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The Systematic Wood Anatomy of the Moraceae (Urticales) I. Tribe Castilleae 1

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The Systematic Wood Anatomy of the Moraceae (Urticales) I. Tribe Castilleae 1 IAWA Bulletin n.s., Vol. 5 (3),1984 183 THE SYSTEMATIC WOOD ANATOMY OF THE MORACEAE (URTICALES) I. TRIBE CASTILLEAE 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 taxonomy 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 Maquira, Perebea and Pseudolmedia. fonnation be used to contribute to our under­ Antiaris, Castilla, Helicostylis, Mesogyne and standing of patterns of relationship, and aid in Naucleopsis 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, plant 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, Brosimum, Dor­ genera or commercially important timbers. stenia, Helianthostylis, Trilepisium, Tryma­ Numerous accounts in the literature were com­ tococcus; of Scyphosyce and Vtsetela no pared with our findings. Here we only refer to material is available); literature inc1uding elaborate wood anatomical III. Ficeae (Ficus); 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 (Broussonetia, Cal­ Janssonius, 1934; Koek-Noonnan & ter Welle, pidochlamys, Cardiogyne, Chlorophora, 1976; Record & Hess, 1940; Tippo, 1938. Maclura s.1. incl. Plecospermum and Cudra- 1 This project was made possible by a grant of BION-ZWO (14.45-01). Downloaded from Brill.com09/26/2021 01:46:11AM via free access 184 IAWA Bulletin n.s., Vol. 5 (3),1984 nia, Maillardia, Malaisia, Militia, Morus, Pa­ Finally, the specific gravity was established chytrophe, Streblus s.I., Trophis 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 (Antiaropsis, Arto­ during at least several weeks. It is the general carpus, Bagassa, Batocarpus, Clarisia, Parar­ experience that this results in a relative mois­ tocarpus, Poulsenia, Prainea, Sorocea, Spa­ ture content of approx. 12%. rattosyce, Treculia; 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 species, described since the genus 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.
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