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The Systematic Wood Anatomy of the Moraceae (Uriticales)

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The Systematic Wood Anatomy of the Moraceae (Uriticales) lAWA Bulletin Vol. 7 1986 n.s., (3), 175 The systematic wood anatomy of the Moraceae (Uriticales). V. Genera of the tribe Moreae without urticaceous stamens* by B.J.H. ter Welle J. Koek-Noorman and S.M.C. Topper Institute ofSystematic Botany, University of Utrecht, Heidelberglaan2, 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 Artocarpeae sensu Corner (1962). provided: Antiaropsis, Artocarpus, Bagassa, Ba- Both Berg’s and Corner’s subdivisions deviate tocarpus, Clarisia, Parartocarpus, Poulsenia, from older classifications, as given by, for in- Prainea, Sorocea, Sparattosyce, and Treculia. stance, Bentham and Hooker (1880) and Engler Wood anatomical variation below the genus (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 (New Guinea), Artocarpus (Southeast by the presence or absence of septate fibres, Asia), Bagassa (Neotropics), Batocarpus (Neo- and of radial latex tubes, the size of the inter- tropics), Clarisia (Ncotropics), Huliettia (South- vascular pits, the parenchyma distribution,and east Asia),Parartocarpus (Southeast Asia), 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 Africa). anatomy Methods and Materials Introduction The methods employed are those given in This is of in which the the first of this paper part a series, paper series (Koek-Noorman et wood anatomy of the Moraceae is described al., 1984). All wood samples studied are backed and discussed in relation to the taxonomy of by herbarium vouchers. Taxa of the African the The family. 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- In the fourth of the the al., 1984). paper series 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 described was (Ter Welle study because wood samples were not at our et al., 1986). In this paper the genera without disposal. this of stamens dealt with. In addition type are 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 be used to characterise this can group on other samplesofthe Utrecht wood collection of genera in Moreae according to Berg (1983); not further included in the anatomical study. inflorescences rather to simple 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. 1, 2) prises all genera of the tribes Artocarpeae and A genus with one or two species (Comer, Moreae sensu Comer (1962), and also a few 1962). The genus is probably restricted to Irian to genera assigned the tribe Olmedieae by Jaya (Indonesia). Its occurrence in Papua New Comer (l.c.). The subdivision in two groups Guinea is questioned by Comer (1962). The * This project was made possible by a grant of BION-ZWO (14.45-01). 176 IAWA Bulletin n.s., Vol. 7 (3), 1986 shrubs or small trees, up to 11 m high, are found 22031b (Uw 24421). - A. fretissii Teijsm. & in lowland forests 1000 usual- Binn. ex Hassk. Irian BW 9871 up to m altitude, Indonesia, Jaya: ly in the undergrowth. (Uw 18112), BW 2133 (Uw 24316), BW 2512 Material studied : A. decipiens Schu- (Uw 24317), BW 2175 (Uw 24318). A. hor- mann. Indonesia, Irian Jaya: CS1RO H 5198 ridus Jarrett. Indonesia, Irian Jaya; BW 11593 (Uw 25533). (Uw 20479). - A. integer (Thunb.) Merr, Indo- General features: 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 cubic kemando FMS 28853, ex FHOw per metre. Miq, Microscopic features: Vessels diffuse, 4536 (Uw 24608); Indonesia: RTIw IND. Col. solitary (50%) and in short radial multiples of 3590 (Uw 24605), Pfeiffer E755 (Uw 24324). to — lakoocha Roxb. Pakistan: MADw 24487 2—3; 9 per sq.mm, round slightly oval, diam- A. ex eter 108 pm, vessel member length 525 pm. (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); RTIw IND. — nitidus Trecul. allyround, 7—9 pm. Vessel-ray and vessel-paren- Col. 4027 (Uw 24606). A. 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, 7822 — A. mon, often with vitreous silica. Fibres septate, Irian Jaya: Fokkinga (Uw 24329). with small Jarrett. Sabah: Sandakan with several septa per fibre, simple sericicarpus Malaysia, pits restricted to the radial walls; walls 3-4pm, For. Dept. 50586 (Uw 24218). - A. teysmannii lumina 3—7pm; length 1600 pm; F/V-ratio 3.0. Miq. Indonesia, Irian Jaya: BW 1204 (Uw Rays uniseriate and multiseriate, heterogeneous 24330). Uniseriate General features; Growth type II/III, 7 per mm. rays 25%, rings absent; composed of upright cells only, but variable in heartwood light brown to dark brown, often to 12—24 cells distinct from the to height, heightup (480-800pm). sharply pale yellow light Multiseriate rays composed of procumbent cells brown sapwood. Texture coarse, grain(weakly) of 280—940 N except for the uniseriate margins 1—5, occa- interlocked. Specific gravity per 20 of sionally up to rows square and/or upright cubic metre. cells; 3—4 cells wide, up to 960—1400 pm high. Microscopic features: Vessels diffuse, Occasionally with some sheath cells. Parenchy- solitary (45—80%) and in short radial multiples concentric and clusters of 2—6 ma scanty paratracheal, and in irregular 2—4; per sq. mm, cells wide and Paren- round and diameter 160—310 vessel bands, 3-6 4-5 per mm. oval, pm, chyma strands of 4 cells. A few latex tubes ob- member length 350—630 pm. 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 pm. Vessel-ray and ves- The genus has been monographedby Jarrett sel-parenchyma pits larger, half-bordered, the (1959c,d,e; 1960a,b). About 50 species are borders sometimes reduced. Thin-walled tyloses The is distributed but absent. Fibres currently recognised. genus common, occasionally non- from India to southern China, Malesia and the septate, with small simple pits restricted to the altilis radial walls 2—4 lumina 7—25 Solomon Islands. Two species, A. and A. walls; pm, pm, 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 pm; F/V-ratio 2.3—3.9. Rays uni- in evergreen forests and areas with a mild mon- seriate and multiseriate, heterogeneous type from level 1000 3—6 Uniseriate soon climate, usually sea up to II/III, per mm. rays 3—30%, m altitude. composed of upright, square and sometimes Material studied: A. anisophyllus Miq. few procumbent cells, height up to 4—10 cells Malaysia, Sabah; Sandakan For. Dept. 50567 (160—480 pm). Multiseriate rays composed of (Uw 24213). — A. blancoi (Elmer) Merr. Philip- procumbent cells, except for the uniseriate pines, Quezon: J.P. Rojo 262 (Uw 24214). — margins of 1—2 (5) rows of square and/or up- A. communis J. R. &G. Forster. Indonesia, Irian right cells; 3—5 cells wide, up to 600—1200 pm 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 lAWA 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 Noorman, 1980). monograph of Mello Filho & Emmerich (1968). The is distributed from genus Costa Rica to Aublet to Bagassa (Figs. 7, 8) Brazil, Peru and Bolivia. Trees up 20—30 m In a treatment for the Flora of Suriname, 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 monotypic, as B. tiliifolia is now con- (Uw 19158); Krukoff 6422 (Uw 7714); Peru: as sidered a juvenileform ofB. guianensis (Berg, Williams 5334 (Uw 18416). — B. orinocensis The is distributed in the Karsten.
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