lAWA Bulletin n.s., Vol. 7 (2), 1986 91

The systematic wood anatomy of the (Urticales)

IV. Genera of the tribe Moreae with urticaceous *

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,, of the tribes Moreae and 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: + group of genera with urticaceous stamens, treat-

Allaeanthus: + Chlorophora + Cardio- ed here, were described by Berg (1983) as

+ Cudrania + The gyne Plecospermum. segrega- having relatively simple; macro-

tion of the African of in species Chlorophora a spermy to microspermy; diaspores dry, drupa-

separate Milicia is supported by wood ceous or surrounded by a fleshy perianth,part-

anatomical evidence. The broad genus concept ly zoochorous; ecology predominantlymarginal

of and 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 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

- — 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 (+)(+) - (+)+(+)- - - - - diameter c + - + - - - - scanty

a + a + - + - - (+) (+) Parenchyma distribution vessel sp: Vv + +_(+)--+ +-(+)------— - - and P + + + + + + initial); sp i S cells. I higher or Broussonetia. 11

|_g Moo much o -s JU 1100 ? terminal of Fibres 980-1130980-1130 690-750 570-770 775-930 695-1150 parenchyma I1 (i.e., species 11 frequency axial aQÛ |.3

a: vessel marginal £a c 'E-"H, f/■■S 'r' •Ü 7—97-9 6 4-7 7-9 6-11 9-13 9-13 cells; individual m: fa|| ray of latewood

So c « E E 1 « S £ ? I -sxs X) ? confluent; 305—325305-325 280-335280-335 245-265 230-245 240-275 295-315 the in c: procumbent characters *: bands; in I1 p: Vessels o .s.5 Ss c 3 CTO' a *d: 6—9 16 5-6 poorly 6—9 5-6 8-10*8-10* 8-20*8-20* 7-9*1-9* 4-5*4-5» upright wood or aliform;

(ri) a; and = o Some o•a 3£’£ *3 di di di ri ri ri present di,

I 1. _ — ? _—+ --_ 1100 ___- + ----+ square parenchyma: in Table occasionally u: Indonesia Indonesia Europe (): crystals; - ring-porous,- - Origin Madagascar Thailand Philippines, Japan Japan Japan, Thailand, absent; rhombic -: ri: rh: diffuse; present; druses; di: Allaeanthus Broussonetia +: greveana kurzii luzonicaluzonica kaempferikaempferi kazinoki papyrifera papyrifera d: Species kurzii B.kazinoki Section B. B. B. Section B. B. B. B. Legends; tribution: crystals: I AWA Bulletin n.s., Vol. 7 (2), 1986 95

and 12 Fibres procumbent upright cells, heightup to common. nonseptate, with small simple

cells (120-185/am). Multiseriate rays composed pits restricted to the radial walls; walls 1—3

of procumbent cells except for the uniseriate pm, lumina 5-20 pm, occasionally gelatinous;

margins of 1-2 rows of square and/or upright length 570-1150 pm; F/V-ratio 2.5-3.8. Rays

to cells; 2—6 cells wide, up 600(—1300) pm uniseriate and multiseriate, heterogeneous 11/

high (in B. luzonica to 280-350 7-9 Uniseriate up pm); occa- III, per mm. rays 0-15%, com-

sionally with some sheath cells. Parenchyma posed of procumbent and upright cells, height aliform and aliform-confluent, sometimes in up to 4—8(—25) cells (130-160, sporadically

short in the of B. kurzii 420 Multiseriate of bands; sample only pm). rays composed pro-

narrow vasicentric parenchyma present. Paren- cumbent cells except for the uniseriate margins chyma strands of 3-4 cells. Rhombic crystals of 1—3 square and/or upright cells; 3—5 (—7) the in marginal ray cells, and occasionally in cells wide, up to 410—510 pm high; occasional- the procumbent ray cells and the axial paren- ly with some sheath cells. Parenchyma scanty chyma. Radial latex tubes present, sometimes paratracheal, occasionally aliform with short

diameter 20-45 in few, pm, (much) larger than wings, and initial bands up to 20 cells wide. the cells in surrounding ray tangential section. Parenchymastrands of 3—4 cells. Rhombic crys-

Note: Broussonetia kurzii shows druses and restricted the common, to marginal ray cells, or

few rhombic in the cells. As distributed the entire and in the axial crystals ray only over ray,

sections of this were data Radial latex species available, on parenchyma. tubes scarce or ab-

element length cannot be provided, sent, diameter 20 pm, only slightly larger than

the cells in surrounding ray tangential section. Broussonetia Broussonetia sect. (Table 1; Figs. Note: The tropical samples of B. papyrifera

1,4-8) show less distinct growth rings (Uw 26750) or This section four comprises species (Comer, no growth rings at all (Uw 24338).

1962), distributed in Asia. Broussonetia papyri- fera has been introduced into temperate and Maclura Nutt.

outside tropical areas its natural distribution The genus Maclura s.l. in the classification of Small 35 area. trees or trees up to m, mainly in Comer (1962) consists offour sections; Cardio- subtemperate forests, or the more seasonal parts gyne, Chlorophora, Cudrania , and Maclura. Be-

of tropical forests. fore these sections considered then, were as sepa- Material studied: B. Siebold. kaempferi rate genera. Berg (1977a) modified this broad FOFw TWTw 532 Japan: (Uw 26981), (Uw genus concept. In his opinion Plecospermum,

26782). — B. kazinoki Siebold. Japan: T. treated Comer as a separate genus by (1962), Kishima 715 (Uw 26764), S. Okamoto 836 should be included in Maclura s.l. Berg (1977a)

(Uw 26765), FOFOw 7424 (Uw 26979), also mentioned that Chlorophora excelsa and

TWTw 331 — (Uw 26783). B. papyrifera (L.) C. regia should be excluded from Maclura s.l., Vent. Japan: TWTw 70 (Uw 26784); Thailand: and consequently transferred them to Milicia Kasin 392 (Uw 24338); Indonesia: IND. col. (Berg, 1982). Chlorophora tinctoria remained

9400 Netherlands (Uw 26750); (cult.): Cantons- included in Maclura s.l., although Kaastra 4718 park (UNw 213), Wagen. Arb. C 10668 (1972) preferred the genus rank for this taxon.

(Uw 26761); loc. unknown: s.n. (Uw 26753).

General features: Growth rings mostly Maclura sect. Cardiogyne (Bureau) Comer

colour to distinct; light brown cream, no de- (Table 2; Figs. 2, 9) marcation between sapwood and heartwood. A section with one species,M. africana (Berg, Texture medium to coarse, grain straight. Spe- 1977a). In his discussion the taxonomic on po- cific gravity 360—650 N per cubic metre. sition of M. brasiliensis, Kaastra (1973) con- features: Microscopic Wood mostly sidered this taxon congeneric with Cardiogyne Vessels ring-porous. solitary (40-80%) and in africana and (sect. Maclura short radial multiples of 2-4 (in the latewood sensu Comer). Berg (1977a) does not follow clusters of few 50 irregular to narrow vessels, this idea, although he states that Cardiogyne is diameter 10—50 4-20 pm); per sq.mm, round closely related to and probably even congeneric to oval, diameter 80-215 vessel member with Maclura pm, s.s. In this paper, Cardiogyne is length 230—315 Perforations pm. simple, end considered as a section of Maclura with only walls almost transverse. Intervascular pits alter- one species, whereas M. brasiliensis is treated round, oval to nate, polygonal, 6—13 pm. Ves- under Maclura sect. Maclura. Maclura africana sel-ray and and is distributed in southern Africa vessel-parenchyma pits larger as evergreen irregularly shaped, half-bordered, the borders shrubs or treelets, up to 7 m high, in coastal sometimes reduced. Thin-walled tyloses pres- scrubs, on riverbanks or even rainforests, from ent. in the latewood to Spiral thickenings vessels sea level up 1000 m altitude. 96 lAWA Bulletin n.s., Vol. 7 (2), 1986

tubes latex radial— -S* T3 S 2 1 (+)3 (+) 1 1 1 1 dis-

w w procum- Is S2 p)*T3 *3 p) p) vessel II u *T3 c c >. 5 s parenchyma distribution 3crystal p'(u, p' (u,'f p'(u, of pac, ac,(p)o 65 ac,(p)O p ac,(u,p)o 65 O ac, ac, p 65 u> 63 ac,

1 lower.

+ + 3 m3 1 + 1 + composed 69 (+)3 o chambered 3

— o* , much + + + + + + *< wb* (+)3 + (+) /■“s i.e., in 3* c

o TC Oc + + + + + 3(+) + ac: 3

a> r-> 65 + diameter a + '+■(+) (+) + 1 + Parenchyma t/5distribution -(+)+ *a o- crystals:

1

W' 5 and (Kribs) g.>: ray2 - O (III) (111) (III) K (III) ho: - 5S type II ho (II) 3“ o o o ho ho ho 1 ho initial). 3* 3* 3" 3* higher or ■u (1968); 44- O» V/> to (/) O c ray S ! -"“Scells) (in width3* a multiseriate.» .a E 3OJ 3-4 1 3-4 1 1 4■fc* to2-5 3-5 1—2 Plecospermum. u> u> u> N< much Rays69 Kribs terminal o pa to K) «« « | ° rows£ U) cellsImarginalI of O J_ 1-21 0 I >30 0—2 to and o0-1 O0-1 O0-1 V O (i.e., o frequency s.l. NO oo 00 On >. mm)'c'1(perI frequencyc s £ 1 1 o10 1 6-9 4-8 3-8 8-10 On 5-7 5—6 OJ Ol Ol according 00 vessel marginal Oi On Maclura Ol o o O types VO to -1040 -995 tO m: = ;-950 a* pun) (in0 length-s o i o of 1 L/l — ￿- Fibres o è 825-950j 810-1040 740-965 t 685-99500 latewood -n oo O' oo i bands;

1 the a S' 1 thickenings|I ~spiral characters 1 1 +/- + + 1 1 in heterogeneous wavy ||? ￿: IO in 00 VO VO III: o- NO9 pun) .5(in size.a pit 9VO 7-8 1 8-91 8-91 1 o 8-11 8 00 9—12 8 9 II, wb: developed; I, «*-■=" o O anatomical to U) NJ pun)E (in£I.lengthJ! memberS-sE E 1 i—265 1-250 1 type: 300 175-265 190-240 345345 305 confluent; 250-320 -J 220-250 poorly cells. wood 25( 22( S ray c: or ray *

* -o o Some -< 1 oi * Ol aliform; 0 IO O | 2 present Vessels sE ?pun) (in S O diameter £145 On 2. -- 1 ——— 160* 1070 0 90-180- 70/28070/280 105-180105-180 -- 1020 -110-250* a: —— 1120 procumbent 1 140-160/275* — 11— 14( semi-ring-porous. Table in

p: * 3030 vO occasionally * se: to vasicentric; EE sq.sr I >. c o* 12 1 o mm) (per frequency«£: 5-9 1 50* Ul 5-7* 01 5-20»5-20* Oi 5-10»5-10* app. cells; (): v: ~§ 5«a> ray sen £: = /> t distribution.2 “■ - ring-porous; D-di &di ri ri o.di absent; - - — ri,3. se,(di)p i upright -: ri: parenchyma: -5 2 and 0 2 :s- s- 2 S' K- diffuse; 1 2 present; 1 S 1 8- 1 1 B* s square 2. cT S' •s 1 s di: only. Cardiogyne C3 Chlorophora Cudrania 2 2 +: D a* Maclura & ricana 9 p cochinchinensis tricuspidataS I S in S' brasiliensis2 | f S’tinctoria o pomifera5 n $ 3. ■g 3 a 3 3 O 3 ■Q cells u: o’ O o’ Species-o o' a C/5 M.s M. M.s sM. M. M.s r. o O r. Plecospermum Legends: tribution: Sectionft Sectionft Section Section bent cells; C/5 on if SP lAWA Bulletin n.s., Vol. 7 (2), 1986 97

2. Element of Madura s.1. and Fig. length Trophis s.1., including Plecospermum and Olmedia.

Material studied: M. africana (Bureau) 1020 3.4. pm; length pm; F/V-ratio Rays uni- Comer. Botanical Harare; Zimbabwe, Garden, seriate and multiseriate, heterogeneoustype 11,

s.n. B—lo Berg (Uw 27394). per mm. Uniseriate rays 30% composed General features: Growth of rings absent, procumbent and upright cells, height up to

sapwood white to heartwood not avail- 6 cells yellow, (270 pm). Multiseriate rays composed able. Texture coarse, grain straight. Specific of procumbent cells except , for the uniseriate

gravity 570 N cubic metre. of I—2 per margins rows of square and/or upright

Microscopic features: Vessels diffuse, 3 cells 600 cells; wide, up to pm high. Paren- solitary (25%) and in radial multiples of 2—7 chyma aliform with short or long wings, often and clusters of 12 some irregular 2-3; per sq. confluent and in wavy bands. Parenchyma round diameter 145 mm, to oval, pm, vessel strands of 3—4 cells. Rhombic crystals in the member 300 Perforations length pm. simple, ray cells, and in the axial parenchyma in cham-

end walls almost transverse. Intervascular pits bered cells, up to 4 crystals in axial rows. alternate, round, oval or polygonal, 9 pm. Ves-

and and sel-ray vessel-parenchyma pits larger Maclura sect. Chlorophora (Gaudich.) Bureau irregularly shaped, half-bordered, the borders (Table 2; Figs. 2, 10-12) sometimes reduced. Thin-walled tyloses com- Since Berg (1982) assigned the African spe-

mon. Fibres with small restrict- to nonseptate, pits cies of this section the reinstated genus Mili- ed to the radial walls; walls 3-4 lumina 5 Sim, the pm, cia section comprises only one species I AWA Bulletin Vol. 7 1986 98 n.s., (2),

(two additional species described in Chloropho-Maclura is found from Sri Lanka and

to ra were previously transferred to Broussonetia India China and Japan, and throughout the

by Berg, 1977a). The species is distributed in Malesian Archipelago, as climbers in lowland

the Neotropics. The trees are up to 30 m high, forests and occasionally up to 1800 m altitude.

and occur in various habitats from sea level up Maclura tricuspidata is distributed in the humid

to 1600 m altitude. temperate part of Central China and Korea, as

Material studied: M. tinctoria (L.)Don. trees or shrubs.

Mexico: (Uw 25792); Panama: USw 664 (Uw Material studied: M. cochinchinensis

7135); Colombia: SJRw 1893 (Uw 18434); (Lour.) Comer. India: Gamble 0.5006 (Uw Venezuela: SJRw 10031 (Uw 18419); Brazil, 18395), 0.5007 (Uw 18396); China: Y. Tang

Parana: Lindeman & H. de Haas 1671 (Uw 0526, ex SJRw 21994 (Uw 18420); Indonesia:

13243), 1853 (Uw 13369);Argentina: Pederson Woodworth 4609 (Uw 24256), Volkers XI B

856 (Uw 26755), USw 458 (Uw 7107). 327 (Uw 25799), XI B 286 (Uw 25798); Aus-

General features: Growth rings absent tralia,Queensland; F.P.R.L. 18982 (Uw 18521).

F.P.R.L. 14677 or faint; heartwood dark brown, sharply defined M. tricuspidata Carr. China:

from the light brown sapwood. Texture me- (Uw 18385); U.K. (cult.): Kew, Temperate

to dium coarse, grain interlocked. Specific grav- House (Uw 18397).

ity 600-970 N per cubic metre. General features: Growth rings dis-

Microscopic features: Vessels diffuse, tinct in the wood samples of the temperate solitary (40-55%) and in short radial multiples, regions M. tricuspidata, and part ofM. cochin-

de- and few irregular clusters of2—4(—B); 5—9 per chinensis); heartwood dark brown, sharply

round diameter 105-180 fined from the brown sapwood. Texture sq.mm, to oval, pm, light

vessel member length 250—320 pm. Perfora- medium to coarse, grain straight. Specific gravi-

tions simple, end walls almost transverse. Inter- ty 750—950 N per cubic metre.

vascular pits alternate, round, oval and poly- Microscopic features: Wood ring-po-

to diffuse- gonal, 7—B pm. Vessel-ray and vessel-paren- rous to semi-ring-porous, tending

chyma pits larger, elongated, more irregularly porous in the samples from tropical areas. Ves-

shaped, half-bordered, the borders commonly sel(s) solitary (18—52%) and in short radial

2—4 very reduced. Thin-walled tyloses abundant. multiples and irregular clusters of (in the latewood Fibres nonseptate, with small simple pits re- up to 12); 5-20(-50) per sq.mm,

stricted to the radial walls; walls 3—4 pm, lumi- round to oval, diameter 90—250 pm (in the

4-9 810- wood to 400 vessel member length na pm, occasionally gelatinous;length early up pm), 175—265 Perforations end walls al- 1040 pm; F/V-ratio 2.9—3.9. Rays uniseriate pm. simple,

and multiseriate, homogeneous, rarely hetero- most transverse. Intervascular pits alternate,

6—9 mm. Uniseriate oval and 8-11 Vessel- geneous type 111, per rays round, polygonal, pm.

0-5%, composed mainly of procumbent cells ray and vessel-parenchymapits larger,elongated,

the borders and few square and/or upright cells, height up irregularly shaped, half-bordered, Multiseriate in to I—s cells (60—120 pm). rays part reduced, occasionally unilaterally com-

composed of procumbent cells only; 3—4 cells pound. Thin-walled tyloses abundant. Fibres

to 170—400 ali- with small restricted to wide, up pm high. Parenchyma nonseptate, simple pits short and the radial 2—3 lumina 5—7 form, with long wings, occasionally walls; pm, pm, oc-

abaxially unilateral, confluent, occasionally in casionally gelatinous;length740—950 pm; F/V-

wavy bands, and occasionally in narrow ter- ----ratio 3.1—4.3 Rays uniseriate and multiseriate, minal bands. Parenchyma strands of 2—7 cells. homogeneous and occasionally heterogeneous

Rhombic crystals abundant in the axial paren- type HI, 3—B per mm. Uniseriate rays 5-48%,

chyma, in crystalliferous strands containingup composed of procumbent and some square

to 15 crystals, one crystal per chamber; occa- and/or upright cells, height 3—5(—12) cells of sionally also in the ray cells. (80—240 pm), Multiseriate rays composed

Note; Radial and axial latex tubes were procumbent cells, and only occasionally with

7107 from I—2 of of observed in only one sample (Uw (—4) rows uniseriate margins square

2—5 cells 540 Argentina). and/or upright cells; wide, up to

1200 pm high. Occasionally with some sheath

Madura sect. Cudrania (Trecul) Comer (Table cells. Parenchyma aliform with short or long

2; Figs. 2, 13-16) wings, confluent, and sometimes wavy, result-

A section with five species, which are distri- ing in a reticulate pattern; in the prominently

buted through Asia, Australia, and Japan. The ring-porous samples initial bands including

habit is variable ranging from shrubs and trees the larger pores of the earlywood. Parenchyma

to climbers. The two species studied here show strands of 2—4(—5) cells. Rhombic crystals

in a different distribution and habit. Cochinchinensisabundant the axial parenchyma, in crystalli- lAWA Bulletin Vol. 7 1986 99 n.s., (2),

to ferous strands containing up 14 crystals, one rays composed of procumbent cells, and only

chamber or few in the of crystal per cell, crystals occasionally one marginal row square or up- rays. right cells; 3-5 cells wide, up to 495—930 pm

Notes: Spiral thickenings were observed in high, sporadically with some sheath cells. Pa- both ofM. and one sometimes samples tricuspidata sample renchyma confluent, wavy, tending of M. cochinchinensis (Uw 18420 from China). to short bands. Parenchyma strands of 2—5

Radial latex tubes, diameter 25 pm and much cells. Rhombic crystals abundant to common

than the cells larger surrounding ray in tangen- in the axial parenchyma, in crystalliferous tial and also axial latex tubes observed strands to 15 section, containing up crystals, one crys- in one sample (Uw 24256). tal per chamber or cell, occasionally few rhom-

bic crystals in the rays.

Maclura sect. Maclura (Table 2; Figs. 2, 17—19) Notes: In two samples (cult. UNw 262 and

A section with two species, M. pomifera and Uw 26769) crystals are lacking. Maclura brasi-

M. brasiliensis. For details the on taxonomy see liensis deviates from the description in a num- under Maclura sect. Maclura Cardiogyne. pomi- ber ofmainly quantitative characters. This may

is native to the U.S.A. the fact fera a temperate species, be explained by that the sample is ofa

The trees or shrubs are up to 20 m high. Maclura limited diameter, it is the only sample from the brasiliensis is found in South and Central Ameri- tropics, and it is a liana. Main differences (next

and is said to have liana-like habit. to ca a the differences indicated in Table 2) are;

Material studied: M. brasiliensis (Mar- F/V-ratio 3.1; uniseriate rays up to 35%. Multi- thas) Endl. Venezuela: 61949 seriate 1-2 cells wide and 1500 Steyermark (Uw rays up to pm 26767). M. pomifera (Raf.) C. Schneider. high.

U. S.A.: locality unknown, USw 1531 (Uw In M. pomifera the parenchyma distribution

7227), USw 5843 (Uw 7441), USw 7391 (Uw is also vasicentric, aliform, and marginal.

8462); A.F. Wilson s.n. (Uw 10222); Marts s.n,

(Uw 24258); Israel (cult.): Woodworth 4 (Uw Plecospermum Trecul (Table 2; Figs. 2, 20, 21)

24259); the Netherlands (cult.): (UNw 262); Comer (1962) regarded this taxon as a genus.

U.S.S.R. (cult.): (Uw 26769). Berg (1977a) placed it near Cardiogyne, and in

General features: Growth rings dis- 1983 suggested inclusion in Maclura s.l.,but did tinct in M. pomifera, absent in M. brasiliensis; not formally reduce it to a section of the latter heartwood dark defined The of clim- golden brown, sharply genus. genus comprises two species

from the lightbrown sapwood. Texture medium bers, found in tropical,rather dry habitats, and to coarse, grain slightly interlocked. Specific is distributed throughout India, Sri Lanka, Bur- N cubic gravity 600—1000 per metre. ma, Thailand, and the Andaman Islands.

Microscopic features (mainly based Material studied: P. spinosum Trecul.

on M. pomifera, see note): Wood ring-porous. India: For. Dept. 1878 E 487 (Uw 18400).

Vessels solitary (40-80%) and in short radial General features: Growth rings dis- multiples and irregular clusters of 2—4 (in the tinct; sapwood light brown, heartwood not latewood s—lo up to 10); per sq.mm, round available. Texture coarse, grain straight. Speci-

and diameter 140-160 the metre. oval, pm (in early- fic gravity 650 N per cubic wood up to 275 pm), vessel member length Microscopic features: Wood semi-

190-240 Perforations end walls al- pm. simple, ring-porous to diffuse-porous. Vessels solitary most transverse. Intervascular pits alternate, (5%) and in radial multiples (especially in the

oval and 8-9 round, polygonal, pm. Vessel-ray latewood), and some irregular clusters of 2—10;

and vessel-parenchyma pits larger, elongated up to 50 per sq.mm, round to oval, diameter

and often 160 vessel irregularly shaped, half-bordered, pm, member length 305 pm. Perfo- with reduced borders, sometimes unilaterally rations simple, end walls almost transverse. In-

compound. Thin-walled tyloses abundant. Spi- tervascular pits alternate, round, oval and poly- ral 9 thickenings commonly present, more prom- gonal, pm. Vessel-ray and vessel-parenchyma inent in the narrow latewood vessels. Fibres pits larger, elongated, and irregularly shaped, nonseptate, with small simple pits restricted to half-bordered, the borders reduced, sometimes the radial walls 3—4 lumina 4—B walls; pm, pm, unilaterally compound. Thin-walled tyloses occasionally gelatinous; length 685-995 pm; abundant. Fibres nonseptate, with small simple

F/V-ratio 3.5-5.0. Rays uniseriate and multi- pits restricted to the radial walls; walls 2—3 pm, seriate, homogeneous, occasionally heterogen- lumina 5-8 pm; length 1120 pm; F/V-ratio 5—7 Uniseriate eous type 111, per mm. rays 3.7. Rays uniseriate and multiseriate. Homo-

10-25%, of cells and and composed procumbent geneous occasionally heterogeneous type few upright cells, height to mm. square and/or up 111, 5—6 per Uniseriate rays 5%, composed cells Multiseriate 3—6(—12) (100—240 pm). of procumbent cells, and few square and/or up- 100 lAWA Bulletin n.s., Vol. 7 (2), 1986

'w' in (a)w a05 D>a » t» distributionI£C« 3crystal*s dis- y'(p,a) (p,a)y p,y p,y (p,a) (p,a) a: p,y y' y' vessel procum- 1 cells; + + + + of BJ 3 3 + + + + + +

o 3 ray

i + N< t wb 1 i 1 1 1 1

tr u lower.

o b c 1 3 1 + i 1 1 composed 3 i

r -(+)-+ n much t (+)(+) a 1 1 1 + i + + procumbent Parenchyma «**"S distribution i.e., w z (+)--++ +(+)-- +(+)--

*3 o. < + ++ + + 3 + + in I gvacwbm^' p: 1 diameter (II) g. ray (ho) (ho) (Kribs)C K/c- type s111 sIII IIIB ni(ii) BIII IIIIII III III vessel homogeneous, crystals:

O and O 8 ho: *• 2 1200 1080 pun) ê(in Q0 -720 -840 ON 1 I 1 ray « g 0 bands. heighti multiseriate.1 E Or» 0 w o higher Rays 425-425-840 555-840or» 500-720O 495-495-840 270-600 625-1200 480-1080 ?o y Or» Or» K> 625- (1968); Morus. wavy

O on much cells) t(in is Os 'Ôn •k is in of 6 1 Kribs ray «S' 4-6 4(6) 4-6 3-4 4-5 4-6 width «multiseriate.« §.s-s.2§S c OJ 4>- oft to wb: oo ON 00 il on D- g. species mm) (perW frequency= 4-8 OJ3 4 5-6 6-8 4-5 4-5 frequency •£t on (Os 4». -o I according

1 (V) O U» vasicentric; o vessel O J 1 NO a On 0 O o NO £ 00 O types v: E;5 c individual O' pun) (in lengthJï 1 K)1280 1100 Fibres 695695-950 1300-1935 1275-1435 790-1020 T1 790 of 1300 1245 1275 latewood I initial);

1 gob the or thickeningsi S 8- + + + !§ spiral 1 (+)2 1 + + in heterogeneous features OJ NJ hS o *“* *”* *: " Os pun)- .S(in size pit’g. 11 6 1 terminal 8-10 11-13 O10-12 8-11 7-11 00 00 ~-J III: - II, (i.e., on O on 8 O developed; I, to anatomical to OJ U) K> I oo £ pun) (inIlengthC J8 member5-gE 580 1 1 1 y 305505 380 I type: NO195-225 280-350y OJ335-400 on 230-270 vT 350-355 OJ N- NJ OJ oj K) marginal O) poorly Wood ray m: * or -170* pun) _(in diameterSgI=3 NO lO 1 1 ? ~ 190 125 1 1 O on75-115* oo180-250 155-200 present Table 155 135-135-150* 120-120-170* confluent;

* * * c: o On semi-ring-porous. on on OJ * - OJ 8oo -1 -1 1245-1580 ?sq. o. g. 8* Imm) (per frequency=S 1 00 2-5 4-5 1 Li ro -£» on- 0 13-16* 25-30* 30-35* OJ occasionally K) se: Cw.g So- OJ aliform; (): 2.ri ri3. n ri2. 3.ri a: distribution.2 1£ o.di se Q.di

I

n K- - — 3 — ring-porous;- a> n absent; O o o zone 3 N -a n> g 9 o Z CJ o o £ 1 1 ri: 3 3 3 £L parenchyma: c n n a» montane o o p zr subtropical T3 ■S montane montane cells. 3 lowland 3 3 3 •3 | é 3 1 3 Climatic temperate tropicaltropical tropical tropical tropicalo temperate/ temperate Q n rt rt present; diffuse;

S di: only, 1 +: -S

c parenchyma f 2 «1 cells r 2 t g* «* r insignis macroura mesozygia nigra rubra o alba S' SE sa alba 2 s n Species*3 Legends: tribution: axial co M.is M.s M. M.S M.£ sM. M.s bent IAWA Bulletin Vol. 7 1986 n.s., (2), 101

to to right cells, height up 5 cells (180 pm). Multi- common, the diameter equal the ray cells in of cells 10—15 seriate rays composed procumbent ex- tangential section, pm.

cept for the uniseriate margins of o—2 rows of

these Milicia Sim square and/or upright cells, margins only (Fig. 23) sporadically present; 4 cells wide, up to 960 Recently Berg (1982) reinstated the genus

with few sheath cells. Milicia the African of pm high. Occasionally Sim, comprising species

Parenchyma rather abundant,vasicentric to ali- Chlorophora, viz. C. regia and C. excelsa. Ear- form, confluent to reticulate, and as initial lier in 1977 Berg had mentioned that C. regia

the and C. excelsa bands, 5—15 cells wide, and including pores probably constituted a separate

of the of I—2 Kloos leaf anatomical fea- earlywood. Parenchyma strands genus. (1982), using cells. Rhombic crystals abundant in the axial tures, came to the same conclusion. The species parenchyma, often in crystalliferous strands are distributed throughout tropical Africa,

containing up to 16 crystals, one crystal per from Guinea-Bissau to Mosambique. The decid- chamber. Rhombic also in the 35 found in crystals rays, uous trees up to m high are tropi- but far less common, and also in the tyloses. cal lowland forests, ranging from rainforest to

forest islands in savannaregions.

Malaisia Blanco (Fig. 22) Material studied: M. excelsa (Welw.)

A monotypic genus (Comer, 1962), distri- C.C. Berg. Tanzania: Schlieben s.n. (Uw 6423); buted throughout tropical Asia, from southern East Africa: Schlieben 11026 (Uw 15648).

China, Thailand to Australia and the Fiji Islands. M. regia (A. Chev.) C.C. Berg. Guinea-Bissau:

This woody climber, shrub or treelet is chiefly G-7 (Uw 5636);Ivory Coast: Leeuwenberg 2471 found in forest edges of the tropical lowland (Uw 6545).

to and swamp forests, up 1200 m altitude. General features: Growth rings absent

Material studied: M. scandens (Lour.) or faint; heartwood light to dark brown, sharp- Planchon. Australia: DFP 6608 (Uw 24362); ly defined from the yellowish sapwood. Texture locality unknown: ex MAD-SJRw 16710 (Uw coarse, grain straight to interlocked. Specific

24630). gravity 520-740 N per cubic metre. General features: Growth rings faint Microscopic features; Vessels diffuse, or absent; heartwood dark brown, sharply de- solitary (40—80%) and in short radial multiples

fined from the light brown sapwood. Texture or irregular clusters of 2—4; I—7 per sq.mm,

coarse, graininterlocked. Specific gravity 700— round to oval, diameter 160—215 pm, vessel

800 N per cubic metre. member length 350-400 pm. Perforations sim-

Microscopic features: Vessels diffuse, ple, end walls almost transverse. Intervascular solitary (70-80%) and in short radial multiples pits alternate, round, oval or polygonal, 10—12

and irregular clusters of 2—4; 5 per sq.mm, pm. Vessel-ray and vessel-parenchyma pits larger, round to oval, diameter 150—170 pm, vessel and irregularly shaped, half-bordered, the bor-

member length 440—470 pm. Perforations sim- ders sometimes very reduced. Thin-walled ty-

ple, end walls almost transverse. Intervascular loses common. Fibres nonseptate, with small

pits alternate, round, oval to polygonal, 11 simple pits restricted to the radial walls; walls

and 3—4 lumina 9—14 pm. Vessel-ray vessel-parenchyma pits pm, pm, occasionally gelat-

and half- 1050—1425 2.9 larger, elongated, irregularly shaped, inous; length pm; F/V-ratio bordered, the borders sometimes reduced. Thin- 3.7. Rays uniseriate and multiseriate, hetero-

walled Fibres with 4-6 Uniseriate tyloses present. nonseptate, geneous type 111, per mm. rays

small restricted to the radial of and simple pits walls; 0—15%, composed upright square cells, to walls 3—4 pm, lumina 15 pm, occasionally ge- height up 2—5 cells (110-220 pm). Multi- 1345—1495 latinous; length pm; F/V-ratio seriate rays composed of procumbent cells,

3.0—3.2. Rays uniseriate and multiseriate, het- with uniseriate margins of 1, sporadically 2

3—5 Uniseriate of 3—5 erogeneous type 11/111, per mm. rows square and/or upright cells; (—6)

of and cells 315—460 rays 15—17%, composed upright square, wide, up to pm high; occasional-

and few procumbent cells, height up to 3—5 ly with few sheath cells. Parenchyma aliform to

cells (260-340pm). Multiseriate rays composed confluent, and often in wavy bands, and occa-

of procumbent cells except for the uniseriate sionally marginal. Parenchyma strands of 3—4

of I—2 rows of upright cells. Rhombic in the margins and/or square crystals common square

cells; 3—4 cells wide, up to 570—840 pm high, and upright marginal cells of the multiseriate

sheath cells. occasionally with some Parenchyma rays and the uniseriate rays, always present in

vasicentric-aliform with short wings, only occa- the axial parenchyma but far less common. Ra-

sionally confluent. Parenchyma strands of 4 dial latex tubes, 25-50 pm in diameter,(much)

cells. Small quantities of vitreous silica present larger than the surrounding ray cells as seen in

in the fibres and vessels. Radial latex tubes tangential section, infrequent, often absent. 102 lAWA Bulletin n.s., Vol. 7 (2), 1986

Morus 3—B L. (Table 3; Figs. 1,24-32) homogeneous, per mm. Uniseriate rays

A recent monograph of Morus is not avail- 2—23%, composed of procumbent, square and/

able. About 10 species are recognised. The or upright cells, height up to 3—12 cells (100-

genus is widespread,with species in the northern 300 pm). Multiseriate rays composed of pro-

hemisphere (U.S.A., Europe, Japan, and China), cumbent cells except for the uniseriate margins

and the tropical regions including Andes, Cen- of 1 —3 rows of square and/or upright cells; 4 -

tral America, Africa, and Indonesia. The trees, 6 cells wide, up to 425

treelets or shrubs occur in various forest types, sionally with some sheath cells. Parenchyma

from sea level up to 2500 m altitude. paratracheal, varying from vasicentric to ali-

Material studied: M. alba L. The Neth- form-confluent and in confluent wavy bands,

erlands (cult.): Cantonspark 4483 (UNw 309), and apotracheal as marginal bands. Parenchyma

Utrecht Hot. Gardens s.n. (UNw 716); South strands of 2—4 cells. Rhombic crystals, pre-

Africa: H. Baijnath s.n. (Uw 25505); India, dominantly in the uniseriate margins of the

Brandis 95H M. cel- multiseriate and the Himalaya: (Uw 25787). rays uniseriate rays, in

tidifolia H.B.K. U.S.A.: Barghoom 8902 (Uw variable amounts, and few to scarce in the axial

9980). - M. insignis Bureau. Colombia: Cuatre- parenchyma.

casas 18405 (Uw 24976). - M. macroura Miq. Notes: In M. mesozygia the multiseriate

India, Assam: F.R.I, 75804 (Uw 24229); In- rays are 3—4 cells wide and up to 270—480 pm

donesia, Java: Koorders 8742b (Uw 24425), high. Radial latex tubes, 30—45 pm in diame-

141938 - M. and much than the (Uw 24426). mesozygia Stapfex ter, larger surrounding ray

A. Chev. Zaire: Wermuth s.n, (Uw 18763); cells (in tangential section) observed in one

Nigeria: F.H.I. 3956 (Uw 18399);West Africa: sample ofM. mesozygia.

MSP 36 (Uw 18398); Uganda: NO GM 39 (Uw Details on the wood anatomical features of

18522); Angola: s.n. (Uw 24513). M. nigra individual species are givenin Table 3 and Fig. 1.

L. The Netherlands: s.n. (UNw 43); Libanon:

Schweinfurth 1 (Uw 25788). M. rubra L. Olmedia Ruiz & Pavon: see page 110.

The Netherlands (cult.): Utrecht Bot. Gardens

72-199 (UNw 583); U.S.A.: USw W-3561 (Uw Pachytrophe Bureau (Figs. 33, 34)

7336), USw W-4233 (Uw 7399). Berg (1977a) revised the genus, and com-

General features: Growth rings dis- bined the two species recognised until that

tinct, faint or absent; heartwood brown to dark time into one species, P. dimepate. The genus is brown, sharply defined from the light cream endemic to Madagascar. The trees or shrubs are

Texture medium 30 and found in humid sapwood. to coarse, grain up to m high, to dry

interlocked. from level 1000 altitude. straight or weakly Specific gravity forests, sea up to m

560-850 N per cubic metre. Material studied: P. dimepate Bureau,

Microscopic features: Wood ring-po- Madagascar: Veenendaal & den Outer 1190 (Uw rous in the temperate species, faintly semi-ring- 26144), Thouvenot 23 (Uw 24257), U.P.J. 608,

to porous commonly diffuse-porous in the ex RBHw (Uw 26776).

tropical species. Vessels solitary (17—60%) and General features; Growth rings absent;

in short radial multiples or irregular clusters of heartwood lightbrown, not distinguishablefrom

2—4 (in the latewood of the ring-porous spe- the sapwood. Texture medium, grain straight.

cies clusters of to 8 910—1000 N cubic many irregular up pores); Specific gravity per metre.

2—35 per sq.mm, and up to 100 per sq.mm in Microscopic features: Vessels diffuse,

the latewood of the ring-porous samples, round solitary (55—70%) and in short radial multiples

to diameter 75—250 the latewood and clusters of 3-7 oval, pm (in irregular 2—4; per sq.mm,

as narrow as 30 pm), vessel member length round to oval, diameter 110—165 pm, vessel 195-400 Perforations walls pm. simple, end member length 355-515 pm. Perforations sim-

almost transverse, Intervascular pits round, oval ple, end walls almost transverse. Intervascular

and polygonal, 6—13 pm. Vessel-ray and vessel- pits alternate, round, oval or polygonal, 7—9 parenchyma pits larger, elongated and irregular- pm. Vessel-ray and vessel-parenchyma pits

ly shaped, half-bordered, the borders often re- larger, elongated and irregularly shaped, half-

duced. Thin-walled the borders often tyloses common. Spiral bordered, very reduced. Thin-

thickenings present in the narrow vessels of the walled tyloses abundant. Fibres nonseptate

(semi-)ring-porous samples. Fibres nonseptate, with small simple pits restricted to the radial

with small restricted the radial walls 3-4 lumina 8-12 simple pits to walls; pm, pm, occa-

walls 2-4 lumina 6-20 1190—1460 walls; pm, pm, occa- sionally gelatinous; length pm;

695—1935 2.7-3.4. sionally gelatinous; length pm; F/V- F/V-ratio Rays uniseriate and multi-

----ratio 2.9—5.5. Rays uniseriate and multiseriate, seriate, heterogeneous type 11, 3—5 per mm.

heterogeneous type 111, occasionally type II or Uniseriate rays 10—25%, composed almost ex- lAWA Bulletin Vol. 7 1986 103 n.s., (2),

clusively of square and upright cells, heightup ferent. Streblus glaber is distributed in Malesia,

to 4-9 cells (240-360 pm). Multiseriate rays Philippines, Indonesia and the Solomon Islands

of trees to composed procumbent cells except for the as up 40 m high in montane forests uniseriate of I—s of from 700 Streblus margins rows square and/ to 2500 m altitude. penduli-

or upright cells; 3-4 cells wide, up to 395 nus is distributed in the Pacific, Australia, and

840 eastern pm high. Parenchyma predominantly in the part of New Guinea as a shrub or

more or less concentric bands, 1.5—5 per mm, tree of up to 13 m high, found in the lowland

3—6 cells also ali- forests wide, scanty paratracheal, and along seashores. The wood anatomy

form with long and short wings, and occasion- of these two species differs in many features

ally confluent. Parenchyma strands of 3 cells. (Ter Welle, 1985). Because of these differences

Rhombic crystals abundant in the ray cells, es- the species will be presented individually here.

in the and less fre- pecially square upright cells,

quentin the procumbent cells and then smaller. Streblus glaber (Table 4; Figs. 35, 36)

Crystals occur occasionally also in the axial pa- Material studied: S. glaber(Men.) Cor-

renchyma and the tyloses. ner. Philippines: A.N. Green, F.P.R.I. 896 (Uw

24237); Indonesia: IND. col. E 3838 (Uw

Plecospermum Trecul: see page 99. 24433), Irian Jaya: BW 9290 (Uw 20480), BW

11165 (Uw 20740), BW 7106 (Uw 21171), BW

106. 7091 Sloetiopsis Engl.: see page (Uw 24414).

General features: Growth rings absent; Streblus Lour. heartwood dark brown, well defined from the

The delimitation of this genus is rather prob- lightersapwood.Texture medium,grainstraight.

lematic. Comer (1962) included many taxa, Specific gravity 900—1090 N per cubic metre.

formerly described as genera, as sections in Microscopic features: Vessels diffuse,

Streblus s.l. As he had studied the not genus solitary (26—66%) and in short radial multiples

he mentioned the Sloetiopsis himself, only pos- and inegular clusters of 2—4; 3-6 per sq.mm,

that this to sibility genus might be closely related round oval, diameter 150-220 pm, vessel

to Streblus sect. Sloetia. like Neosloetiopsis, member length 500 pm. Perforations simple,

Sloetiopsis native to Africa, is assigned to.Streb- end walls almost transverse. Intervascular pits

lus sect. Paratrophis by Comer (1962). Berg alternate, round, oval and polygonal, 8—11 pm.

(1977a) monographed Sloetiopsis and Neo- Vessel-ray and vessel-parenchyma pits larger,

and the latter in the and sloetiopsis placed synony- elongated, irregularly shaped,half-bordered,

of In the he the borders reduced. Thin-walled my Sloetiopsis. same paper men- usually very

tioned that Sloetiopsis and Sloetia may prove tyloses abundant, thick-walled tyloses scarce or

to be congeneric and may be regarded as related absent. Fibres nonseptate with small (occasion-

to Bleekrodea, another ofStreblus sensu section ally more prominent) simple pits restricted to

Comer. In his classification of the Moraceae the radial walls 3—5 lumina 8-15 walls; pm,

Berg(l9B3) Streblus s.l. totheMoreae 1360 assigned pm, occasionally gelatinous; length pm; with urticaceous stamens; however, an indica- F/V-ratio 2.7. Rays uniseriate and multiseriate,

tion of the taxa to be included was not given. heterogeneous type 11, s—B per mm. Uniseriate

Till hesitates in- of today, Berg (pers. comm.) to rays 15-24%, composed square and upright

clude Sloetia and in cells and Sloetiopsis Streblus, a con- some procumbent cells, height up to

reflection ofhis statement that s—ll cells tinuing (1977a) (205—360 pm). Multiseriate rays Streblus s.l. be appears to too heterogeneous. composed of procumbent cells, except for the

Material of Comer’s sections Paratrophis, uniseriate of I—s of margins rows square and/or and 3—4 Phyllochlamys, Pseudostreblus, Sloetia, upright cells; cells wide, up to 440—960

Streblus, as well as of Sloetiopsis was available pm high;occasionally with few sheath cells. Pa- for this study. The sections will be treated sep- renchyma scanty paratracheal, vasicentric, vasi-

arately here because of the differences in the centric-aliform with short and long wings, con-

various classifications. The sequence of the fluent and in wavy and often tangentialbands. sections is alphabetical,followed by Sloetiopsis. Parenchyma strands of 4 cells. Rhombic crystals

in the upright and square cells, and occasional-

Streblus sect. Paratrophis (Blume) Comer ly also in the procumbent cells of the rays, and The section comprises nine species in Asia, generally less abundant to scarce in the axial

and Australia. Polynesia, Trees, 13—40 m, oc- parenchyma cells in the same sample. Occasion-

curring in various habitats, from sea level to ally in chambered parenchyma cells in longitu- montane Two available for areas. species were dinal rows of up to 6 crystals. Vitreous silica this viz. S. study, glaber and S. pendulinus. The present in the fibres and sporadically in the

of these two ecology species is strikingly dif- axial parenchyma of some samples. 104 IAWA Bulletin n.s., Vol. 7 (2), 1986

Table 4. Some wood anatomical characters ofStreblus s.1. and Sloetiopsis

I ' i 1 i 1 i >

IIpm) a ~ C c cells

i! ofo is size,§ SS J 2 | £ » S Si §- ct"£ •> > rt ««-*—• « . a> c F/V-ratio -J 5 frequency diameter member tyloses length c frequency SpeciesSpecies .Es £ 'S.pit 2- J5 ttT^Srows

Section ParatrophisParatrophis

is 150-220 CO8 00 S.5. glaber 3-6u> 150-220 500 8-11 t,(T)3 1360CJ o to2.7’-o 5-8CO 1 00 T1-5CO

S. 225-300to CO u> o ON VO 1 vO o U) -J 00 o T S. pendulinus 15-22 60-85 to 1 6-81oo -t 915-1190 3.7-4.41 ■&> 8-10 1-4-p-

Section PhyllochlamysPhyllochlamys 1 S. taxoidestaxoides 110o 40-P-o £465CO 6ON -1 ioo;1005 2.2to to £14 1

Section PseudostreblusPseudostreblus

S. indicus 15Co 110O ê405CO 8-900 1VO t 1260to o 3.1c*j 7-j T1-2to

Section Sloetia

5. 100-135 1 S. elongatuselongates -o7-14£ 100-135 U)350-425Co 0 co U»3-51Co t,(T)3 1100-1300o o u> QO u>3.0-3.20 1 u> to 5-8CO 00 0-1(2)1 to

Section Streblus

90-120 - S. asper VO9-13U) 90-120 310-385u> 0 1u> CO 9-11VO t 985-1300vOoo 1 8 3.1-3.5U» T u> (/I 6-11On T1-2to

SloetiopsisSloetiopsis 20-40 30-50 300-350U) § U» o 2tO t,TH i900-1070o -oo 3.1-3.2u> T U) to 008-10o T1-4

+: t: Legends: present; —: absent; (): occasionally present or poorly developed. — tyloses: thin-walled; T; thick-

walled. — ray type: 1, II, III: heterogeneous types according to Kribs (1968); ho: homogeneous,i.e., composed of

cells — procumbent only. parenchyma: a: aliform; c: confluent; cb; in concentric bands; v: vasicentric; sp: scanty

Notes. Rhombic crystals are much more mon, Fibres nonseptate, with small simple pits

abundant in the samples from Irian Jaya than restricted to the radial walls; walls 2-4 pm, lu-

those of the other Faint mina 3—ll in provenances. pm, occasionally gelatinous; length

growth rings occur in Uw 24414, marked by 915—1190 pm; F/V-ratio 3.7-4.4. Rays uni-

differences in fibre wall thickness and lumen seriate and multiseriate, heterogeneous type II

diameter. (III), 8-10 pm per mm. Uniseriate rays 15-38

%, composed of square, upright and some pro-

Streblus pendulinus (Table 4 ; Figs. 37, 38) cumbent cells, height up to 7—14 cells (175

Material studied; S. pendulinus(Endl.) 340 pm). Multiseriate rays composed of pro-

F, Muell. U.S.A., Hawaii: Stem & Herbst 2948 cumbent cells except for the uniseriate margins

New Caledonia: of 1— 4 of (Uw 18561); s.n. (Uw 25784); rows square and/or upright cells;

Australia: DlO R-l 139-33 3—5 cells 151-96(Uw24387), wide, up to 450—950(—3000) pm R-535-21 Pa- (Uw 24388), (Uw 24389). high; occasionally with some sheath cells.

General features: Growth rings absent; renchyma scanty paratracheal and abundant as

3—14 cells 2-4 heartwood dark brown, the sapwood lighter concentric bands, wide, per

brown to yellow. Texture medium, grain slight- mm. Parenchyma strands of 2—4 cells. Rhom-

interlocked. 700—900 N bic abundant in the axial ly Specific gravity per crystals parenchyma,

cubic metre. less abundant in the square and upright ray

Microscopic features: Vessels diffuse, cells, and only few in the procumbent ray cells.

solitary (20-41 %) and in short radial multiples Vitreous silica in the vessels of some samples.

and irregular clusters of2-6; 15-22 per sq.mm, Note. A third species ofStreblus sect. Para-

round to oval, diameter 60-85 pm, vessel mem- trophis. viz. S. urophyllus Diels is represented

ber length 225-300 pm. Perforations simple, by one sample (Indonesia, Irian Jaya: Kalkman

end walls almost transverse. Intervascular pits 4938, Uw 18356). The parenchyma distribu-

alternate, round or polygonal, 6—B pm. Vessel- tion and the absence of crystals matches com-

ray and vessel-parenchyma pits larger,elongated pletely the description of S. pendulinus. The

and irregularly shaped, half-bordered, the bor- number of vessels per mm falls within the range

ders in part reduced. Thin-walled tyloses com- of S. glaber. Finally, the vessel diameter (96 lAWA Bulletin n.s., Vol. 7 (2), 1986 105

Table 4 (continued)

-o =r Rays Parenchymaco n 3 O- 3 CO

<

, 1 I

-9 mm)I I cells) 8 § S 5 „ 3 (per3 c C 'v width C i S, .2 distributiona a (in « T3 *-> 'JZ metre)» J3X) » w multiseriate3e>* “ _ number crystal £3vitreous specific CO ray cbO' wbO' width S (N Species*0 n> £ o(in 2 sp•o

￿ ￿ CO s o o- 3—4| 4k S11 i - -1 -1 900-1090 S.ÇO glaber>5_ft 3 Ui (+)V— (+)3 (+)3 (+),+. 2(+) u, p, a, (ac) (fi),(a)3i-' '— ( M < 700-900s ° S.?o 1 3-5U) | t/l II i i i + i - 3-14U) i- -fa- 2-4to 14^ pendulinus■Q5g S'6to (III)Ci (+)---W' + u, a,a,(p)(p) (ve)nW

+ c CO CO Ço5. taxoides11 2to I- (+)—/—v + 1 —1 —1 —1 2—3to 1 15Ln u, p,y a, aco 1 1

+ -+ 1 700-800o O10O o o Ço indicuscl 2to 111 1 1 -1 -1 - S. o'e to -(+)++ + + + acCOo

+ + + » CO W 900-1100O | S. o' 2-to 1 u> III 1 1 1 -1 -1 VOO o Ço elongatus11to (ho)3o w- -++(+)-- a, aco (ve),(fi),a3n 3 w-

+ 8-1200 to U) Olo o r-O Ol o & 3-U) I Ol III5 i i i —1-31 -1 sï 500-750 S. gasperS 2(+)---+ (+)2 ve,< (fi)

+ 10-140 — ■P» 'O900-930oo i VO U> o co a. S5- 3U) H= i i i -1 i 1 Sloetiopsis§ •§tO (+)-_-3 u,t-= ve,< (t)3

axial paratracheal; wb: in wavy bands. — crystals: u: in square and upright ray cells; p; in procumbent ray cells; a: in

silica distribution: parenchyma cells; ac: in chambered parenchyma cells; t: tyloses. — (vitreous) a: axial parenchy- 1

ma; fi: fibres; ve: vessels; t; tyloses. - ) based onimmature sample only.

pm) is intermediate between that ofS. glaber Streblus sect. Pseudostreblus (Bureau) Comer

and 5. pendulinus. respectively. Comer, in his (Table 4)

unpublished manuscript for the Flora Malesiana, A section with only one species, distributed

states that the species S. urophyllus may be on the Asian mainland; tall trees.

merely a high mountain form of S. glaber. Material studied: S. indicus (Bureau)

Comer. India: SJRw 23944 (Uw 24636).

Streblus sect. Phyllochlamys (Bureau) Corner General features; No data can be pro-

(Table 4) vided as only a small sectioning block was avail-

A section with only one species, distributed able.

from Sri Lanka, India, and Indochina to Male- Microscopic features: Vessels diffuse,

sia. Small, very shrubby trees in lowland forests solitary (23%) and in short radial multiples and

in rocky or dry places. Only one sample was irregular clusters of 2—5; 15 per sq.mm, round

available for this study. The diameter of this to oval, diameter 110 pm, vessel member length

sample is 0,4 cm. Therefore only an incomplete 405 pm. Perforations simple, end walls almost

wood anatomical description, and no details on transverse. Intervascular pits alternate, round,

general features can be provided. B—9. Vessel-ray and vessel-parenchyma pits Material studied: S. taxoides (Heyne) larger, elongated and irregularly shaped, half-

Kurz. Indonesia, Java: Bakhuizen van den Brink bordered, the borders occasionally reduced.

2028 (Uw 26779). Thin-walled tyloses common. Fibres nonsep-

Microscopic features. Vessels diffuse, tate, with small simple pits restricted to the

solitary and in clusters or radial multiples. Per- radial walls; walls 3-4 pm, lumina 7-11 pm,

forations simple. Intervascular pits round or often gelatinous;length 1260 pm; F/V-ratio 3.1.

polygonal, 6 pm. Fibres nonseptate. Rays uni- Rays uniseriate and multiseriate, heterogeneous

seriate and biseriate. Parenchyma scanty para- type 111, 7 per mm. Uniseriate rays 17%, com-

tracheal, but predominantly as concentric posed of square, upright, and procumbent cells,

to bands, 2—3 cells wide, c. 15 per mm. Rhombic height up 7-11 cells (240-360 pm). Multi-

crystals abundant in the ray and parenchyma seriate rays composed of procumbent cells ex-

the of I—2 cells. cept for uniseriate margins (—4) 106 lAWA Bulletin n.s., Vol. 7 (2), 1986

of 2 cells in rows square and/or upright cells; wide, renchyma cells, part chambered, one crystal

480 vasi- to 6 in a up to pm high. Parenchyma (narrowly per chamber, up crystals longitudinal

centric to) aliform with long narrow wings, row, occasionally very few. Vitreous silica pres-

with ent the axial and in confluent and in wavy bands, occasionally in parenchyma occasionally and/or a tendency to unilateral paratracheal, also in the fibres vessels. Radial latex tubes 20-30 than the some marginal bands. Parenchyma strands of pm in diameter, (much) larger

2-4 cells. Rhombic crystals few, in chambered surrounding ray cells as seen in tangential sec-

parenchyma cells, up to 4 crystals in a longitu- tion, sporadic, in three samples only.

dinal row.

Streblus sect. Streblus (Table 4; Figs. 41, 42)

A section with two species, distributed from

Streblus sect. Sloetia (Teijsm. & Binnend.) Cor- Southeast Asia to China and Indonesia. Trees

ner (Table 4; Figs. 39, 40) or shrubs up to 14 m high, in lowland vegeta-

A section with only one species, distributed tion, especially in secondary forests, and in

and Indonesia Trees in Malaya (Sumatra). up to open places.

35 in lowland forests Material studied: S. Lour. m high, as a canopy tree, asper Philip-

often in secondary forests, and surviving re- pines: Bur. Forestry 20257 (Uw 24439); Indo-

peated coppicing. nesia, Java; Reorders 1643 m 116178 (Uw Material studied: S. elongatus (Miq.) 24427), 136 c 248718 (Uw 24428), 136 d 95458

Comer. Malaysia: RB3-18, ex SFCw (Uw 24396), (Uw 24429), IND. col. 1360 (Uw 26763).

R 1110-78, ex SFCw (Uw 24394), DFP 12362, General features: Growth rings absent;

ex FPAw (Uw 24416), U.P.H. 373, ex RBHw brown to light brown (presumably heartwood).

(Uw 26778); Indonesia: R 575-99, ex SFCw Texture medium to coarse, grain straight. Spe-

(Uw 24395), IND. col. 713 (Uw 24444), Kru- cific gravity 500-750 N per cubic metre.

koff 4441 (Uw 25794), Krukoff 4334 (Uw Microscopic features: Vessels diffuse,

25794a). solitary (45—60%) and in short radial multiples

General features: Growth rings absent and irregular clusters of 2—5; 9—13 per sq.mm,

to 90—120 or faint; heartwood dark brown and sharply de- round slightly oval, diameter pm,

fined from the light brown sapwood. Texture vessel member length 310—385 pm. Perfora-

medium, grain straight to interlocked. Specific tions simple, end walls almost transverse. Inter-

gravity 900—1100 N per cubic metre. vascular pits alternate, round, sometimes poly-

Microscopic features: Vessels diffuse, gonal or oval, 9-11 pm. Vessel-ray and vessel- solitary (35—55%) and in short radial multiples parenchyma pits larger, elongated, and irregu-

and clusters of the borders often irregular 2—4;7—14 per sq.mm, larly shaped, half-bordered,

round to oval, diameter 100-135 pm, vessel reduced. Thin-walled tyloses abundant. Fibres

member 350-425 Perforations sim- with small restricted length pm. nonseptate, simple pits to

end walls almost Intervascular the radial walls lumina 7—ll ple, transverse. walls; 2—4, pm;

pits alternate, round, oval to polygonal, 3-5 pm. length 985—1300 pm; F/V-ratio 3.1—3.5. Rays

Vessel-ray and vessel-parenchyma pits larger, uniseriate and multiseriate, heterogeneoustype

half-bor- often elongated, irregularly shaped, 111, 6-11 per mm. Uniseriate rays 5-7%, com-

dered, the borders in part very reduced. Tyloses posed of upright, square, and some procum-

abundant, mostly thin-walled, occasionally bent cells, height up to s—ll cells (125—420

thick-walled. Fibres nonseptate, with small sim- pm). Multiseriate rays composed of procum- ple pits restricted to the radial walls; walls 3—4 bent cells except for the uniseriate margins of

lumina 2—6 1100-1300 I—2 of 3—5 pm, pm; length pm; rows square and/or upright cells;

F/V-ratio 3.0—3.2, Rays uniseriate and multi- cells wide, up to 370-960 pm high; occasional-

seriate, heterogeneous type 111, occasionally ly with some sheath cells. Parenchyma, concen-

s—B Uniseriate 1 and in B—l2 cells homogeneous, per mm. rays tric occasionally wavy bands,

of and I—3 and Pa- 17%, composed square, upright, some wide, per mm scanty paratracheal.

procumbent cells, height up to 4—5 cells (90— renchyma strands of 2—4 cells. Vitreous silica

of the 180 pm). Multiseriate rays composed pro- abundant in vessels, and sporadically also in

cumbent cells except for the occasional uni- the fibres.

seriate margins of 1 or 2 rows square and/or

upright cells, 2-3 cells wide, up to 400-800

vasi- (—1200) pm high. Parenchyma vasicentric, Sloetiopsis Engl. (Table 4; Figs. 43, 44)

centric-aliform with short wings and occasion- This monotypic genus has been monographed

ally long wings to confluent, with a tendency by Berg (1977a). According to Comer (1962)

to a unilateral distribution. Parenchyma strands Sloetiopsis might be related to his sections

of 3—5 cells. Rhombic in the axial Sloetia and of the Streblus. crystals pa- Paratrophis genus lAWA Bulletin Vol. 7 1986 107 n.s., (2),

In Berg’s opinion (1977a) Sloetiopsis and the Trophis P. Browne

section Sloetia may prove to be congeneric, but Corner (1962) included two taxa in Trophis

he hesitates to include these taxa in the genus P. Browne, which until then had been recognised

Streblus Comer. The is distributed distinct viz. and sensu genus as genera, Calpidochlamys

in Africa, from Guinea to Mo9ambique. Trees Maillardia. In his treatment of the African Mo-

or shrubs up to 10 m high, in primary and sec- raceae Berg (1977a) reduced the number of

ondary, rather humid forests, from sea level up species in Maillardia to two, but retained the

to 1000 m altitude. taxon as a distinct genus. However, he also

Material studied: S. usambarensis Engl. mentioned that Comer’s decision to include

Cameroun: Leeuwenberg & Berg 9686 (Uw Maillardia in Trophis s.l. mighteventually prove

24445); Ivory Coast: Bamps 21840 (Uw24236). to be correct. According to Berg (1977a) Calpi-

General features: Growth rings absent dochlamys is probably the closest relative of

or faint; light yellowish brown, no demarcation Maillardia. The same author accepted Trophis

between sapwood and heartwood. Texture fine, s.l. in 1983, but now (Berg, pers. comm.) hesi-

grain straight. Specific gravity 900—930 N per tates to include both Maillardia as well as Cal-

cubic metre. pidochlamys in Trophis.

Microscopic features: Vessels diffuse, The genus Olmedia was included in Trophis

solitary (60—70%) and in short radial multiples s.l. by Berg (1983), following his earlier state-

and irregular clusters of2—4; 20—40 per sq.mm, ments (1977a) that considerations about the

round to oval, diameter 30-50 pm, vessel delimitation of Trophis s.l. should also include

300—350 Perforations member length pm. sim- , Olmedia, and Clarisia. Clarisia and

ple, end walls almost transverse. Intervascular Sorocea were assigned to the Moreae without

pits alternate,round, or polygonal, about 2 pm. urticaceous stamens by Berg (1983). Burger

Vessel-ray and vessel-parenchyma pits much (1962) recognised the close relation between

larger, elongated (often tendingto scalariform), Trophis and Calpidochlamys but he did not

borders reduced them half-bordered, the very to combine in one genus. In Burger’s opin-

hardly visible, resulting in simple pit pairs. ion Trophis mexicana is closely related to

Thin-walled and thick-walled tyloses present. T. racemosa. Comer (1962) assigned T. race-

Fibres nonseptate, with small simple pits re- mosa (then T. americana0 to his subgenus

stricted to the radial walls; walls 2—4 pm, lu- Trophis, and T. mexicana to a separate sub-

mina 7—9 pm, partly gelatinous; length 900— genus Prototrophis. The two subgenera, and

1070 prn; F/V-ratio 3.1—3.2, Rays uniseriate the three sections sensu Comer (1962) will be

and multiseriate, heterogeneous type 11, B—lo treated here in alphabetical order, followed by

per mm. Uniseriate rays 25—40%, composed of the description of Olmedia. of and rows upright square cells, occasionally

one or more rows of procumbent cells, height Trophis subgenus Prototrophis Corner (Table5;

up to 12 cells (400 pm). Multiseriate rays com- Figs. 2, 45)

of cells for the posed procumbent except uni- A subgenus with one species distributed in

seriate of I—4 of Central margins rows square and/or America. Shrubs or trees up to 20 m 3 cells 600—720 in moist upright cells; wide, up to pm high, forests, and from sea level up to

high; occasionally with some sheath cells. Pa- 2000 m altitude.

renchyma apotracheal as narrow concentric Material studied: T. mexicana (Liebm.)

and occasionally slightly wavy bands, one cell Bureau. Guatemala: Sosa 50 (Uw 18029).

wide, 10—14 bands per mm and occasionally General features; Growth rings absent;

also scanty paratracheal. Parenchyma strands heartwood brown, sapwood yellow. Texture

of 3 cells. Rhombic in and crystals square up- medium, grain straight. Specific gravity 800 N

and abundant in the The right cells, tyloses. per cubic metre.

tyloses in the relatively small vessels are more Microscopic features; Vessels diffuse,

or less square to angular in radial and tangential solitary (20%) and in short radial multiples and

view. The crystals in these tyloses are generally irregular clusters of 3—6; 8 per sq.mm, round

larger than those in the ray cells, often several to oval, occasionally angular, diameter 145 pm,

in Vitreous silica vessel member 370 Perforations crystals a longitudinal row. length pm.

present in the vessels and the tyloses, often simple, end walls almost transverse to slightly

even the in the latter. Intervascular enveloping crystals oblique. pits alternate, round or

Notes. Differences in fibre wall thickness 9 and polygonal, pm. Vessel-ray vessel-paren- and lumen diameter produce faint growthrings. chyma pits larger, elongated and irregularly

The crystals in the tyloses were erroneously shaped, half-bordered, the borders sometimes

interpreted by Ter Welle (1985) as chambered reduced. Thin-walled tyloses present. Fibres

parenchyma cells. nonseptate, with small simple pits restricted to 108 lAWA Bulletin n.s., Vol. 7 (2), 1986

fD «silica<-> 360 vitreousg? 1 1 1 i ve «c« < scanty paren- „

aW in distribution|’-S J:2£.22-a crystal2 & a, ■p p,p, p, i u, p,p c a: u, u,c u, c parenchyma: cells;

1 ray 3 wb1 (+)(+) 1 1 (+)3 i 0 3 — cr + + + + + v< ** cb0 + ET c (1968). O cr aW 1 1 i i 3 procumbent

rt> W' + v (+)- (+)- Kribs Parenchyma (+)3 3 (+)3 3 distribution ++(+)+- in CO to "3 O- + (- sp 1 (+)3 1 (+)3 p:

1 (I)(1) cells; type«§ ray£• O(Kribs)*c >—' II= II= IIB II= II11 according Olmedia. ray

4- Ul n/i O'

oo cells) (in 5 rayM era * 1 a ,g widthx■§ multiseriateVI| 4-5 3-51 3-4 3-6 2-41 types and 4k OJ U) ro upright

s.1. 0; 0 -O ON o and CO Rays po ?o cellsc« *3" marginalw P<3of2rowsS 3 4k u> 4k 4k n/i 1-3(-7) i 1 l-4(-7) l-5(-10) l-4(-6)■ l-4(-7) Trophis s.sg'.s *-* (— - heterogeneous square in 0 of 00 oo -o III: c cl, ■p'mm)I (per >>frequency2 S £ O'6 1 1 u: 5-8 1 Ln 8-10 4k4-8 O'6-7 00 i II,

I,

OJ to .7-o crystals: to OJ JO 4k —i o vessels. characters s 1 1 ratio F/Vt 3.4 type: 4k4.1 T* u> 2.1-2.3 lyi2.S-3.2 U»2.3-2 JO JO JO ray ve:

Lft O bands. vO N/i NO ft> 4- 4^ Oi

to Ov opm)S (inc length anatomical O' o rS I Ol wavy 1525 1165 o Fibres 920-1920-1195 001380-1495 1250-1450 distribution: -n OJ 1250- developed. in wood wb: septate8 S + t- 1 1 1 1 (+) poorly

1 bands; (vitreous) u> O o or Some oo "?pm)5 (in.5w |size pitU VO9 1 6-8 ! to12-13 Ov 8-10 7-10 5. 00 present silica Table U) o O concentric _ - N/« <-/i 4k pm)s' (in length00 j- g I cells. S w •£ -=S memberf 370 - 345 in _ 370-- - U) - O -- c O 440-515 390-460 530-550 — 4k U) Lfi cb: O occasionally Ui N/l N/i O

o sq. à\ mm)s' =r 00 5O, >» 2 S cr o (per frequency£ 800 10 —: 4—6 7-111 8-101 chambered ~o OO

i "a vasicentric; 3 J in— present; v: ac: Prototrophis TrophisTrophis a- +: 1- 1 cells; C mexicana 3 ■s - Calpidochlamys SpeciesSpecies 0 Maillardia Trophis TO T. e O' T. Olmedia paratracheal; Subgenusc SubgenusSubgenus Section SectionSection Section Olmedia Legends: chyma C/Î lAWA Bulletin n.s., Vol. 7 (2), 1986 109

the radial walls 4 lumina 7 strands of 4 cells. Rhombic walls; /am, pm, occa- Parenchyma crys-

1525 sionally gelatinous; length (am; F/V-ratio tals abundant in the rays, in nearly all square

4.1. Rays uniseriate and multiseriate, hetero- and upright cells (in some upright cells more geneous type 11, 6 per mm. Uniseriate rays 20 than one crystal), less frequent in the procum- of %, composed square and upright and some bent cells, and very common in the axial paren- procumbent cells, height up to 6 cells (240 chyma cells, the cells often chambered, up to

Multiseriate of pm). rays composed procum- 4—lo crystals in longitudinalrows. Radial latex bent cells except for the uniseriate margins of tubes few, diameter 25—50 pm, much larger

1 of than the cells —4(—7) rows square and/or upright cells; surrounding ray as seen in tan-

4—5 cells wide, up to 960 pm high. Parenchy- gentialsection, in one sample. ma vasicentric and predominantly as concen- tric, slightly wavy bands, 3—6(-9) cells wide, Trophis subgenus Trophis sect. Maillardia (Frap-

2 bands per mm. Parenchyma strands of pier & Duchartre) Corner (Table 5; Figs. 2,

3 cells. Rhombic crystals abundant in the 50, 51) rays, present in nearly all square and upright This section comprises two closely related cells, also in some procumbent ray cells, and species of trees up to 15 m tall (Berg, 1977a), very common in the axial parenchyma cells. distributed in Madagascar, Reunion, and Alda-

bra and the Comoro Islands, and is found in

and from Trophis subgenus Trophis sect. Calpidochlamys dry wet forests, sea level up to 1500

(Diels) Corner (Table 5; Figs. 2, 48, 49) m altitude.

The section comprises three species, distri- Material studied: ‘Maillardia borboni- buted in Malaysia, Philippines and Indonesia. ca (this name is used as Comer did not publish

Trees up to 30 m high in primary and second- a new combination in accordance with the stat. ary forests, in the lowlands and up to 1500 m Nov. ofMaillardia in Trophis). Reunion: T. Cadet altitude. 6310 (Uw 27392).

Material studied: T. philippinensis(Bu- General features; Growth rings absent; reau) Comer. Indonesia, Irian Jaya: BW 11922 sapwood white to yellow. Texture fine, grain

(Uw 18173), Fokkinga 1871 (Uw 24355). straight. Specific gravity 840 N per cubic metre.

General features: Growth rings absent; Microscopic features: Vessels diffuse, heartwood brown to dark brown, not well de- solitary (40%) and in short radial multiples and fined from the Tex- clusters of golden yellow sapwood. irregular 2-5; 10 per sq.mm, round ture coarse, grain straight. Specific gravity to oval, diameter 85 pm, vessel member length

metre. 530-600 N per cubic 345 pm. Perforations simple, end walls almost features: Microscopic Vessels diffuse, transverse. Intervascular pits alternate, round, solitary (35—69%) and in short radial multiples oval or polygonal, 6—B pm. Vessel-ray and ves- and/or irregular clusters of 2-4; 4—6 per sel-parenchyma pits larger, elongated and irre- round to oval, diameter 170—205 sq.mm, pm, gularlyshaped, half-bordered, the borders occa- vessel member 440—515 Perfora- length pm. sionally reduced. Fibres nonseptate, with small tions simple, end walls almost transverse. Inter- simple pits restricted to the radial walls; walls vascular 2—4 pits alternate, round,oval or polygonal, pm, lumina 7-11 pm, occasionally gela- 12—13 and pm. Vessel-ray vessel-parenchyma tinous; length 1165 pm; F/V-ratio 3.4. Rays pits larger, elongated and irregularly shaped, uniseriate and multiseriate, heterogeneous type half-bordered, the borders B—lo Uniseriate occasionally very 11, per mm. rays 50%, com- reduced. Thin-walled tyloses present. Fibres posed of upright cells, and few procumbent

to nonseptate, with small simple pits restricted cells, height up to 13 cells (480 pm). Multiseri- the radial walls 2—4 lumina 16—20 of cells walls; pm, ate rays composed procumbent except 920-1195 F/V-ratio pm; length pm; 2.1-2.3. for the uniseriate margins of I—3 (—7) rows of

Rays uniseriate and multiseriate,heterogeneous and/or 3—4 cells square upright cells; wide, up II (I), s—B type per mm. Uniseriate rays 15—25 to 720 pm high. Parenchyma scanty paratra- of %, composed square, upright and some pro- cheal, only occasionally completely vasicentric, cumbent cells, height to 10—15 cells (240 up predominantly in narrow regular concentric

480 pm). Multiseriate composed of 2—5 cells wide 4—6 bands rays pro- bands, per mm. Pa- cumbent cells for the except uniseriate margins renchyma strands of 4 cells. Rhombic crystals of I—s (—10) rows of and/or scattered in and square upright present, square upright ray 3—5 cells 840—1200 cells; wide, up to pm high; cells, less frequent in the axial parenchyma occasionally with some sheath cells. Parenchy- cells.

vasicentric cell ma scanty paratracheal to (one Note: The high value for the percentage of wide), to slightly aliform with minimal wings, uniseriate rays might be due to the limited and in bands, 3—5 cells wide, 0.1 1 per mm. diameter of the sample. 110 lAWA Bulletin n.s., Vol. 7 (2), 1986

Trophis subgenus Trophis sect. Trophis (Table up to 20 m high, in humid forests, up to 1500

5; Figs. 2,46, 47) m altitude.

from Material O. Ruiz & The single species T. racemosa occurs studied: aspera

Central America and the West Indies to Colom- Pavon, Panama: Dressier 5196 (Uw 21743);

Welle 2954 bia and Peru. Trees up to 18 m high, in humid Ecuador: Maas, Berg & ter (Uw

forests, from sea level up to 1000(—2500) m 23576), Berg & Akkermans 1047 (Uw 27038); altitude. Peru; Williams 7082 (Uw 18429), 6019 (Uw

Material studied: T. racemosa (L.)Urb. 18430).

Mexico, Los Tuxtlas: R. C. Trigos 306 (Uw General features: Growth rings absent;

25690); Honduras: Record & Kuylen 40 (Uw whitish yellow, no demarcation between sap-

26145); Venezuela, Merida: Breteler4s73 (Uw wood and heartwood. Texture medium, grain

550—700 N 12205);Peru: ex FPAw 25331 (Uw 24365). straight. Specific gravity per cubic

General features: Growth rings absent; metre.

heartwood brown, sapwood yellow. Texture Microscopic features: Vessels diffuse,

medium, grain interlocked. Specific gravity solitary (50—70%) and in radial multiples and

metre. of B—lo 700—850 N per cubic irregular clusters 2—4; per sq.mm, features; Vessels round 95-105 vessel Microscopic diffuse, to oval, diameter pm,

solitary (20-50%) and in short radial multiples member length 530—550pm. Perforations sim-

and irregular clusters of 3—6; 7—ll per sq.mm, ple, end walls slightly oblique to almost trans- round to oval, occasionally angular, diameter verse. Intervascular pits alternate, round, oval

95—135 vessel member 390—460 and 7—lo and vessel- pm, length polygonal, pm. Vessel-ray

Perforations end walls almost often and ir- pm. simple, trans- parenchyma pits larger, elongated

verse to slightly oblique. Intervascular pits al- regularly shaped, half-bordered, the borders

ternate, round to polygonal, B—lo pm. Vessel- very reduced. Thin-walled tyloses common. Fi-

and bres and in the ray vessel-parenchyma pits larger,elongated, septate, or septate nonseptate

and irregularly shaped, half-bordered, the bor- same sample, with small simple pits restricted

ders sometimes reduced. Thin-walled tyloses to the radial walls; walls 2—3 pm, lumina 12—

common. Fibres nonseptate, with small simple 20 pm; often gelatinous; length 1250-1450

restricted the radial walls 2—4 2.3—2.7. uniseriate and mul- pits to walls; pm;F/V-ratio Rays

lumina 4—9 6—7 pm, pm, occasionally gelatinous, tiseriate, heterogeneous type 11, per mm.

length 1380—1495 pm; F/V-ratio 2.5—3.2. Uniseriate rays 5—20%, composed of square,

uniseriate and and few Rays multiseriate,heterogeneous upright procumbent cells, height up to

4—B Uniseriate 5—9 cells Multiseriate type 11, per mm. rays 5-30%, (240-360pm). rays com-

of and few of cells for the uni- composed square, upright procum- posed procumbent except

bent cells, height up to B—l2 cells (360—420 seriate margins of I—4 (—7) rows of square of 2-4 pm). Multiseriate rays composed procum- and/or upright cells, cells wide, heightup

to bent cells except for the uniseriate margins of 840-1320 pm. Sheath cells common. Paren-

of and/or l-4(—6) rows square upright cells; chyma scanty paratracheal, only occasionally

3—6 cells wide, up to 840—1320 pm high. Pa- vasicentric as a complete ring of one cell wide,

renchyma in complete vasicentric rings, and predominantly as more or less regular concen-

3—6 predominantly as concentric, cells wide, tric bands, 3—12 cells wide, 2—4 bands per mm, 4—5 bands and often slightly wavy bands, per mm, including the vessels. Parenchyma strands

some 1 cell wide marginal bands. Parenchyma of 3—4 cells. Vitreous silica common in the

strands of 3-8 cells. Rhombic crystals mostly vessels. Radial latex tubes present, infrequent,

abundant in the rays, in nearly all square and diameter 10—15 pm, equal to the diameter of

also in and the cells in upright cells, procumbent ray cells, surrounding ray as seen tangential

in the axial section. very common parenchyma.

Note: T. racemosa from Venezuela has Note:Mennega and Lanzing-Vinkenborg

much less abundant rhombic in the crystals ray (1977) described Olmedia as having nonseptate

and parenchyma cells than the other samples. fibres (exceptionally one or two septate fibres

seen in macerations). However, in two samples

Olmedia Ruiz & Pavon (Table 5; Figs, 52—54) collected more recently in Ecuador, the fibres

A of Olmedia of addition- monograph was published by are definitivelyseptate. Preparation

Berg (1972). He assigned only one species to al sections of the material studied by Mennega

this genus. More recently the same author and Lanzing-Vinkenborg revealed the presence

(1983) included the genus in Trophis 5.1., how- of septate fibres in these samples too, although

ever without making new combinations. The the majority of the fibres is nonseptate.

genus Olmedia is distributed from Costa Rica

the Andes Bolivia. Trees shrubs continued through to or (text on page 124) lAWA Bulletin n.s., Vol, 7 (2), 1986 111

Fig. 3. Broussonetia sect. Allaeanthus, B. greveana, Uw 26758, Madagascar. Scale line valid for all

sect. photomicrographs, unless otherwise indicated. — Fig. 4. Broussonetia Broussonetia, B. kazi-

noki, Uw 26765, Japan. — Fig. 5. Ibid., B. papyrifera, Uw 24338, Thailand. — Fig. 6. Ibid., ibid.,

UNw 213, the Netherlands. 112 IAWA Bulletin n.s., Vol. 7 (2), 1986

Fig. 7. Broussonetia sect. Broussonetia, B. kazinoki, Uw 26979. — Fig. 8. Ibid., B. papyrifera, Uw

24338. — Fig. 9. Maclura sect. Cardiogyne, M. africana, Uw 27394. — Fig. 10. Maclura sect. Chloro- phora. M. tinctoria, Uw 25792. lAW A Bulletin n.s., Vol. 7 (2), 1986 113

Fig. 11. Maclura sect. Chlorophora, M. tinctoria, Uw 25792, Mexico. — Fig. 12. Ibid., ibid., Uw

18419, Venezuela. — Fig. 13. Maclura sect. Cudrania, M. cochinchinensis, Uw 18521, Australia. —

Fig. 14. Ibid.,ibid., Uw 25798, Indonesia. 114 IAWA Bulletin n.s., Vol. 7 (2), 1986

Fig. 15. Maclura sect. Cudrania, M. tricuspidata, Uw 18397, U.K., cult. - Fig. 16, Ibid., M. cochin- chinensis, Uw 18396. — Fig. 17. Maclura sect. Maclura, M. pomifera, Uw 7227, U.S.A. — Fig. 18.

Ibid., M. brasiliensis, Uw 26767 (upper part; stem periphery; lower part: juvenile wood near pith). lAWA Bulletin Vol. 7 1986 n.s., (2), 115

Fig. 19. Maclura sect. Maclura, M. brasiliensis, Uw 26767;rhombic crystals in chambered parenchy-

ma cells (left), rays and vessel (right). — Fig. 20. Plecospermum spinosum, Uw 18400. — Fig. 21.

Ibid., Uw 18400. — Fig. 22. Malaisia scandens, Uw 24362. 116 lAWA Bulletin n.s., Vol, 7 (2), 1986

Fig. 23. Milicia excelsa, Uw 15648. — Fig. 24. , Uw 309, the Netherlands. — Fig. 25. - Ibid., Uw 25505, South Africa. Fig. 26. Ibid., Uw 25787, India, Himalaya. lAWA Bulletin n.s., Vol. 7 (2), 1986 117

Morus Uw Fig. 27. rubra, 7336, U.S.A. — Fig. 28. , Uw 25788, Lebanon. — Fig. 29.

Morus insignis, Uw 24976, Colombia,Andes. — Fig. 30. Ibid.,Uw 24976. lAWA Bulletin Vol. 7 1986 118 n.s., (2),

— - Fig. 31. Morus macroura, Uw 24426, Java. Fig. 32. Morus mesozygia, Uw 18399, Nigeria. Fig

33. Pachytrophe dimepate, Uw 24257. - Fig. 34. Ibid., Uw 26144. lAWA Bulletin n.s., Vol. 7 (2), 1986 119

Fig. 35. Streblus sect. Paratrophis, S. glaber, Uw 20480. — Fig. 36. Ibid., ibid., Uw 24237. — Fig. - 37. Ibid., S. pendulinus. Uw 24387. Fig. 38. Ibid., ibid., Uw 24388. 120 lAWA Bulletin n.s., Vol. 7 (2), 1986

Fig. 39. Streblus sect. Sloetia, S. elongatus, Uw 24396. — Fig. 40. Ibid., ibid., Uw 24396: laticifer in

— — 24439: ray. Fig. 41. Streblus sect. Streblus. S. asper. Uw 24428. Fig. 42. Ibid., ibid., Uw silica

in the vessels arrowed. lAWA Bulletin n.s., Vol. 7 (2), 1986 121

Fig. 43. Sloetiopsis usambarensis, Uw 24445. — Fig. 44. Ibid., Uw 24236: crystals in the tyloses of the vessels, in transverse and tangential section. — Fig. 45. Trophis subg. Prototrophis, T. mexicana,

Uw 18029. — Fig. 46. Trophis subg. Trophis sect. Trophis, T. racemosa, Uw 25690. 122 IAWA Bulletin n.s., Vol. 7 (2), 1986

Fig. 47. Trophis subg. Trophis sect. Trophis, T. racemosa, Uw 25690. - Fig. 48. Ibid. sect. Calpido- chlamys, T. philippinensis, Uw 18173. — Fig. 49. Ibid., ibid., Uw 18173. — Fig. 50. Trophis subg.

Trophis sect. Maillardia, T. borbonica, Uw 27392. lAWA Bulletin Vol. 7 1986 n.s., (2), 123

51. T. Uw 27392. — 52. Olmedia Fig. Trophis subg. Trophis sect. Maillardia, borbonica, Fig. aspera, - Uw 23576. Fig. 53. Ibid.,Uw 21743. - Fig. 54. Ibid., Uw 23576: septate fibres. 124 lAWA Bulletin n.s., Vol. 7 (2), 1986

Discussion ings distinguish the temperate and subtropical

from the tropical species. Element lengths of

Ecology and wood anatomy tropical species are higher than those of tem-

The majority of the taxa of Moreae sensu perate species.

distributed Berg with urticaceous stamens is in tropical lowland vegetations with moderate to Morus The species and samples available

abundant water supplies. However, Morus, for this study are from temperate, tropical low-

montane For Broussonetia and Maclura are represented in land and tropical vegetations.

wood anatomical data Table 3 tropical and temperate areas; Plecospermum is per species, see found in rather dry habitats of India and Sri and Figure 1.

Lanka, and the two species of Streblus sect. The vessel member length in the tropical

and is and the fibre Paratrophis occur in tropical lowland trop- specimens slightly higher length

ical montane forests, respectively. considerably higher than in temperate species.

Within the tropical specimens the vessel mem-

Broussonetia Species of Broussonetia are ber length is slightly lower in the montane taxa.

found in temperate and tropical areas of Asia Morus nigra, with a natural distribution in the

and Madagascar. The section Allaeanthus (3 dry eastern, Mediterranean (subtropical) areas,

species) is tropical; the section Broussonetia (4 falls within the range of the tropical taxa. The

studied from Lebanon and the species) is mainly temperate. Only B. papyri- samples are Netherlands. fera occurs in the tropics, as well as in temper-

A vessel distribution is observed ate areas. As can be seen in Table 1 and Fig. 1, ring-porous

in general the element length in the temperate in the specimens from temperate areas. Here,

species is shorter than in the tropical species, also spiral thickenings occur. The tropical low-

and the vessel diameter is slightly smaller in the land species, M. mesozygia, is diffuse-porous

temperate species, thus confirming the trends and lacks spiral thickenings. Morus macroura,

established by Van der Graaff and Baas (1974). from tropical montane areas, is semi-ring-po-

However, for both trends exceptions exist in rous with faint spiral thickenings. However,

our material. M. insignis is diffuse-porous and lacks spiral

The most striking difference is found in the thickenings, although it is a tropical montane

porosity patterns: ring-porous in the temperate species too.

the all ofM. alba show species, and diffuse-porous in tropical spe- Finally, samples a (semi-)

cies, with the exception of one sample of B. ring-porous vessel distribution and spiral thick- papyrifera from Indonesia which shows a semi- enings in the narrow vessel elements. The origin

ring-porous vessel distribution. As no informa- of the samples is temperate (the Netherlands),

tion on habitat of this sample is available, this subtropical/temperate (South Africa) and trop-

cannot be explained. ical montane (India), respectively.

Spiral thickenings occur in the smaller ves- In conclusion, the features of (semi-)ring-

sels of all samples with a ring-porous vessel dis- porosity and spiral thickenings are linked with-

in tribution, and also in the diffuse-porous sample in the genus, and this combination occurs

of B. papyrifera from Thailand. Thus spiral the temperate and tropical montane areas. The

is in thickenings occur in all samples of the section only exception found M. insignis, occur-

in high tropical montane areas (approx. Broussonetia, and never in those of the section ring 3000 with diffuse vessels without Allaeanthus. Spiral thickenings are commonly m altitude)

regarded as an ecological adaptation and are spiral thickenings.

crite- The distribution is therefore not considered as a taxonomic parenchyma strongly

rion. In Broussonetia, however, it seems pos- dominated by the vessel distribution. The wide

sible to recognise both sections with the aid of earlywood vessels in the temperate species are

this feature. included in broad bands ofmarginal(i.e., initial)

The parenchyma distribution is strongly parenchyma, the small latewood vessels show

dominated by the vessel distribution. In ring- vasicentric to slightly aliform parenchyma. In

porous samples the parenchyma is scanty para- the tropical montane taxa the marginal bands

tracheal with terminal or initial bands, and in are less well developed, and the paratracheal

the diffuse-porous samples aliform-confluent, and parenchyma is more confluent. Finally, in

sometimes with short bands. Broussonetia kurzii tropical lowland species marginal parenchyma

from Thailand shows only vasicentric paren- is faint, and wavy bands are dominating the pa-

chyma. renchyma distribution.

condi- Maclura —As can be seen in Table 2, in Plecospermum A semi-ring-porous

Maclura again ring-porosity and spiral thicken- tion is also found in Plecospermum, a tropical lAWA Bulletin n.s., Vol. 7 (2), 1986 125

taxon of rather dry habitat with seasonal rain- Berg transferred the species of Maclura sect.

to fall. As in the species of Maclura, Morus and Chlorophora (except M. tinctoria) the genus

Broussonetia with a ring-porous vessel distri- Milicia. In his description of Broussonetia gre- bution, broad bands of marginal (i.e., initial) veana (1977a) he mentions the resemblance be- parenchyma occur. Spiral thickenings were not tween this species and Chlorophora (Milicia)

observed. excelsa. Wood anatomically, both species are

rather similar, but in Milicia the parenchyma

— Streblus sect. bands. Streblus The two species of pattern is more confluent and in wavy

Paratrophis differ in their habitat and habit; The resemblance between Broussonetia greve-

to wood S. glaber(trees, up 40 m tall) occurs in mon- ana and B. luzonica is very high in all

tane forests (700—2500 m), and S. pendulinus anatomical features. Capuron’s decision to as-

(shrubs or small trees, up to 13 m tall) is found sign these species to the same taxon (the genus

in lowland forests and on sea shores, often un- Allaeanthus) is thus fully supported.

der seasonal conditions. The differences in wood A comparison of the wood anatomical data

anatomy (see Table 4 and Fig. 2) are substantial, ofsection Broussonetia and section Allaeanthus differences of and are not easily explained by in (Table 1) reveals that the occurrence spiral

ecology. Therefore the broad genus concept of thickenings can be used to distinguishbetween

two Streblus sensu Comer is contradicted (TerWelle, the sections; spirals occur only in section

1985).See further under‘Generic delimitation’. Broussonetia. The fact that this difference can-

not be fully explained by ecological differences

Habit and wood anatomy (see above) might be an indication of some

Among species studied some lianas or clim- taxonomic value of this feature. At first view,

to bers occur, viz. Broussonetia kurzii, Maclura the two sections seem differ in their vessel

cochinchinensis, M. brasiliensis, Plecospermum distribution and in parenchyma distribution,

and Malaisia. too. However, these features are in part related

A high percentage of vessel tissue through to geographical and ecological factors. It may

large and/or numerous vessels is a well known be concluded that the decision to include the

characteristic of lianas. This pattern is found in former genus Allaeanthus as a separate section

Maclura brasiliensis, Plecospermum, and some in Broussonetia is, at least, not contradicted by

of the specimens of Maclura cochinchinensis. the wood anatomical differences.

Along with large vessels, many narrower vessels

in radial chains clusters of 10 Maclura Comer included four or irregular up to (1962) genera

In the Maclura: Cudrania pores generally occur. temperate speci- in Cardiogyne, Chlorophora,

mens of Maclura cochinchinensis, Malaisia and and Maclura. Berg (1977a) modified this con-

the not Broussonetia kurzii, vessel pattern is cept. He included also Plecospermum, but as-

liana-like. Maclura cochinchinensis and Malaisia signed the two African species of Chlorophora

usually start as small shrubs, finally becoming to Milicia (1982), following the conclusions

lianas. It is assumed that the samples studied of Kloos (1982) who reported considerable dif-

had not reached the stage of liana when they ferences in the leaf anatomy between the Afri-

were collected. The same conclusion can be can and Neotropicalspecies.

drawn for Broussonetia kurzii. In the outermost Maclura sect. Chlorophora and Milicia: As in

part of the small sample the first radial chains macromorphological and leaf anatomical attri-

of small vessels can be observed, as well as the butes, the wood anatomy reveals a number of

druses, A mixture of druses and rhombic crys- differences between Milicia and Maclura tinc-

tals at the moment the habit becomes liana-like toria (see Table 6). Thus, the distinction of two

taxa taxa has also been described for some in the in the former genus Chlorophora is sup-

Urticaceae (Bonsen & ter Welle, 1984). ported. Maclura, Cardiogyne, Chlorophora and Cu-

Generic delimitation drania: These four sections of Maclura have

characters in Most many common. striking is As Broussonetia already pointed out in the occurrence of chambered crystalliferous

the generic description, several suggestions for parenchyma strands, with up to 16 crystals in

the taxonomic position of B. greveana have a longitudinal row. In the section Cardiogyne

been made, i.e., treatment as: a species of the this feature is less well developed. Crystalli-

genus Chlorophora (Leandri, 1948); a species ferous parenchyma strands are very scarce in

of the section Chlorophorain Maclura (Comer, the Moraceae. Besides the taxa already men-

1962); a species of the genus Allaeanthus (Ca- tioned, they were observed in Plecospermum,

puron, 1972); a species of section Allaeanthus in Streblus sections Paratrophis, Phyllochla-

sect. in Broussonetia (Berg, 1977a). mus and Sloetia, and in Trophis Calpido- 126 lAWA Bulletin n.s., Vol. 7 (2), 1986

Table 6. Some wood anatomical features of Milicia and Maclura sect. Chlorophora.

fibre Species vessel member pit sizesize fibre length ray frequency multiseriate ray crystalcrystal ray typetype

length (in pm) (in(inpm)/xm) (in(inpm)/xm) (per mm) height (in pm) distribution (Kribs)

Milicia 350-400 10-12 1050-1425 4-6 170-400170- 400 III 350-400 10-12 1050-1425 u,au, a III

MacluraMadura tinctoria 250—320250-320 7-8 810—1040 6-9 315-460 ho tinctoria 7—8 810-1040 6—9 315—460 ac, p

axial chambered Legends: Crystals: a = in parenchyma cells; ac = in parenchyma cells; p = in procumbent ray cells;

and — III Kribs ho u = in square uprightray cells. Ray type: I, II, = heterogeneous types according to (1968); =

homogeneous,i.e,, composed ofprocumbent cells only.

chlamys. However, the length of the longi- wood anatomical differences are mainly correl-

tudinal rows in these taxa is often more lim- ated with the geographical distribution and

in ited, even more so than Cardiogyne. The ecological variation (see above). Within this

of Maclura s.l. is Morus be identified ray type homogeneous to oc- genus, mesozygia can easily

casionally heterogeneous type 111, a ray type by its parenchyma distribution in confluent

which is rather uncommon in the Moraceae. and wavy bands.

The section Cardiogyne and Maclura brasi- liensis of section Maclura differ from the Streblus (cf. Table 4 and Fig. 2) Streblus

other taxa in having ray type I or 11. The sensu Corner: The wood anatomical features of of Streblus range of variation of mean fibre lengths and the individual species vary within

vessel element lengths for all sections is nar- small limits, notwithstanding the differences in row when compared with the other Moreae locality and habitat. However, the variation be-

themselves low. the of Streblus Corner studied, and the values are tween species sensu ap-

The basic parenchyma distribution pattern in proaches the full range reported for all other

Maclura s.l. is vasicentric-aliform and in short taxa studied here! Parenchyma distribution,

wavy bands combined with marginal paren- crystal distribution, silica distribution, number

chyma. The marginal bands were not observed of vessels per sq.mm, vessel diameter and size

in section Cardiogyne and Maclura brasiliensis. of the intervascular pits all vary tremendously.

The parenchyma distribution in Chlorophora The wood anatomy does not support the broad

A is usually less wavy-banded, and occasionally genus concept of Streblus sensu Comer.

unilateral. genus rank for most species studied seems more

The four sections of Maclura sensu Comer, justified.

viz., Cudrania, Cardiogyne, Chlorophora and Streblus and Sloetiopsis: Comer (1962)men-

Sec- Maclura have many characters in common. tioned the possibility of including Sloetiopsis Maclura brasiliensis his of Streblus s.l. tion Cardiogyne and deviate in genus concept Berg (pers.

to somewhat, but resemble each other in many comm.) hesitates accept this idea. From a

features. Based on wood anatomical evidence wood anatomical point of view Sloetiopsis is

Maclura s.l. the the Moreae under the genus concept of is supported, most deviating taxon in

with the remark that more material of Cardio- study here. The very small intervascular pits (c. gyne and Maclura brasiliensis should be studied 2 pm) and the frequent narrow concentric apo-

before these two taxa are definitely included in tracheal parenchyma bands suggest an isolated Maclura. position within the Moreae.

Maclura-Plecospermum:Plecospermum on-

ly deviates from Maclura s.l. in the high number Trophis Corner (1962) included two taxa,

this be of then individual of vessels per sq.mm. As may a result until recognised as genera, viz.

the climbing habit of Plecospermum, the sug- Calpidochlamys and Maillardia, in Trophis s.l.

gestion of Berg (1983)to include this taxon in According to Berg (1977a) Calpidochlamys and

Maclura s.l. is fully supported by its wood Maillardia seem to be related, but he hesitates

anatomical features. to include these taxa in Trophis.

Trophis: Comer (1962)assigned T. mexicana

— and Americana to Morus. The circumscription subdivision and T. (= T. racemosa according

of difficult recent to different Morus are (Berg, 1977a). A Burger, 1962) two subgenera.

monograph is not available. The species studied Burger (1962) monographed Trophis (the Neo-

two are from temperate and tropical areas. The tropical species), and concluded that these IAWA Bulletin Vol. 7 1986 n.s., (2), 127

species are closely related. The wood anatomi- References

cal data fully support Burger, notwithstanding

small quantitative differences. Baillon, H. 1895. Histoire physique naturelle et Trophis, Calpidochlamys and Maillardia: The politique de Madagascar 35 (ed. A. Grandi-

wood anatomical differences between these dier). Histoire naturelle des plantes 5. Atlas taxa are given in Table 5. Section Trophis and 3. Paris.

in section Maillardia are similar the concentric Bentham, G. & J.D. Hooker. 1880. ,

parenchyma bands, but differ in vessel diameter, Genera Plantarum. London.

size of the intervascular pits, length of vessel Berg, C.C. 1972. Olmedieae, Brosimeae (Mora-

members and fibres, and number of Flora rays/mm. ceae). Neotropica, Monograph no. 7.

Thus, in our opinion the inclusion ofMaillardia Hafner Publ. Comp,, New York.

in is not — Trophis (Comer, 1962) supported. 1973. Some remarks on the classification

Berg’s suggestion (pers. comm.) to exclude and differentiationof Moraceae. Meded. Bot.

Maillardia seems more justified. In section Cal- Mus. Herb. Rijksuniversiteit Utrecht 386;

pidochlamys, the parenchyma bands are much 1-10.

less and Other — dominating more widely spaced. 1977a. Revisions of African Moraceae (ex- differences found the of are in number vessels, cludingDorstenia, Ficus, Musanga and Myri- the size of the intervascular pits, the fibre anthus). Bull. Jard, Bot. Nat. Belg. 47: 267- length, and the crystal distribution,especially 407.

the of — presence crystalliferous strands in Calpi- 1977b. The Castilleae, a tribe of the Mora-

Consequently, Berg’s idea renamed and redefined dochlamys. (pers. ceae, due to the ex- comm.) to exclude this taxon from Trophis clusion of the type genus Olmedia from the

5.1., too, seems justified. According to Berg ‘Olmedieae’. Acta Bot. Neerl. 26: 73—82.

Maillardia and — (1977a) Calpidochlamys are 1982. The reinstatement ofthe genus Milicia related. probably closely The wood anatomical Sim (Moraceae). Bull. Jard. Bot. Nat. Belg. differences between these two taxa are as sub- 52: 225-229.

stantial between them and — as section Trophis/ 1983. Dispersal and distribution in the Ur-

subgenus respectively. - Prototrophis, ticales an outline. In; Dispersal and distri- and Olmedia: Trophis Berg (1983) included bution (ed. K. Kubitzki). Publ. Comp. Paul Olmedia in Trophis s.l. Comer (1962) assigned Parey, Hamburg, Berlin.

Olmedia the tribe to Olmedieae, by Berg re- Bonsen, K.J. & B.J.H. ter Welle. 1984. System- named the tribe Castilleae after as (1977b) the atic wood anatomy and affinities of the Ur-

exclusion of Olmedia. Olmedia differs from ticaceae. Bot. Jahrb. 105: 49—71.

Trophis in the occurrence of septate W.C. 1962. Studies fibres, Burger, in new world Mora- radial latex tubes and vitreous silica in the ves- ceae: Trophis, Clarisia, Acanthinophyllum, sels. In the rhombic in the addition, crystals Ann. Missouri Bot. Card. 49: 1—34.

rays and parenchyma cells of Trophis are lacking Capuron, R. 1972. Contribution a 1’etude de la in Olmedia and the vessel members are longer. flore forestiere de Madagascar. Adansonia

Olmedia should not Thus, be included in ser. 2, 12: 375-388.

Trophis. Corner, E.J.H. 1962. The classification of Mo-

raceae. Card. Bull. Sing. 19: 187-251. Malaisia and The wood Pachytrophe ana- Engler, A. 1888. Moraceae. In: Die naturlichen

tomical features of Malaisia and Pachytrophe Pflanzenfamilien(ed. A. Engler& K.Prantl). each vary within narrow limits. These genera Engelmann, Leipzig.

therefore seem to be well defined. Graaff, N.A. van der & P. Baas. 1974. Wood

anatomical variation in relation to latitude Taxonomy of the Moreae and altitude. Blumea 22: 101—121.

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A with the comparison other of the — 1973. genera Description and taxonomic position tribe (those without urticaceous is of stamens) es- Madura brasiliensis (Moraceae). Acta the sential, as in literature several classifications Bot. Neerl. 22; 69-74, have been proposed where genera of both A. 1982. Kloos, Multidisciplinary systematic re-

groups have been treated as close allies. We will search in Moreae - Leaf (Moraceae) anatomy return to this in point a subsequent paper (Ter of the Maclura-group. Acta Bot. Neerl. 31: Welle et al., in preparation). 145. 128 IAWA Bulletin n.s., Vol. 7 (2), 1986

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Kribs, D. A. 1968. Commercial foreign woods 26: 1-27.

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‘Artocarpoideae’ de Madagascar, Not. Syst.

13; 171-181.