lAWA Bulletin n.s., Vol. 7 (2), 1986 91 The systematic wood anatomy of the Moraceae (Urticales) IV. Genera of the tribe Moreae with urticaceous stamens* by B.J.H. ter Welle J. Koek-Noorman and S.M.C. Topper Institute of Systematic Botany, University of Utrecht, Heidelberglaan2, 3508 TC Utrecht, The Netherlands Summary The wood anatomy of the genera of the tribe Berg (1983) is followed here. He recognised 4 Moreae with urticaceous stamens, viz. Brousso- tribes: Dorstenieae, Castilleae, Moreae and Fi- Madura s.l. Chlo- The Moreae all netia, (including Cardiogyne, ceae. sensu Berg comprise genera rophora, and Cudrania),Malaisia,Milicia,Morus, of the tribes Moreae and Artocarpeae sensu Olmedia, Pachytrophe, Plecospermum, Sloeti- Comer (1962), and also a few genera assigned Streblus to Olmedieae opsis, s.l. (includingParatrophis, Phyl- the tribe sensu Comer (1962). lochlamys, Pseudostreblus, and Sloetia), and This large tribe, comprising about 55% of all Trophis s.l.(includingCalpidochlamys and Mail- genera of the family, is subdivided into two is described in detail. of 9 and 12 The lardia), Separate descrip- groups genera, respectively. tions have been made for sections and/or sub- most important difference between the two genera to facilitate the discussion about the groups of the Moreae sensu Berg is found in the generic delimitations made by several taxono- stamens: presence or absence of ‘urticaceous’ mists. The combinations inflexed which flower- following generic pre- stamens, i.e., stamens on viously proposed by taxonomists are supported ing spring back elastically (Berg, 1973). The by wood anatomical features: Broussonetia + group of genera with urticaceous stamens, treat- Allaeanthus: Maclura + Chlorophora + Cardio- ed here, were described by Berg (1983) as + Cudrania + The inflorescences gyne Plecospermum. segrega- having relatively simple; macro- tion of the African of in species Chlorophora a spermy to microspermy; diaspores dry, drupa- separate genus Milicia is supported by wood ceous or surrounded by a fleshy perianth,part- anatomical evidence. The broad genus concept ly zoochorous; ecology predominantlymarginal of Streblus and Trophis is not supported by to rainforest conditions;taxonomically and geo- wood anatomy. Several sections ofthese genera graphically more or less coherent; centered in should be reinstated as genera. The correlations the Indo-Malaysian region, with transpacific between wood anatomy, latitude, habit and connections. habitat are discussed as far as allowed by the In Berg’s opinion this subdivision of the material studied. Characters useful for the de- Moreae is very similar to groups recognised in limitation of the be the size the older classifications of the Moraceae. How- genera proved to of the intervascular pits, the parenchyma distri- ever, both Bentham and Hooker (1880) as well bution, and the lengths of fibres and vessel ele- as Engler (1888) recognised only two tribes ments. Rhombic crystals, vitreous silica and (subfamilies) in the Moraceae, viz. Moreae (Mo- radial latex tubes usually are useful additional roideae) and Artocarpeae (Artocarpoideae)! characters. These two tribes consequently also include the Key words: Moraceae, Moreae, Urticales, sys- tribes Ficeae, Dorstenieae and Castilleae (or Ol- tematic wood anatomy medieae) as recognised by both Berg(1983) and Comer (1962). From the foregoing it will be evident, that Introduction the delimitation of the Moraceae tribes, and es- This paper is part of a series, in which the pecially of the group of the Moreae and Arto- wood anatomy of the Moraceae is described carpeae is extremely difficult. For practical rea- and discussed in relation the of the delimitation of the tribes and to taxonomy sons genera the family. The general outline of this study is as presented by Berg (1983) is followed here. provided in the first paper (Koek-Noorman et The group of genera treated in this paper is characterised the of urticaceous al., 1984). by presence The recent classification of the Moraceae by stamens. Included are the following genera: * This project was made possible by a grant of BION-ZWO (14.45-01), 92 lAWA Bulletin n.s., Vol. 7 (2), 1986 distribution genus including geographic Ampalis — Madagascar Bleekrodea —- Madagascar and Malesia Broussonetia — Broussonetia — Temperate and tropical Asia — Allaeanthus -— Madagascar and Asia - Fatoua — Madagascar and Asia MacluraMadura — MacluraMadura — Temperate and tropical America — Cardiogyne —- Southeast Africa — Chlorophora —- Tropical America — Cudrania —- Asia and Australia — Plecospermum -— India and Thailand Malaisia — Southeast Asia and Australia Milicia -— Tropical Africa MorusMoms -— Temperate northern hemisphere and Pantropical Pachytrophe — Madagascar Streblus — Streblus — Southeast Asia — — — Paratrophis Asia and Australia — Phyllochlamys — Asia — Pseudostreblus —- Asia, mainland — Sloetia — Malaya and Sumatra — Sloetiopsis -— Tropical Africa Trophis — Trophis —- Tropical America — Calpidochlamys — Asia — Maillardia — Madagascar and ReunionRéunion — Olmedia -— Tropical America Berg (1983) mainly followed the broad genus sel member length, fibre wall thickness and concepts of Comer (1962). However, he hesi- lumen diameter, fibre length and F/V-ratio to Maillardia tated accept the inclusion of and (i. e., fibre length/vessel member length), the Calpidochlamys in Trophis, and of Sloetia and number ofrays per mm. The other quantitative Sloetiopsis in Streblus (pers. comm.). On the data represent minimum and maximum values. other hand. Corner (1962) did not include Ol- The data on specific gravity are based on the media in Trophis, and Plecospermum in Maclu- samples cited under ‘Material studied’, comple- these the of The ra, but gave taxa rank genus. mented with data of samples in the Utrecht genus Milicia was reinstated by Berg (1982)to wood collection not further included in the accommodate the two African species of Chlo- anatomical study. rophora. Methods and Materials Generic descriptions The methods are described in the first paper of this series (Koek-Noorman et al., 1984).The Broussonetia L’Her. ex Vent. ray types mentioned in the descriptions, follow the definitions of Kribs (1968). The wood sam- Broussonetia sect, Allaeanthus (Thw.) Corner ples of the African genera are generally backed (Table 1; Figs. 1, 3) by herbarium vouchers identified by Dr, C.C. According to Corner (1962) this section com- Berg; this also applies to the samples of the prises three species, B. kurzii, B. luzonica, and B. Neotropical genus Olmedia. The samples of the zeylanica. Corner (1962) included Allaeanthus Asian backed herbarium in this genera are generally by Broussonetia, reducing taxon to a sec- vouchers, but the genus and species names have tion. The combination Broussonetia greveana not been checked recently. Details on individual was made by Berg(1977a). Originally, this taxon wood samples are provided at the beginningof was assigned to Ampalis by Baillon (1895) as In each generic description. total 114 samples A. greveana. However, Leandri (1948) trans- were studied. Wood samples of Ampalis, Bleek- ferred this taxon to Chlorophora. Corner (l.c.) rodea, and Fatoua were not available for this made the combination Madura greveana as a study. In the descriptions, averages are given species of the section Chlorophora. More re- for percentage of solitary vessels, numbers of cently, Capuron (1972) placed it in Allaean- vessels per sq.mm, diameter of the vessels, ves- thus, near A. zeylanica. Vol. 7 1986 lAWA Bulletin n.s., (2), 93 Fig. 1. Element length ofMorus and Broussonetia species. Broussonetia is restricted to Mada- well defined from the greveana paler cream sapwood. and the Comore Islands. The other Texture interlocked. gascar spe- coarse, grain slightly Spe- cies are distributed from Sri Lanka to Malesia. cific gravity 460—630 N per cubic metre. The trees (or liana, B. kurzii) of the Asian spe- Microscopic features: Vessels diffuse, in cies are found lowland rainforests, up to solitary (55-70%) and in short radial multiples 200—400 altitude. Broussonetia of round to m greveana 2—4; 5—9(—16) per sq.mm, oval, prefers dry forests or thickets or occurs along diameter 155—215 pm, vessel member length streams, from sea level up to 800 m altitude. 280—335 pm. Perforations simple, end walls al- Material studied: B. greveana (Baillon) most transverse. Intervascular pits alternate, C.C. C.T.F.T. 59 oval to 4—9 Berg. Madagascar: (Uw 26758), round, polygonal, pm. Vessel-ray via Wageningen no. 148 (Uw 26759), Somely and vessel-parenchymapits larger and irregularly (= Vory) s.n. (Uw 26760). - B. kurzii (Hook.f.) shaped, half-bordered, the borders sometimes Comer. Thailand: ex Lw(Uw26754). — B. luzo- reduced. Thin-walled tyloses present. Fibres nica (Blanco) Bureau. Philippines: A.N. Green nonseptate, with small simple pits restricted to F.P.R.I. 1165 (Uw 24210); Indonesia: AL the radial walls; walls 1—3 pm, lumina 8—16 22231 WDw 980—1130 (Uw 24432); locality unknown,ex pm, occasionally gelatinous; length 416/2531 (Uw 24611). pm; F/V-ratio 3.3-3.7. Rays uniseriate and General features: Growth rings absent multiseriate, heterogeneous type II/III, 3—5 or faint; heartwood light brown to brown, not per mm. Uniseriate rays 0—5%, composed of 94 lAW A Bulletin n.s., Vol. 7 (2), 1986 11 (p) a) (p) (p) dis- p a) c Oo 3 2 •3 u a u U.P a, a, a, u, u,a,(p) (u, u,a,(p) u,a,(p) U, u, u, vessel Crystals O,* vasicentric. (rh) v; 4>Cu >> rhrh rh rh rh rh rh lower. d,d, I 1 1 m much m - - - + + + (+)(+) paratracheal; b