits restrict­ to 10 cells (430 .um). Multiseriate rays com­ ed to the radial walls; walls 1-3 .um, lumina posed of procumbent cells', uniseriate margins 15-25 .um, occasionally gelatinous; length of 1-2 (4) rows of square and/orupright cells 1000-1200 J..lm; F/V-ratio 2.0-2.9. Rays uni­ and occasionally few sheath ceIls; 4-7 cells seriate and multiseriate, 4-7 per mm. Uniseriate wide and up to 1100 J..ln! high. Parenchyma pa­ rays 0-25%, composed of procumbent and up­ ratracheal, vasicentric-aliform with short wings, right cells, height up to 6-10 cells (150-500 occasionally confluent, parenchyma strands of J..lm). Multiseriate rays composed of procum­ 3-5 (8) cells. Radial latex tubes present in all bent cells', with uniseriate margins of 1-3 rows sampies. Rhombic crystals few, in marginal ray of square and/or upright cells and few sheath cells and axial parenchyma in some sampies cells in some rays; 3-6 cells wide, up to 800- from Asia. 1300 J..lm high. Parenchyma paratracheal, the Not e: Rhombi.: and elongated crystals are pattern varying from alm ost restricted to vasi­ extremely abundant in the axial parenchyma centric or alif6rm with short wings to confluent­ and marginal ray cells of the sam pie of Koorders banded, the bands up to 20 cells wide and 1-2 11 27g/22746B. per mm. Parenchyma strands of 2-4 (8) cells. Radial latex tubes present in all sampIes. Vitre­ Castilla Sesse in Cervantes (Figs. 3, 4) ous silica present in some vessels in two sam pies. Berg (1972) monographed the genus and dis­ Not e: Castilla uZei (Williams 1802) devia­ tinguished three species: C. eZastica Cerv., C. tes in the high uniseriate rays (up to 30 cells, tunu Hemsl., and C. uZei Warb. The genus Cas­ 650 J..lm) and the narrow multiseriate rays (up tilla occurs in Central America, the coastal re­ to 3 cells wide) with uniseriate margins of 3-6 gion of NW. South America, and the Amazon rows of uprigh t cells. basin. Castilla eZastica has been introduced and naturalised in most tropical countries. Castilla Helicostylis Trecul (Fig. 13) consists of trees, about 30-40 m tall, growing This Neotropical genus was monographed by in tropical rainforest up to 850 m altitude. Berg in 1972. He distinguished seven species, M a t e r i als t u die d: C. eZastica Cerv. Gua­ distributed mainly in the Amazon basin and temala: McClay & Clara 30 (Uw 24222); Co­ the coastal regions of Venezuela and the Guia­ lombia, Choc6: Cuatrecasas 14218 (Uw 25161), nas. One species (H. tovarensis) occurs up to Cuatrecasas 15217(Uw25289);Surinam: Linde­ Costa Rica. Some species, originally described man 229 (Uw 23350). - C. tunu Hems!. Pana­ under Helicostylis, proved to belong to the ma: Pittier s.n. (US 715986, UW 7048); Ecua­ genus Brosimum. Helicostylis is a genus of trees, dor: Berg 424 (Uw 23615); Nicaragua: Engle­ up to 20-30 m tall, of the tropical lowland sing 97 (Uw 18808). - C. uZei Warb. Brazil, rainforest. Amazonas: Prance & Berg 19852 (Uw 20943); M a t e r i als t u die d: H. eZegans (Macbr.) Brazil, Mato Grosso: Krukoff 1409 (Uw 18427); C.C. Berg. Brazil, Mato Grosso: Prance & Berg Brazil, Rio Purus: Krukoff 5766 (Uw 18807); 18613 (Uw 20924), Prance & Berg 19868 (Uw Peru: Ellenberg 2949 (Uw 8827), Williams 1802 20944); Brazil, Amazonas: Krukoff 6809 (Uw (Uw 18433). 7946),8075 (Uw 16159),8521 (Uw 16201).­ Gen e ra I fe a t ure s: Growth rings fain t H. peduncuZata R. Ben. Surinam: van Donselaar or absent; colour light yellowish brown, no de­ 2992 (Uw 11963), Lanjouw & Lindeman 421 marcation between sapwood and heartwood. (Uw 1224), Lindeman 3544 (Uw 2315); French Texture coarse, grain straight. Specific gravity Guiana: BAFOG 165 M (Uw 5244),267 M (Uw 310-480 N per cubic metre. 5342); Brazil, Amapä: Pires 51670 (Uw 8956). Mi c r 0 s cop i c fe a tu res: Vessels diffuse, - H. scabra (Macbr.) C.C. Berg. Brazil, Amazo­ solitary (30-70%) and in short radial multiples nas: Krukoff 6811 (Uw 7948), 7081 (Uw or irregular clusters of 2-6; 4-8 per sq.mm, 8162), Prance & Berg 17967 (Uw 20953). - H. round to oval, diameter 135-215 .um, vessel tomentosa (P. & E.) Rusby. Guyana: Fanshawe member length 350-560.um. Perforations sim­ 148, For. Dept. 2757 (Uw 985), Maguire et al. ple, end walls transverse. Intervascular pits al­ 45735 (Uw 16775); Surinam: Lindeman 4756 ternate, round, oval to polygonal, 8-11 J..lm. (Uw 3276),5356 (Uw 3711),5357 (Uw 3712), Vessel-ray and vessel-parenchyma pits larger 6712 (Uw 4539), LBB (Maas) 10963 (Uw and irregularly shaped, half-bordered. Thin­ 11698); Brazil, Mato Grosso: Krukoff 1557 walled tyloses occasionally present. Fibres all (Uw 19400), 8859 (Uw 16256), Prance & Maas

, 'Procumbent' refers to very strongly procumbent cells with a length : height ratio of over 3; 'weakly procumbent' is used for length : height ratios of 2-3 : I of at least part of the procum­ ben t ray cells.

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15027 (Uw 19122); Brazil, Amapa: Pires 51834 Maquira Aublet (Fig. 7) (Uw 9077). - H. tovarensis (Klotzsch & Kar­ This Neotropical genus was revised by Berg sten) C.C. Berg. Colombia, Choc6: Cuatrecasas in 1972. Species of this genus were formerly 14424 (Uw 24955). - H. turbinata C.C. Berg. assigned to i.a. Olmedia and Perebea. Berg dis­ Brazil, Amazonas: Krukoff 8832 (Uw 16258), tinguishes five species, all being medium-sized type. or small trees of the tropical rainforest, up to General features: Growth rings absent; 20-50 m tall, distributed in northern South sapwood golden yellowish brown, heartwood America with one species up to Nicaragua. light to dark brown, sometimes with dark brown Material studied: M. calophy/la (P. & streaks on the tangential surfaces. The demarea­ E.) C.C. Berg. Brazil, Amazonas: Krukoff 6347 ti on between sapwood and heartwood general­ (Uw 7655),6655 (Uw 7851),8580 (Uw 16208), ly irregular. Texture moderately fine, grain 8642 (Uw 16210). - M. coriacea (Karst.)C.c. straight. Specific gravity 590-1070 N per cu­ Berg. Brazil, Amazonas: INPA-Manaus X-3885 bic metre. (Uw 18851), Krukoff 6615 (Uw 7817), 6705 M i e r 0 s e 0 p i c fe a t ure s: Vesse1s diffuse (Uw 7886); Brazil, Amapa: Pires 51743 (Uw but with a tendeney towards a diagonal arrange­ 9012); Venezuela: Breteler 3726 (Uw 11746). ment; mostly solitary (70-90%) and in short - M. costaricana (Standl.)C.C. Berg. Colombia, radial multiples of 2-4; 5-12 per sq.mm, round Choe6: Cuatrecasas 14135 (Uw 25382). - M. to oval, diameter 90-200 JJ.m, vessel member guianensis Aublet. Surinam: Lanjouw & Linde­ length 400-550 JJ.m. Perforations simple, end man 2187 (Uw 1644),2811 (Uw 1936), LBB walls almost transverse; intervaseular pits alter­ (Maas) 10795 (Uw 11210); Schulz 8980 (Uw nate, round, oval or polygonal, 8-10 JJ.ID. Ves­ 8855); Stahel 222 (Uw 222); Brazil, Amazonas: sel-ray and vessel-parenchyma pits larger and ir­ Pires 51803 (Uw 9634), Prance & Berg 18609 regularly shaped, half-bordered. Thin-walled (Uw 20921); Brazil, Amapa: Pires, Rodrigues tyloses often present. Fibres all or partly sep­ & Irwin 51612 (Uw 8949). - M. sc/erophylla tate, in some specimens non-septate; with small (Ducke) C.C. Berg. Surinam: Lindeman 5040 simple pits restricted to the radial walls; walls (Uw 3424); Brazil, Para: Krukoff 1133 (Uw 2-5 JJ.ID, lumina 7-14 JJ.m; in some sampIes 19224). partly gelatinous; length 1140-1585 JJ.m; F/V­ General features: Growth rings faint ratio 2.4-2.9. Rays uniseriate and multiseriate, or absent; colour (light) brown, no demarca­ 4-7 per mm. Uniseriate rays 0-20%, generally tions between heartwood and sapwood. Tex­ composed of few rows of procumbent cells and ture medium, grain slightly interlocked. Speci­ few to many rows ofupright cells, height up to fic gravity 610-1000 N per cubic metre. 8 eells (150-350 JJ.m). Multiseriate rays entire­ Mi c r 0 s cop i e fe a t ure s: Vessels diffuse Iy composed of (weakly) proeumbent eells' with a slight tendency towards a diagonal ar­ except for the uniseriate margins of 1-2 (5) rangement, solitary (40-90%) and in short ra­ rows of square and/or upright cells and few dial multiples of 2-4, 5-20 per sq.mm, round sheath cells in some of the rays; 2-5 (6) cells to oval, diameter 90-160 JJ.ID, vessel mem ber wide, up to 310-900 JJ.m high. Parenchyma pa­ length 400-640 JJ.m. Perforations simple, end ratracheal, vasieentric aliform with narrow to walls almost transverse. Intervascular pits alter­ broad wings; sometimes tending towards uni­ nate, round, oval to polygonal, 7-9 JJ.m. Vessel­ lateral arrangement abaxially, sometimes con­ ray and vessel-parenchyma pits larger and irre­ fluent; sometimes terminal parenchyma pres­ gularly shaped, half-bordered. Thin-walled ty­ ent. Parenchyma strands of 4 (2 - 8) cells. loses often present. Fibres all or partly septate Rhombic erystals in variable amounts in rays with small simple pits restricted to the radial and axial parenchyma. Vitreous siliea present walls; walls 3-5 JJ.m, lumina 5-9 JJ.m, occasion­ oecasionally in the fibres, rarely also in the ally gelatinous; length 1015-1600 JJ.ID; F/V­ axial parenchyma. Radial latex tubes always ratio 2-3. Rays uniseriate and multiseriate, present, albeit sometimes sporadically. Selero­ 6-8 per mm. Uniseriate rays 0-1 0%, composed tie cells oeeasionally noticed in the rays and/or of upright and few procumbent cells, height up axial parenchyma. to 3-6 cells (150-300 JJ.m). Multiseriate rays Not e: H. turbinata deviates in its F /V­ composed of procumbent cens' except for the ratio (3.9), due to the relatively long fibres uniseriate margins of 1-3 rows and square (1585 JJ.m) and short vessel members (400 JJ.m). and/or upright cens and few sheath cens; 3-5 The parenchyma is often confluent and locally cells wide, up to 300-1200 JJ.m high. Parenchy­ forms long parenchyma bands, up to 7-8 cells ma paratracheal, variable: wide to narrow vasi­ wide. Furthermore, the multiseriate rays are eentric, aliform with short wings to confluent frequently up to 7 cells wide and no diagnonal vessel arrangement was found. (text continued on page 191)

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Fig. 1. AntÜlris toxicarÜl, Uw 24430. - Fig. 2. Ibid., Uw 18780. - Fig. 3. , Uw 25289. - Fig. 4. C. ulei, Uw 20943. - All x 360.

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Fig. 5. Perebeaxanthochyma, Uw 18431. - Fig. 6. P. guianensis, Uw 16183. - Fig. 7. Maquira coriacea, Uw 9012. - Fig. 8. Pseudolmedia laevis, Uw 11817. - All x 360.

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Fig. 9. Naucleopsis uZei, Uw 18428. - Fig. 10-12. N. imitans, Uw 19991. - All x 360.

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Fig. 13. Helicostylis turbinata, Uw 16258. - Fig. 14-15. Mesogyne insignis, Uw 25797. - Fig. 16. Ibid., Uw 25686. - All x 360.

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and in narrow bands, sometimes wavy; strands Naucleopsis Miquel (Figs. 9-12) of 3-4 (2-8) cells. Radial latex tubes generally In this Neotropical genus Berg (1972) com­ present, though sometimes very in frequent. bined two Perebea species with the species for­ Rhom bic crystals occasionally present in the merly assigned to Acanthosphaera, Ogcodeia rays and rarely in the axial parenchyma. Vitre­ and Palmolmedia. The genus consists of small ous silica sometimes present in the axial paren­ to medium-sized trees of the tropical lowland chyma, rarely also in the vessels and fibres. and is distributed from Honduras to Rio de Not e: Wood anatomically, the individual J aneiro (Brazil). species show only part ofthe variation indicated M a t e r i als t u die d : N. amara Ducke. Bra­ above. As a genus, however, Maquira is very zil, Mato Grosso: Krukoff 1406 (Uw 19340). - heterogeneous in, at least, the parenchyma pat­ N. concinna (Stand!.) e.e. Berg. Brazil, Ama­ tern, the ray structure, the presence and fre­ zonas. Krukoff 6620 (Uw 7822), 6687 (Uw quency of crystals and silica, and the diameter 7875). - N. glabra Spruce ex Baillon. Brazil, and density of the vessels. The wood structure Acre: Krukoff 5323 (Uw 19899); Brazil, Ama­ shows an overlap with that found in Helicosty­ zonas: Berg et a!. 754 (Uw 24080), Krukoff lis, Perebea, and Pseudolmedia. 1300 (Uw 19277), 1392 (Uw 19331), 6198 (Uw 7549), 8271 (Uw 16170), Prance & Berg Mesogyne Engler (Figs. 14 -16) 18241 (Uw 20901),19805 (Uw 20938); Brazil, An African genus, monotypic according to Mato Grosso: Krukoff 1347 (Uw 19304). - N. Berg (I 977b). It consists of shrubs or trees, up guianensis (Mildbr.) e.e. Berg. Surinam: van to 15 (occasionally 40) m tall, of the lower Donselaar 3791 (Uw 12167), Lindeman 3790 stories of the tropical rainforest, between 500 (Uw 2776). - N. imitans (Ducke) e.e. Berg. and 1300 m altitude, occurring in eastern Tan­ Brazil, Acre: Krukoff 5485 (Uw 19991). - N. zania and Sao Tome. inaequalis (Ducke) c.e. Berg. Brazil, Mato Gros­ M a t e r i als t u die d: M. insignis Engler. so: Krukoff 1458 (Uw 19364). - N. krukovii Tanzania: Holst 2290 (Uw 25797), Scheffer 22 (Stand!.) e.e. Berg. Brazil, Amazonas: Krukoff (Uw 25795), Schlieben 158 (Uw 25686). 4544 (Uw 19524). - N. macrophylla Miq. Bra­ General features: Growthringsabsent; zil, Amazonas: Ducke 232 (Uw 2667), Maguire colour light brown, no demarcation between et a!. 56684 (Uw 16478), Prance & Berg 18543 heartwood and sapwood. Texture fine, grain in­ (Uw 20920). - N. mello-barettoi (Stand!.) e.e. terlocked. Specific gravity 700-750 N per cubic Berg. Brazil, Amazonas: Krukoff 6839 (Uw metre. 7974). -N. stipularis Ducke. Brazil, Amazonas: M i c r 0 s cop i c fe a t ure s: Vessels diffuse, Ducke 370 (Uw 18423). - N. ternstroemiflora solitary (60-80%) and in short radial multiples (Mildbr.) C. e. Berg. Brazil, Amazonas: Krukoff of 2-4, 13-20 per sq.mm, weakly angular (to 1306 (Uw 19281). - N. ulei (Warb.) Ducke. round), diameter 85-105 J.lm, vessel member Brazil, Amazonas: Krukoff 1301 (Uw 19278); length 375-505 J.lill. Perforations simple, end Peru: Williams 5194 (Uw 18428). walls alm ost transverse. Intervascular pits alter­ General features: Growth rings faint nate, round or polygonal, 7-10 J.lill. Vessel-ray or absent; colour light brown, no demarcation and vessel-parenchyma pits larger and irregular­ between heartwood and sapwood. Texture fine Iy shaped, half-bordered, apertures sometimes to medium; grain straight. Specific gravity be­ coalescent. Thin-waJled tyloses present. Fibres tween 570 and 990 N per cubic metre. septate with simple pits restricted to the radial Mi c r 0 s cop i c fe a t ure s: Vessels diffuse walls; walls 2-4 J.lm, lumina 9-12 J.lill; occa­ with a tendency towards a diagonal arrange­ sionally gelatinous; length 1125-1280 J.lm; ment, solitary (20-80%) and in short radial F/V-ratio 2.4-3.4. Rays uniseriate and multi­ multiples of 2-4; 20-45 per sq.mm, angular, seriate, 8-11 per mm. Uniseriate rays 15-45 round or oval, diameter 60-110 !Lm, vessel %, composed of few rows of weakly procum­ member length 355-535 J.lm. Perforations sim­ bent cells 2 and many rows of upright cells, ple, end walls alm ost transverse. Intervascular height up to 11 cells (500 J.lm). Multiseriate pits alternate, round, oval to polygonal, 6-9 J.lm. rays composed of (weakly) procumbent ceJls Vessel-ray and vessel-parenchyma pits larger except for the uniseriate margins of 1-6 rows and irregularly shaped, half-bordered. Thin­ of square and upright cells and few sheath cells walled tyloses occasionally present. Fibres sep­ in some rays; 2-4 cells wide, up to 700 J.lm tate with small simple pits restricted to the ra­ high. Parenchyma abundant; paratracheal long dial walls; walls 2-4 J.lm, lumina 5-10 J.lm; oc­ aliform-confluent to banded, and apotracheal casionally gelatinous; length 1000-1250 J.lm; in some diffuse strands. Parenchyma strands of F/V-ratio 2.1-2.9. Rays uniseriate and multi­ 4 (1-9) cells. Vitreous silica scarce, in fibres seriate, 6-8 per mm. Uniseriate rays 0-15%, and axial parenchyma. composed of procumbent 2 and/or square to

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upright cells, height up to 4-7 cells (200-350 length 420-550 J..lm. Perforations simple, end J..lm). Multiseriate rays composed of (weakly) walls alm ost transverse. Intervascular pits alter­ procumbent cells except for the uniseriate mar­ nate, round, oval or polygonal, 7-9 J..lm. Vessel­ gins of 1-6 rows of square and/or upright cells ray and vessel-parenchyma pits larger and irre­ and few sheath cells; 2-4 (5) cells wide, and up gularly shaped, half-bordered. Thin-walled tylo­ to 600-1500 J..lm high, sometimes vertically ses occasionally present. Fibres septate with fused. Parenchyma paratracheal, vasicentric ali­ sm all simple pits restricted to the radial walls; form, occasionally locally confluent, the vasi­ walls 2-4 J..lm, lumina 7-18 J..lm; occasionally centric ring often incomplete and lacking on gelatinous; length 1050-1600 J..lm; F/V-ratio the adaxial side; strands of 2-4 (1-8) cells. 1.9-3.4 J..lm. Rays uniseriate and multiseriate, Latex tubes present, generally with a small dia­ 5-7 per mm. Uniseriate rays 0-17%, composed meter in comparison to the surrounding ray of procumbent1 and upright cells, height up to cells. Rhombic crystals present in the ray cells 5-7 cells (150-340 J..lm). Multiseriate rays com­ of about 50% of the sam pies, rarely also in the posed of procumbent cells except for the uni­ parenchyma tissue. Vitreous silica occasionally seriate margins of 1(-3) rows of square (up­ present in the fibres and parenchyma. right) cells and few sheath cells; 3-6 cells wide Not e : The sam pies of N. stipularis, N. ama­ and up to 500-1000 J..lm high. Parenchyma pa­ ra and N. ulei (Williams 5194) do not complete­ ratracheal in generally complete, narrow, ali­ Iy fit the generic description. N. amara deviates form vasicentric rings, often confluent in tan­ in the diameter (150 J..lm) and number (15 per gential or diagonal direction over a short dis­ sq.mm) of the vessels and in its fibre length tance, the sheaths usually widest on the abaxial (1450 J..lm). N. stipularis shows abundant paren­ side of the vessels; strands of 4 (2-8) cells. Ra­ chyma in wavy paratracheal bands and ray mar­ dial latex tubes often present, some axial latex gins of up to 8 cells high. Both sampies of N. tubes present in one sam pie (cf. Topper & ulei differ from each other in the vessel dia­ Koek-Noorman, 1980). Rhombic crystals ab­ meter and ray height: the values are among the sent or scarce in ray cells; in one sampie abun­ lowest (Williams 5194) and highest (Kruko!f dant in rays and axial parenchyma.Yitreous silica 1301) values found in the genus. scarce in one sampie, in both vessels and fibres. Not e: One sam pie of P. guianensis (Kru­ Perebea Aublet (Figs. 5,6) koff 7209) deviates in its narrow-aliform paren­ Ever since this Neotropical genus was estab­ chyma, without confluent bands. The two sec­ lished, the generic delimitation has been the tions of Perebea, as defined by Berg, cannot be subject of discussion. In his monograph, Berg distinguished by their wood anatomy in the (1972) distinguished eight species in two sec­ available sampies. tions. The genus consists of small to medium sized trees or shrubs. Representatives are found Pseudolmedia Tnicul (Fig. 8) in tropical lowland forest in tropical South Berg revised the genus and reduced the num­ America and Panama. ber of species to nine. Some species were trans­ Material studied: P. guianensis Aublet ferred to other genera (Berg, 1972). Pseudol­ ssp. guianensis. Brazil: Irwin 48718 (Uw 16989), media occurs in the northern part of South Krukoff 8350 (Uw 16183); Brazil, Amazonas: America, Central America and the Greater An­ Krukoff 7041 (Uw 8128), Krukoff 7209 (Uw tilles, as sm all or medium-sized to large trees 01' 8241). - P. mollis (P. & E.) Huber ssp. mollis. various habitats. Brazil: Ducke 175 (Uw 2668); Brazil, Amazo­ M a t e r i als t u die d: P. laevigata Trecul. nas: Krukoff 1289 (Uw 19268), 6985 (Uw Guyana: A.C. Smith 3633 (Uw 21706); Brazil, 8080). - P mollis ssp. rubra (Trecul) C.C. Berg. Rio Solimoes: Ducke 359 (SJRw 40090, Uw Surinam: Stahel 211 (Uw 211). - P. xantho­ 2669); Brazil, Amazonas: Krukoff 6697 (Uw chyma Karsten. Peru: Williams 3412 (Uw 7880), 8383 (Uw 16186), Prance & Berg 18611 18431). (Uw 20923); Colombia: J. de Bruijn 1543 (Uw Gen e r a I fe a t ure s: Growth rings faint 14493). - P. laevis (R. & P.) Macbr. Guyana: or absent; colour (light) brown, no demarca­ Fanshawe 5378 (Uw 988); Brazil, Amazonas: tion between heartwood and sapwood. Texture Krukoff 5188 (Uw 19815); Brazil, Acre: Kru­ medium; grain straight. Specific gravity 530- koff 5217 (Uw 19823), 5312 (Uw 19889); 830 N per cubic metre. Venezuela: Breteler 3929 (Uw 11817). - P. Mi c r 0 s cop i c fe a t ure s: Vessels diffuse macrophylla Trecul. Brazil, Amazonas: Prance with a tendency towards a diagonal arrange­ & Berg 18376 (Uw 20906); Bolivia: Krukoff ment, solitary (40-80%) and in short radial 10738 (SJRw 39639, Uw 2670). - P. oxyphyl­ multiples of 2- 4; 5-15 per sq.mm, round to laria Donn.Smith. Mexico: I.N.I.F. XI52 (Uw oval, diameter 95-160 J..lm, vessel member 20653); Belize: Stevenson 119 (SJRw 14901).

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~ P. rigida (Klotzseh & Karsten) Cuatreeasas. rattosyce, formerly inciuded in the tribe Olme­ Ecuador: Maas, Berg & ter Welle 2918 (Uw dieae (Corner, 1962) were excluded by Berg 23575). ~ P. spuria (Swartz) Griseb. Guatema­ and transferred to the Moreae. la: McClay & Clara 31 (Uw 18027); Belize: The eigh t genera of the tribe Castilleae are Brown 35 (SJRw 15351). similar in many features and show considerable Gen e r a I fe a tu res: Growth rings absent; overlap, the differences being in quantitative colour (light) brown, no demarcation between features only. As far as we ean judge now, ehar­ heartwood and sapwood. Texture fine to me­ acters like vessel mem ber and fibre length, pit dium; grain straight or slightly interlocked. size and arrangement, ray strueture, and oecur­ Specific gravity 71 0~940 N per cubic metre. renee of crystals are of no ar very little diag­ Mi c r 0 s cop i c fe a t ure s: Vessels diffuse, nostic or taxonomie value in this group. In solitary (30~80%) and in short radial multiples Table I the characters with at least some diag­ of 2~4; 8~25 per sq.mm, round to oval, dia­ nostic value are listed: frequency and diameter meter 70~165 fJ.m, vessel member length 350~ of the pores, the (tendency towards a) diagonal 540 fJ.m. Perforations simple, end walls alm ost vessel arrangement, the presence of either transverse; intervascular pits alternate, round, weakly or strongly procumbent ray cells' , the oval to polygonal, 7 ~ I 0 fJ.m. Vessel-ray and parenchyma distribution, the presenee or ab­ vessel-parenchyma pits larger and irregularly sence of radial latex tubes, and the specific shaped, half-bordered. Thin-walled tyloses of­ gravity. For these characters, in particular far ten present, sclerotic in some sam pies of P. diagonal vessel arrangement and presence of laevigata. Fibres all or partly septate, with small latex tubes, a careful examination of large areas simple pits restricted to the radial walls; walls is required. Notwithstanding the high degree of 3~4 fJ.m, lumina4~9 fJ.ffi; sometimes gelatinous; similarity and overlap, some genera ean be length 1000~ 1600 fJ.m; F/V-ratio 2.4~3.4. recognised by a combination of features. Rays uniseriate and multiseriate, 5~8 per mm. The sampies of Antiaris studied are all very Uniseriate rays O~ 15 %, composed of procum­ similar. Only one sampie is remarkable in the bent and square, rarely upright cells, height up abundance of crystals. There are no wood ana­ to 3~ 7 cells (l50~600 fJ.m). Multiseriate rays tomical features which contradict the combina­ 2 composed of proeumbent cells , with or with­ tion of all species into one species, as Berg out I ~2 rows of square (rarely upright) margin­ (I 977b) proposed. The genus can be recognised al cells; 3~5 cells wide and up to 3 70~ I 000 fJ.m by its very low specific gravity, a low number high. Parenchyma paratracheal, confluent to of relatively large pores per sq.mm, and paren­ wavy banded, occasionally aliform with short chyma only in aliform (confluent) arrangement. to long wings; strands of 3~6 (11) cells. Radial These features are also found in other Castil­ latex tubes are present, but infrequent and in­ leae, but not in this combination. The wood conspicuous. Axial latex tubes present in one anatomical features thus do not contradiet in­ sampie of P. spuria. Rhombic crystals present clusion of Antiaris in this tribe. Corner's sug­ in marginal ray cells and in axial parenchyma gestion (1962) that Pseudolmedia might be one cells. Vitreous silica in most sam pies in vessels, of the nearest allies of Antiaris, is not confirm­ fibres or axial parenchyma. ed, neither is his idea, that Mesogyne and Anti­ Not e : P. spuria shows rhombic crystals in aris migh t be congeneric (see below). proeumbent as weil as square and upright ray Castilla shares the low specific gravity (al­ cells. ready mentioned by Mennega and Lanzing­ Vinkenborg, 1977) and the low number of relatively large vessels with Antiaris, but differs Discussion in its more abundant parenchyma, partly ar­ In his monograph of the tribe Olmedieae, ranged in bands, comparable with those found Berg (1972) presented a detailed survey of the in Mesogyne and Maquira p.p. Like in Meso­ taxonomie history of the tribe. In a subsequent gyne and Pseudolmedia, no trace of diagonal paper, Berg excluded Olmedia from the tribe vessel arrangement was found. and renamed it Castilleae (1977a). As already Mesogyne and Nauc/eopsis both show rela­ indicated in the Introduction, Mennega and tively narrow vessels. At least part of the pro­ Lanzing-Vinkenborg (1977) thoroughly studied cumbent ray cells are weakly procumbent' . the six Neotropieal genera now assigned to the Mesogyne is characterised by abundant paren­ tribe Castilleae. F or a detailed discussion of the chyma and the highest number of rays per mm; wood anatomy of these taxa we refer to their latex tubes are lacking. A remarkable feature in paper. Berg (1977a) also included two Afriean most specimens of Nauc/eopsis studied is the genera, Antiaris and Mesogyne, in the Castilleae. unilateral arrangement of the paratracheal pa­ The Palaeotropical genera Antiaropsis and Spa- renchyma. This feature was not found in N sti-

Downloaded from Brill.com09/26/2021 01:46:11AM via free access Table I. Some wood anatomical features of the Castilleae. -\0 .j>.

Vessels Fibres Rays Parenchyrna

... :::l ,-.. ;:: e ;:: [ :::l '"u '" Ei Ei u ;:: '"Ei g- '"Ei ;§, '" ~ e ~ Ei 00 ~ ~ ...... ~ '" ,., '" [ '"gp .s Ei ... 0:: 0:: ~ e! e! '"0 .€ -e.'" [ '"~ u '" 00 '"t:: ;§, '"t:: Ei'" B -e. ~ '"~ ,., '" ,., u u ... g .~ .~ Ei'" ep.. :ä '" '" 'i::~ u ;:: ~ 0:: ..... '" ... "2 '" .~ a B ,., u 0 0 ... ~ "'" .~ "'-~ '" ~ I;:: .~'" 0 32 .~ -" '" I;:: ,., '" "g. '"Ei '"'"00 .;:: u '" >< Ei '" er '"0:: 0:: := Tl .-=: '"Ei ~ '" ~ "'" B 0 p.. '" '" 0 *0::

Castilla 4-8 135-215 8-10 - 12-25 < 25 1-3 - 3-6 4-7 + - (+) - r - 310-480

Helicostylis 5-12 90-200 8-10 (+) 7-14 < 20 1-2 (5) + 2-5 4-7 (+) -- (+) r (p, u, a) 590-1070

Maquira 5-20 90-160 7-9 + 5-9 <10 1-3 - 3-5 6-8 (+) (+) (+) (+) r (p, u, a) 610-1000

Mesogyne 13-20 85-105 7-10 - 9-12 15-45 1-6 + 2-4 8-11 + (+) --- - 700-750

Naucleopsis 20-45 60-110 6-9 + 5-10 < 15 1-6 + 2-4 6-8 (+) -- + r (p, u, a) 570-990 Downloaded fromBrill.com09/26/2021 01:46:11AM ~ Perebea 5-15 95-160 7-9 + 7-18 < 17 1 (-3) - 3-6 5-7 + + -- r (ax) (p, u, a) 530-830 t:O E.. Pseudolmedia 8-25 70-165 7-10 - 4-9 < 15 1-2 - 3-5 5-8 + + -- r (ax) u,a 710-940 "g. ::s !n +: present; -: absent; r: in radial direction; ax: in axial direction, u: in upright and/or square ray cells; p: in procurnbent ray cells; a: in axial parenchyrna cells; between brackets: occasionally present. ~ VI 2 : see footnote on page 185. ,-.. W For quantitative features the range of sampie averages is given. '-' via freeaccess \0 -- - 00 .j>. IAWA Bulletin n.s., Vo!. 5 (3),1984 195

pularis where the parenchyma is arranged in - 1977b. Revisions of African Moraceae (ex­ wavy irregular bands, including the vessels. c1uding , Ficus, Musanga and My­ Helicostylis, with the exception ofthe sampIe rianthus). Bull. Jard. Bot. Nat. Belg. 47: of H. turbinata, is characterised by its short­ 267-407. aliform parenchyma, occasionally tending to­ - 1983. Dispersal and distribution in the Urti­ wards a unilateral arrangement, and the pres­ cales - an outIine. In: Dispersal and distri­ ence of weakly procumbent ray cells. bution (ed. K. Kubitzki). Pub!. Comp. Paul As already indicated by Mennega andLanzing­ Parey, Hamburg, Berlin. Vinkenborg (1977), and as may appear after a Bonsen, K.J. & B.J.H. ter Welle. 1983. Compa­ glance at Table I, the remaining genera Maqui­ rative wood and leaf anatomy of the Cecro­ ra, Perebea and Pseudolmedia are extremely piaceae. Bull. Mus. natn. Hist. nat., Paris, difficult to separate on their wood structure. 4e ser., 5, sect. B, Adansonia no 2: 151-177. The absence of diagonal vessel arrangement in - & - 1984. Systematic wood anatomy and the sampies of Pseudolmedia alone can hardly affinities of the Urticaceae. Bot. Jahrb. (in be considered as a sufficiently reliable diagnos­ press). tic and distinguishing feature. Berg (1972) dis­ Corner, E.J.H. 1962. The c1assification ofMo­ tinguished sections in each Maquira, Perebea raceae. Gard. Bull. Sing. 19: 187-252. and Pseudolmedia. In the wood anatomy these JA WA Committee, 1964. Multilingual glossary sections cannot be recognised. As will be evi­ of terms used in wood anatomy. Konkordia, dent from the above, we are not able to suggest Winterthur. any other arrangement, in genera or subgenera, Janssonius, H.H. 1934. Micrographie des Holzes than the one already proposed by Berg. der auf Java vorkommenden Baumarten.VI. In conclusion, in the material described in Brill, Leiden. this paper, we find no reason to question the Koek-Noorman, J. &B.J.H. ter Welle. 1976. The delimitation of the tribe Castilleae sensu Berg. anatomy of branch abscission layers in The genera Antiaris, Castilla, Helicostylis, Me­ Perebea mollis and Naucleopsis guianensis sogyne, and Naucleopsis can be recognised. (Castilleae, Moraceae). In: Wood structure Mesogyne is the most deviating genus. How­ in biological and technological research ever, there are no features that are restricted to (eds. P. Baas, A.J. Bolton & D.M. CatIing): this genus only. All are shared by at least one 196-203. Leiden Bot. Sero 3. Leiden Univ. other genus. Press, The Hague. Maquira, Perebea andPseudolmedia are hard­ Mennega, A.M. W. & M.. Lanzing- Vinkenborg. ly separable and they are highly similar to Heli­ 1977. On the wood anatomy of the tribe costylis. 'Olmedieae' (Moraceae) and the position of the genus Olmedia R. & P. Acta Bot. Neer!. 26: 1-27. References Record, S. J. & R. W. Hess. 1940. American Berg, C. C. 1972. Olmedieae, Brosimeae (Mora­ woods of the family Moraceae. Trop. Woods ceae). Flora Neotropica. Monograph no 7. 61: li-54. Hafner Pub!. Comp., New York. Tippo, O. 1938. Comparative anatomy of the - 1973. Some remarks on the c1assification Moraceae and their allies. Bot. Gaz. 100: and delimitation of Moraceae. Meded. Bot. 1-99. Mus. & Herb., Utrecht: 386. Topper, S. M. C. & 1. Koek-Noorman. 1980. The - 1977a. The Castilleae, a tribe of the Mora­ occurrence ofaxial latex tubes in the sec­ ceae, renamed and redefined due to the ex­ ondary xylem of some species of Artocar­ c1usion of the type genus Olmedia from the pus J.R. & G. Forster (Moraceae). IAWA 'Olmedieae'. Acta Bot. Neer!. 26: 73-82. Bull. n.S. 1: 113-119.

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