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Maquira, Perebea Following Questions 183 The systematic wood anatomy of the Moraceae (Urticales) I. Tribe Castilleaen by J. Koek-Noorman S.M.C. Topper and B.J.H. ter Welle Institute ofSystematic Botany, University ofUtrecht, Heidelberglaan 2, 3508 TC Utrecht, The Netherlands Summary The wood anatomy of the tribe Castilleae The main aim of the present research project sensu is described. Similarities differ- Berg and is to find an answer to the following questions: ences are discussed in relation to his What is the anatomical in concepts variability range taxa of the taxonomy of the tribe. The wood anato- of Moraceae as compared to closely related mical variation does not enable to distinguish taxa? — To what extent can the anatomical in- between Pseudolmedia. Maquira, Perebea and formation 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 tribal 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 conclusions was that the exclusion of Ol- 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 first present paper is the one of a series Antiaris and Mesogyne, also placed in the Cas- with the dealing systematic wood anatomy of tilleae by Berg (1973). Therefore we decided to Moraceae. It includes wood anatomical include the Castilleae in of the descrip- our survey tions of the of the tribe Castilleae. This genera wood anatomy of Moraceae. We intend to pre- project follows in logical the work of sent a series of in each of which the sequence papers, C.C. taxonomist of the who had Berg, family, genera of one tribe are described and their experienced problems in delimitingand defining wood anatomical similarities and differences the family as a whole, as well as the individual are discussed in relation to ideas about delimi- and tribes. The wood and leaf genera anatomy tation and relationships as brought forward by, of the Cecropiaceae, a segregate of the Mora- i.a.. Corner (1962, and his unpublished Flora in relation the of this Malesiana ceae, to taxonomy group account of 1972) and Berg (1972, has been discussed in Bonsen already a paper by 1977a, b, 1983). We will follow the most re- and Ter Welle (1983). Their study on the cent on sys- views ofBerg the tribal arrangement, as tematic of the indicated him anatomy Urticaceae, closely re- by in his paper on distribution lated to the Moraceae, is in the press (1984). and dispersal in the Urticales (1983). Most wood anatomical literature Mora- We intend on to publish separate papers on the ceae is rather difficult to taxonomi- tribes ofthe Moraceae the folio interpret in wing sequence; cally, due to the often dubious identifications I. Castilleae (Antiaris, Castilla, Helicostylis, and the nomenclature! frequent changes. Fur- Maquira, Mesogyne, Naucleopsis, Perebea, thermore, descriptions are nearly always based Pseudolmedia ); few and relate on samples only to individual II. Dorstenieae i Bosqueiopsis, Brosimum, Dor- genera or commercially important timbers. stenia, Helianthostylis, Trilepisium, Tryma- Numerous accounts in the literature were com- of tococcus; Scyphosyce and Utsetela no pared with our findings. Here we only refer to material is available); literature elaborate including wood anatomical III. Ficeae (Ficus ); descriptions based on identified mate- IV. Moreae characterised the reliably p.p., by presence rial, and not mentioned elsewhere in the text: of ‘urticaceous’ stamens (Broussonetia, Cal- Janssonius, 1934; Koek-Noorman & ter Welle, pidochlamys, Cardiogyne, Chlorophora, 1976; Record & 1938. Maclura Hess, 1940;Tippo, s.l. inch Plecospermum and Cudra- 1 This project was made possible by a grant ofBION-ZWO (14.45-01). 1984 184 lAWA Bulletin n.s., Vol. 5 (3), Pa- the established nia, Maillardia, Malaisia, Militia, Morus, Finally, specific gravity was chytrophe, Streblus s.l., Trophis s.l. incl. in the usual way. Measuring of the moisture mate- content of the considered unneces- Olmedia,i; of Ampalis and Fatouia no samples was rial is available); sary as all samples were stored in a room with the absence central and air of V. Moreae p.p., characterised by heating an humidity c. 40% of ‘urticaceous’ stamens ( Antiaropsis, Arto- during at least several weeks. It is the general Clarisia, Parar- experience that this results in a relative mois- carpus, Bagassa, Batocarpus, of tocarpus, Poulsenia, Prainea, Sorocea, Spa- ture content approx. 12%. Treculia; of Hullettia material rattosyce, no is available). intend to discuss In a concluding paper we Generic descriptions the taxonomic value of the various wood ana- tomical features the tribal level and to on com- Antiaris Lesch. (Figs. 1, 2) the individual tribes. pare 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 combined all Most wood samples studied are backed by these species. Berg (1977b) spe- herbarium vouchers which were identified by cies in Antiaris toxicaria Lesch., lowering the C.C. Berg, the taxonomist of the family. Most rank of these four species to subspecies. The to 40-60 samples were taken from the trunk of the tree; genus Antiaris (trees, up m tall, or sometimes, however, the origin was unknown. shrubs) occurs in the Palaeotropics in primary institu- and forests in wet and Many wood samples were provided by secondary dry habitats, all the world. The from level 1500 altitude. tional wood collections over sea up to m Utrecht wood collection served as a first basis Material studied: A. toxicaria Lesch. for this study. In the generic descriptions, only Uganda: B. T. Styles 201 (Uw 19390), For. 8-1950 Zaire: C. Donis413 those samples are cited of which sections were Dept. (Uw 18387): made. The general features of the wood are (Uw 24211), J. Louis 11201 (Uw 18780); West Africa: F.P.R.L. 209 A East Afri- often based upon the study of more wood (Uw 18386); samples not cited here. Sections and macera- ca: H.E. Desch-F.R.I.C.S. (Uw 24305); Malay- to standard tech- sia: For. 9-1954 tions were prepared according Dept. (Uw 18388); Indonesia, niques and embedded in Canada balsam and in Java: Koorders 8316t/127S7B (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.l. 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 including the walls and averages are based tween sapwood and heartwood. Texture coarse; Vessel is based interlocked. 270-530 N on 25 measurements. frequency grain Specific gravity 25 In the for both cubic on counts. descriptions, per metre. characters minimum and maximum averages Microscopic features: Vessels diffuse, found of each in towards a in samples genus are provided. some samples tending diagonal ar- in The percentage of solitary vessels was calculated rangement; solitary (45—65%) and short ra- after examiningan area with at least 100 spores. dial multiples or irregular clusters of 2—6; 3— 10 to diameter 145— Clusters and multiples were regarded as 2, 3, 4, per sq.mm, round oval, 425-575 etc. vessels, depending on the number ofvessels 230 pun, vessel member length pun. Vessel member and fibre Perforations end walls transverse. Inter- per group. length simple, length are based on at least 100 measurements vascular pits alternate,round, oval or polygonal, and minimum and maximum 9-13 Vessel-ray and vessel-parenchyma per sample, again, pun. half-bordered. sample averages found in the genera are pro- pits larger and irregularly shaped, vided. Additionally, the averages were used to Thin-walled tyloses occasionally present in calculate the ratio of fibre length/vessel mem- samples from Asia. Fibres septate with small ber length, in the descriptions referred to as simple pits restricted to the radial walls; walls wall thick- 1—3 lumina 10-30 often F/V-ratio. For the fibres, maximum pun, pun; gelatinous; and maximum lumen diameter 1030-1390 1.8-3.1. ness are given. length pun; F/V-ratio The percentage of uniseriate rays is based on Rays uniseriate and multiseriate, 4—7 per mm. Uniseriate of counts ofat least 100 rays. rays 2—15%, composed procum- 185 lAWA Bulletin n.s., Vol. 5 (3), 1984 bent and few with small restrict- square or upright cells, height up or partly septate simple pits 10 cells Multiseriate ed to the radial walls 1—3 lumina to (430 pun). rays com- walls; pun, 2 of cells uniseriate 15—25 posed procumbent , margins pun, occasionally gelatinous; length of 1—2 of cells 1000—1200 2.0—2.9. (4) rows square and/or upright pun; F/V-ratio Rays uni- and occasionally few sheath cells; 4—7 cells seriate and multiseriate,4—7 per mm. Uniseriate 1100 of wide and up to pun high. Parenchyma pa- rays 0—25%, composed procumbent and up- vasicentric-aliform with short 6—10 cells ratracheal, wings, right cells, height up to (150—500 occasionally confluent, parenchyma strands of pun).
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