lAWA Bulletin Vol. 7 1986 n.s., (3), 175

The systematic wood anatomy of the (Uriticales). V. Genera of the tribe Moreae without urticaceous stamens*

by

B.J.H. ter Welle J. Koek-Noorman and S.M.C. Topper

Institute ofSystematic Botany, University of Utrecht, Heidelberglaan2, 3508 TC Utrecht,

The Netherlands

Summary

The wood anatomy of the Moreae without based upon these characters, however (Berg,

urticaceous stamens is described in detail. Ge- 1983), is not in accordance with the tribes

neric descriptions of the following genera are Moreae and sensu Corner (1962). provided: , , , Ba- Both Berg’s and Corner’s subdivisions deviate

tocarpus, , Parartocarpus, , from older classifications, as given by, for in-

Prainea, , Sparattosyce, and . stance, Bentham and Hooker (1880) and Engler Wood anatomical variation below the (1888). level is very limited, except in the genus Clari- The Moreae characterised by the absence of

sia. Intergeneric variation, however, is much urticaceous stamens comprise the genera Antia-

more evident. Most genera can be recognised ropsis (New Guinea), Artocarpus (Southeast by the presence or absence of septate fibres, Asia), Bagassa (Neotropics), (Neo- and of radial latex tubes, the size of the inter- tropics), Clarisia (Ncotropics), Huliettia (South-

vascular pits, the parenchyma distribution,and east Asia),Parartocarpus (Southeast Asia), Poul- crystal distribution. The diagnostic and taxon- senia (Neotropics), Prainea (Malesia), Sorocea

omic value of several characters is discussed. (Neotropics), Sparattosyce (New Caledonia),

Key words: Moraceae, Moreae, systematic wood and Treculia (Tropical Africa).

anatomy

Methods and Materials

Introduction The methods employed are those given in

This is of in which the the first of this paper part a series, paper series (Koek-Noorman et

wood anatomy of the Moraceae is described al., 1984). All wood samples studied are backed

and discussed in relation to the of by herbarium vouchers. Taxa of the African the The family. general outline of this study is and Neotropical genera were identified or cited provided in the first paper (Koek-Noorman et by the authors of the various monographs. De-

In the fourth of the the al., 1984). paper series tails on individual wood samples are provided

wood anatomy of those genera of the tribe at the beginning of each generic description.

Moreae (sensu Berg) which are characterised by The genus Hullettia is not represented in this

urticaceous stamens described was (Ter Welle study because wood samples were not at our

et al., 1986). In this paper the genera without disposal.

this of stamens dealt with. In addition type are The data on the specific gravity are based on

to the structure of the stamens, the following the samples cited and complemented with data

features be used to characterise this can group on other samplesofthe Utrecht wood collection

of genera in Moreae according to Berg (1983); not further included in the anatomical study.

inflorescences rather to simple complex ;macro- Quantitative data generally refer to average

spermous; diaspores dry or mostly in fleshy values, unless specified otherwise.

structures, mostly zoochorous; mostly in rain- forest habitats; taxonomically and geographi-

cally segregated, pantropical, but chiefly in the Generic descriptions Indo-Malesian region.

The tribe Moreae sensu Berg (1983) com- Antiaropsis Schumann (Figs. 1, 2) prises all genera of the tribes Artocarpeae and A genus with one or two species (Comer,

Moreae sensu Comer (1962), and also a few 1962). The genus is probably restricted to Irian

to genera assigned the tribe Olmedieae by Jaya (Indonesia). Its occurrence in Papua New

Comer (l.c.). The subdivision in two groups Guinea is questioned by Comer (1962). The

* This project was made possible by a grant of BION-ZWO (14.45-01). 176 IAWA Bulletin n.s., Vol. 7 (3), 1986

shrubs or small trees, up to 11 m high, are found 22031b (Uw 24421). - A. fretissii Teijsm. &

in lowland forests 1000 usual- Binn. ex Hassk. Irian BW 9871 up to m altitude, Indonesia, Jaya: ly in the undergrowth. (Uw 18112), BW 2133 (Uw 24316), BW 2512

Material studied : A. decipiens Schu- (Uw 24317), BW 2175 (Uw 24318). A. hor-

mann. Indonesia, Irian Jaya: CS1RO H 5198 ridus Jarrett. Indonesia, Irian Jaya; BW 11593

(Uw 25533). (Uw 20479). - A. integer (Thunb.) Merr, Indo-

General features: Growth rings absent; nesia, Irian Jaya; BW 5659 (Uw 24319); Jamai-

heartwood not available, sapwood light brown. ca (cult.): US Nat. Herb. (Uw 8294); South

Texture fine, grain straight. Specific gravity c. Africa (cult.): Baijnath s.n. (Uw 25507). — A.

750 N cubic kemando FMS 28853, ex FHOw per metre. Miq, Microscopic features: Vessels diffuse, 4536 (Uw 24608); Indonesia: RTIw IND. Col.

solitary (50%) and in short radial multiples of 3590 (Uw 24605), Pfeiffer E755 (Uw 24324).

to — lakoocha Roxb. Pakistan: MADw 24487 2—3; 9 per sq.mm, round slightly oval, diam- A. ex

eter 108 pm, vessel member length 525 pm. (Uw 18033); Thailand: Royal For. Dept. s.n.

Perforations simple, end walls almost transverse. (Uw 24656). — A. lanceifolius Roxb. Indonesia:

Intervascular pits alternate, polygonal, occasion- RTIw IND. Col. 2200 (Uw 24606); RTIw IND.

— nitidus Trecul. allyround, 7—9 pm. Vessel-ray and vessel-paren- Col. 4027 (Uw 24606). A.

chyma pits larger, elongated,half-bordered, the Malaysia, Sabah: Sandakan For. Dept. 25510

borders often reduced. Thin-walled tyloses com- (Uw 24217). — A. sepicanus Diels. Indonesia,

7822 — A. mon, often with vitreous silica. Fibres septate, Irian Jaya: Fokkinga (Uw 24329). with small Jarrett. Sabah: Sandakan with several septa per fibre, simple sericicarpus Malaysia,

pits restricted to the radial walls; walls 3-4pm, For. Dept. 50586 (Uw 24218). - A. teysmannii

lumina 3—7pm; length 1600 pm; F/V-ratio 3.0. Miq. Indonesia, Irian Jaya: BW 1204 (Uw

Rays uniseriate and multiseriate, heterogeneous 24330).

Uniseriate General features; Growth type II/III, 7 per mm. rays 25%, rings absent;

composed of upright cells only, but variable in heartwood light brown to dark brown, often

to 12—24 cells distinct from the to height, heightup (480-800pm). sharply pale yellow light

Multiseriate rays composed of procumbent cells brown sapwood. Texture coarse, grain(weakly) of 280—940 N except for the uniseriate margins 1—5, occa- interlocked. Specific gravity per 20 of sionally up to rows square and/or upright cubic metre.

cells; 3—4 cells wide, up to 960—1400 pm high. Microscopic features: Vessels diffuse,

Occasionally with some sheath cells. Parenchy- solitary (45—80%) and in short radial multiples

concentric and clusters of 2—6 ma scanty paratracheal, and in irregular 2—4; per sq. mm,

cells wide and Paren- round and diameter 160—310 vessel bands, 3-6 4-5 per mm. oval, pm,

chyma strands of 4 cells. A few latex tubes ob- member length 350—630 pm. Perforations sim-

served in the rays. ple, end walls almost transverse. Intervascular pits alternate, round, polygonal and occasion-

Artocarpus J. R. & G. Forster (Figs. 3—6) ally slightly oval, 9-13 pm. Vessel-ray and ves-

The genus has been monographedby Jarrett sel-parenchyma pits larger, half-bordered, the

(1959c,d,e; 1960a,b). About 50 species are borders sometimes reduced. Thin-walled tyloses

The is distributed but absent. Fibres currently recognised. genus common, occasionally non- from India to southern China, Malesia and the septate, with small simple pits restricted to the

altilis radial walls 2—4 lumina 7—25 Solomon Islands. Two species, A. and A. walls; pm, pm,

heterophyllus, are cultivated all over the tropics gelatinous fibres present or absent; length

for their fruits. Trees to large trees, up to 45 m, 1170—1950 pm; F/V-ratio 2.3—3.9. Rays uni-

in evergreen forests and areas with a mild mon- seriate and multiseriate, heterogeneous type

from level 1000 3—6 Uniseriate soon climate, usually sea up to II/III, per mm. rays 3—30%,

m altitude. composed of upright, square and sometimes

Material studied: A. anisophyllus Miq. few procumbent cells, height up to 4—10 cells

Malaysia, Sabah; Sandakan For. Dept. 50567 (160—480 pm). Multiseriate rays composed of

(Uw 24213). — A. blancoi (Elmer) Merr. Philip- procumbent cells, except for the uniseriate

pines, Quezon: J.P. Rojo 262 (Uw 24214). — margins of 1—2 (5) rows of square and/or up-

A. communis J. R. &G. Forster. Indonesia, Irian right cells; 3—5 cells wide, up to 600—1200 pm

Jaya: BW 2904 (Uw 24602), BW 4219 (Uw high. Occasionally few sheath cells present. Pa-

24603). A. dadah Miq. Indonesia: RTIw IND. renchyma aliform with short to long wings, oc-

Col. T889 (Uw 25521), RTIw IND. Col. 14317 casionally confluent, and sporadically in short

(Uw 25522). — A. elasticus Reinw. ex Blume. wavy bands. Parenchyma strands of 3—4 cells.

Malaysia, Sabah: Sandakan Herb. Inst. 18656 Vitreous silica common, in the fibres or in the

(Uw 24215); Indonesia, Java: Koorders 1026c/ vessels, often enveloping the tyloses. Radial lAWA Bulletin n.s., Vol. 7 (3), 1986 177

latex tubes present in most samples, axial latex Batocarpus Karsten (Figs. 9 & 10)

tubes present or absent (cf. Topper & Koek- A genus with four species according to the

Noorman, 1980). monograph of Mello Filho & Emmerich (1968).

The is distributed from genus Costa Rica to

Aublet to Bagassa (Figs. 7, 8) Brazil, Peru and Bolivia. Trees up 20—30 m

In a treatment for the Flora of Suriname, high, in the tropical lowland rainforests.

Berg (1975) distinguished two species. Recent Material studied: B. amazonicus (Ducke)

studies by the same author revealed that the Fosberg.Brazil,Amazonas: Prance & Maas 15741

genus is monotypic, as B. tiliifolia is now con- (Uw 19158); Krukoff 6422 (Uw 7714); Peru:

as sidered a juvenileform ofB. guianensis (Berg, Williams 5334 (Uw 18416). — B. orinocensis

The is distributed in the Karsten. Ecuador: & Akkermans 1079 pers. comm.). genus Berg (Uw

Guianas and adjacent areas of northern Brazil. 27049).

The large trees, up to 45 m high, are of rather General features: Growth rings absent;

infrequent occurrencein the lowland rainforests all samples light greyish or yellowish brown,

and 650 altitude. Williams described the heartwood of savannas, up to m (1936)

Material studied: B. guianensis Aubl. the sample collected by himself (no. 5334) as

Guyana: A.C. Smith 3542 (Uw 21698); Suri- ‘thin, pale or dark brown’. Texture medium to nam: Lanjouw & Lindeman 2433 (Uw 1742); coarse, grainstraight to irregular. Specific gravi-

Brazil: INPA X-6288 540—650 N cubic (Uw23781), Capucho 356, ty per metre.

ex MADw 34591 (Uw 24261). — B. tiliifolia Microscopic features: Vessels diffuse,

Benoist.’. Guyana: F.D. 5827, ex MADw 3198 solitary (40—65%) and in short radial multiples

(Uw Surinam: Stahel 194 (Uw 194); and clusters of 2—5 24263); irregular 2—4; per sq. mm,

French Guiana: BAFOG 1302 round to diameter 125—205 vessel (Uw 5790). oval, pm,

General features: Growth member 405-440 in B. rings poorly length pm (295 pm defined; heartwood dark brown, sharply dis- amazonicus from Peru). Perforations simple,

tinct from the light brown sapwood. Texture end walls almost transverse. Intervascular pits

medium to interlocked. round coarse, grain Specific alternate, or polygonal, 7—10 pm. Vessel- gravity 880—1000 N cubic and per metre. ray vessel-parenchymapits larger, elongated

Microscopic features: Vessels diffuse, and irregular, half-bordered, the borders in part

and in short radial reduced. Thin-walled solitary (20-60%) multiples tyloses occasionally pres- and clusters of 3-4 irregular 2—4; (7) per sq. ent in some samples, always few. Fibres non-

mm, round and oval, diameter 145—240 pm, septate, with small simple pits restricted to the

vessel member 325—445 Perfora- length pm. radial walls; walls 2—5 pm, lumina 7—20 pm,

tions simple, end walls almost transverse. Inter- partly gelatinous (up to 90% of the fibres gela-

vascular pits alternate, round, polygonal or tinous in B. orinocensis ); length 1165-1700

9-13 and slightly oval, (-15) pm. Vessel-ray pm; F/V-ratio 3.9—4.4 in the samples of B.

vessel-parenchyma pits larger, elongated, half- amazonicus, and 2.6 in the sample of B. orino- bordered, borders sometimes reduced. Thin- censis. Rays uniseriate and multiseriate, hetero-

walled Fibres 4—6 tyloses common. nonseptate, geneous type III, per mm. Uniseriate rays

with small simple pits restricted to the radial 6—18%, composed mainly of upright cells and walls; walls 3—5 pm, lumina 5—13 pm, gelatin- a few rows of procumbent cells, the upright

ous fibres in variable 945—1150 cells variable to amounts;length in height, height up 3—6(—8) pm; 2.7—3.0. uniseriate and cells F/V-ratio Rays (145—480 pm). Multiseriate rays com-

multiseriate,heterogeneous type III, sometimes posed of procumbent cells except for the uni- almost homogeneous, 3-7 mm. Uniseriate seriate of 1—2 of per margins rows square and/or rays of and 4—6 0—14%, composed upright, square upright cells; cells wide, up to 1200— often to 4-9 cells 1400 in procumbent cells, height up pm high B. amazonicus (Prance & Maas

(120—360 pm). Multiseriate rays composed of 15741, and Krukoff 6422) and up to 700—800 procumbent cells for mar- except the uniseriate pm in the other two samples. Occasionally of gins 1-2 rows of square and occasionally some sheath cells. Parenchyma scanty paratra- 2-5 upright cells; cells wide, up to 600-840 cheal to vasicentric-aliform with short wings, Sheath cells pm high. very scarce. Parenchyma wings sometimes long and confluent. The dom-

aliform scanty paratracheal to vasicentric, with inating pattern and the vast majority of the

short wings and some diffuse parenchyma. Pa- parenchyma is formed by irregular wavy bands,

strands of 3-4 cells. Rhombic 3—10 cells wide and 2—3 bands Paren- renchyma crys- per mm. tals in the of the less common margins rays, chyma strands of 3—4 cells. Vitreous silica frequent in the axial parenchyma. Radial latex scarcely present in some samples, in vessels tubes present in all samples, axial latex tubes and/or fibres. occasionally (text continued present. onpage 184) 178 IAWA Bulletin n.s., Vol. 7 (3), 1986

Figs. 1—26. Scale-line = 225 µm, unless indicated otherwise.

— seri- Fig. 1. Antiaropsis decipiens, Uw 25533. — Fig. 2. Ibid., septate fibres. Fig. 3. Artocarpus

cicarpus, Uw 24218. — Fig. 4. Artocarpus lakoocha. Uw 18033. lAWA Bulletin Vol. 7 (3), 1986 n.s., 179

5. Fig. Artocarpus Uw radial — sericicarpus, 24218, latex tubes, group 1. Fig. 6. Ibid.,radial latex

tube. — Fig. 7. Bagassa guianensis, Uw 1742. — Fig. 8. Bagassa tiliaefolia,Uw 5790. 180 lAWA Bulletin n.s., Vol. 7 (3), 1986

9. Uw 18416. — 10. Uw 19158. Fig. Batocarpus amazonicus, Fig. Ibid., — Fig. 11. Clarisia biflora.

Uw 25091. — Fig. 12. Clarisia racemosa, Uw 14464. IAWA Bulletin Vol. 7 1986 181 n.s., (3),

— Fig. 13. Clarisia ilicifolia, Uw 7982. Fig. 14. Parartocarpus bracteatus, Uw 24435. — Fig. 15.

venosus Parartocarpus ssp. forbesii, Uw 25807. — Fig. 16. Ibid. 182 lAWA Bulletin n.s., Vol. 7 (3), 1986

Fig. 17. Parartocarpus bracteatus, Uw 24436, radial latex tubes, type 2. — Fig. 18. Parartocarpus venosus ssp. forbesii Uw 25807, homogeneous rays. — Fig. 19. Poulsenia armata, Uw 24638. —

Fig. 20. Ibid., Uw 20696, vitreous silica in the fibres. lAWA Bulletin Vol. 7 n.s., (3), 1986 183

21. Prainea Uw 18124. — Fig. papuana. Fig. 22. Sorocea guilliminiana, Uw 20941. — Fig. 23.

Sparattosyce dioica, Uw 24627. — Fig. 24. Ibid. 184 I AWA Bulletin n.s., Vol. 7 (3), 1986

Fig. 25. Treculia africana, Uw 24640. — Fig. 26. Ibid.,radial latex tubes, type 2,

Note: In Williams 5334(citedby Mello Filho Maguire et al. 56675 (Uw 16474). — C race-

& Emmerich, 1968, as B. amazonicus) rhombic mosa Ruis & Pavon. Venezuela: Breteler 5054

in and and Colombia: de 1056 crystals occur square upright cells, (Uw 12293); Bruijn (Uw

less frequently in the procumbentray cells and 14464), 1553 (Uw 14496);Ecuador: For. Stat.

the (occasionally chambered) axial parenchyma Connocotto s.n. (Uw 23877); Brazil, Amazonas:

cells. In all other specimens crystals are lacking. Krukoff 6327 (Uw 7645), 6628 (Uw 7829);

Loureiro et al., INPA X-5567 (Uw 23714),

Clarisia Ruiz & Pavon (Figs. 11—13; Table 1) General features: Growth rings absent

The genus comprises three species, according or very faint; heartwood absent or indistin-

who included the Acan- from the brown in C. to Berg (1968) genus guishable light sapwood thinophyllum monographed by Burger (1962) biflora and C. ilicifolia., heartwood dark brown

by combiningA. ilicifolia and A. spruceana to and sharply defined from the light brown sap-

Clarisia ilicifolia. The genus is distributed in wood in C. racemosa. Texture medium ((C. ilici-

Central America and the northern part of South folia)to coarse (i(C. biflora, C. racemosa), grain

America. Two species (C. racemosa and C. bi- straight, but interlocked in C racemosa. Specif-

bi- flora) are trees up to 40 m high. Clarisia ilicifolia ic gravity 400-500 N per cubic metre in C.

is a shrub or tree up to 20 m high. All species flora. and 650—850 N per cubic metre in the

occurin tropical lowland forests. other two species.

Material studied; C. biflora Ruiz & Pa- Microscopic features: Vessels diffuse,

von. Colombia, Chocô: Cuatrecasàs 15487 (Uw solitary (25—51%) and in short radial multiples

Peru; 2511 - and clusters of 3-13 25091); Ellenberg (Uw 8712). irregular 2-5; per sq. mm,

C. ilicifolia (Sprengel) Lanj. & Rossb. Guyana: round to slightly oval, diameter 75—200 pm,

For. 4039 Surinam: Lindeman vessel member 310-485 Perfora- Dept. (Uw982); length pm.

6776 (Uw 4587); Daniels & Jonker 1079 (Uw tions simple, end walls almost transverse. Inter-

8593), 1081 (Uw 8595); van Donselaar 1221 vascular pits alternate, round and polygonal,

(Uw 10835); Brazil: Krukoff 6849 (Uw 7982); 6—11 pm. Vessel-ray and vessel-parenchyma lAWA Bulletin n.s., Vol. 7 (3), 1986 185

Table 1. Some wood anatomical characters ofspecies of Clarisia.

!mm) pm)| sq. (in pm) il(per .Si (in rays length pm)

o T (u (inc -2- w’ 1/5 ea (U 2 W5 -2 X) >> _. 3 C crystals•-> 25 c *S C i> C o vessels _ o* -fci. 1 « > frequencyu .2 2 .2 member „ diameter uniseriate S length * ,Sin S ■o g | « S

2 8 8 S o o I > I of'-' t/5 C/5 C/5 o _d F/V-ratioCl 3 « *5 vessel n= TT- -C o fibre „ vessel vessel K U- >>c/}fcvSiO % > <*~ *- COSpecies•a ooo't/5

>

1 -1 4- 1 Cn biflora15Cl 180-200000 to o o OJ3-44^ 11-16i— On 465-485ONcn 00 t-n 1200-1500tooo Ol o o to2.6-3.1On1 OJ

»

S’ + 1 1 1 C.ilicifolian *5*t •o75-1051 o c/i 7-13■O U) 13-22u> tO to U310-320O u> to o VO960-1315Ov0 OJ u>3.8-4.2bo 4^ to (u, p,y a)03

+ C.racemosan r> s o§ 144-2001lOO o 4^4-51 Cfl + _1-31U) 4^.400QO 1285-1300tO 00 t-fl 1 300 OJ3.2-S.3k)1U) U) (p,y a)03

Legend: F/V-ratio: ratio of fibre and vessel member length. — Crystal distribution; u: in square and

in axial cells. upright cells; p: in procubent ray cells, a: parenchyma

pits larger, elongated, and irregularly shaped, Parartocarpus Baillon (Figs. 14-18)

the borders sometimes reduced. The has been Jarrett half-bordered, genus monographedby

Thin-walled tyloses common. Fibres nonsep- (1960c). Two species and four subspecies were

tate, with small simple pits restricted to the ra- recognised by her. A third species was described

dial walls 2—4 lumina 6—20 Comer The is distributed walls; pm, pm, by (1976). genus

gelatinous fibres often present; length 960— from Thailand throughout Malesia and the

1500 pm; F/V-ratio 2.6—4.2. Rays uniseriate Solomon Islands. The medium to large trees,

and III and 45 inhabit lowland multiseriate,heterogeneous type II, up to m high, evergreen

occasionally almost homogeneous, 4-7 per rainforests, extending up to 1800 m altitude.

mm. Uniseriate rays 1 —22%, composed of vari- Material studied: P. bracteatus ( King)

amounts able of procumbent, square and up- Becc. Malaysia: RTIw IND. Col. 24620 (Uw

4—12 cells RTIw IND. Col. 24632 - right cells, height up to (150-360 24435), (Uw 24436).

pm). Multiseriate rays composed of procum- P venenosus (Zoll. & Mor.) Becc. ssp. borneen-

bent cells except for the uniseriate margins of sis (Becc.) Jarrett. Indonesia, Borneo: DFP 5008

of P. 0—2(—5) cells square and/or upright cells; (Uw 24413).- venenosus(Zoll. & Mor.) Becc.

3—6 cells wide, up to 600—1560 pm high. ssp. forbesii (King) Jarrett. Malaysia: F.R.I.

Sheath cells always present, but scarce. Paren- Kepong, ex SJRw 38498 (Uw 24639); Indone-

chyma vasicentric, aliform with short, and oc- sia, Sumatra: Krukoff 4124 a (Uw 25807). P.

casionally long wings, but predominantly band- venenosus (Zoll. & Mor.) Becc. ssp. papuanus

to ed, varying from commonly wavy bands (Becc.) Jarrett. Indonesia, Irian Jaya: BW 12294

sometimes concentric bands, 2—6 (—8) cells (Uw 18185), BW 9422 (Uw 20478), BW 4243

wide and 1—3 (—4) per mm. Terminal bands (Uw 24352); Philippines, Bataan: BF Manila,

occasionally present. Parenchyma strands of ex SJRw 17584 (Uw 24340). P. venenosus

2—4 cells. Rhombic crystals present or absent (Zoll. & Mor.) Becc. ssp. venenosus. Indonesia,

in the rays and/or axial parenchyma, usually Java: Koorders 4273w (Uw 24633).

scarce if present. Vitreous silica common in the General features: Growth rings absent;

vessels, but generally scarce. Radial latex tubes heartwood light yellow-brown,indistinguishable

observed in some samples of C. ilicifolia. from the sapwood. Texture coarse, grain inter-

Note: The wood anatomical difference be- locked. Specific gravity (290—)400-600 N per

tween the three species of Clarisia are consider- cubic metre.

able. For some characters this is illustrated in Microscopic features: Vessels diffuse,

Table 1. solitary (21—60%) and in short radial multiples 186 I AWA Bulletin n.s., Vol. 7 (3), 1986

to or regular clusters of 2—4 (5); I—3 per sq.mm, stricted the radial walls; walls 2—3 /am, lumi-

170—275 vessel 15—35 1410—1415 round to oval, diameter pm, na jam/length pm; F/V-ratio

member length 465—740 pm. Perforations sim- 2.0—2.1. Rays uniseriate and multiseriate, het-

Intervascular 3-5 Uniseriate ple, end walls almost transverse. erogeneous type 111, per mm.

of and pits alternate, polygonal and occasionally rays 3—27%, composed upright square

cells and few round, 14-15 pm. Vessel-ray and vessel-paren- a procumbent cells, height up to

4—7 cells Multiseriate chyma pits larger, elongated and more irregu- (180—310 /am). rays

larly shaped, half-bordered, the borders usually composed of procumbent cells except for the reduced. Thin-walled tyloses very scarce. Fibres uniseriate margins of 1 —2 rows of square and/or

nonseptate, with small simple pits restricted to upright cells; 4—6 cells wide, up to 960—1400

the radial walls; walls 2—4 pm, lumina 15—25 jam high. Occasionally with some sheath cells. pm; length 1370—1810 pm; F/V-ratio 2.2—3.8. Parenchyma vasicentric-aliform with short

Rays uniseriate and multiseriate, homogeneous, wings, occasionally confluent. Parenchyma

strands of 2-8 cells. The distribu- 4—6 per mm. Uniseriate rays usually 1-8%, in parenchyma

some samples up to 30%, composed of pro- tion is not very clear in the transverse sections,

cells 4-14 cells because the lumen thickness ratio cumbent only, height up to diameter/wall

(140—420 pm). Multiseriate rays composed of of the fibres is almost equal to that of the pa-

procumbent cells only; 2—3 (4) cells wide, up renchyma cells. Vitreous silica abundant in the

rather and also in the axial to 600-1200 pm high. Parenchyma fibres, occasionally paren-

abundant, vasicentric-aliform with short and/or chyma and the vessels. Radial latex tubes com-

long wings to confluent, occasionally in short mon, the diameter of the tubes in part equal to

wavy bands. In all samples a tendency towards the diameter of the ray cells, partly larger (as

unilateral abaxial parenchyma is evident. Paren- seen in the tangential section).

chyma strands of 3—4 cells. Radial latex tubes Notes: Going from Mexico, Costa Rica, Pa-

to the diameter of the inter- common and with a large diameter as compared nama, Colombia,

the vascular increases from 6—7 9 At to the surrounding ray cells; as a result, pits to /am.

latex tubes are very conspicuous. Rhombic the same time, the specific gravity decreases

crystals common, but sometimes scarce, in the from 450 to 250 N per cubic metre.

to- axial parenchyma. In about 50% of the samples a tendency

wards anobliquevessel distribution was noticed.

The genus shows a high accumulation ofvitreous

Poulsenia Eggers (Figs. 19, 20) silica, which may exceed 7% (of dry weight)

A distributed from Mexico to Van Slooten monotypic genus according (1968).

to Ecuador and Bolivia. Trees up to 40 m high

Prainea f. in tropical rainforests within the area. King ex Hook. (Fig. 21)

Material studied: P. armata (Miq.) Stand- The genus comprises four species according

to most recent ley. Mexico: Williams 8426, ex SJRw 34588 the monograph(Jarrett, 1959b).

(Uw24624); Trigos 309 (Uw 25689); Costa Rica: The species are confined to Malesia: 3 species

lICA CCO 29 (Uw 20696); Panama; Cooper & in Sumatra, Malaya and Borneo, and 1 in the

New Guinea. Slater 132, ex SJRw 10312 (Uw 24628), Cooper Moluccas and Irian Jay a/ Papua

& Slater SJRw 10269 Co- The 35 found in 88, ex (Uw 24629); trees up to m high are ever- to lombia: Cuatrecasas 14342 (Uw 24638). green rainforests up 1200 m altitude.

General features: Growth rings faint or Material studied: P. frutescens Becc.,

absent; heartwood yellow to creamish and in- Malaysia, Sarawak: CSIRO 32639 (Uw 24363).

distinguishable from the sapwood. Texture me- —P. limpato (Miq.) Beumee ex Heyne. Malesia

dium to coarse, grain interlocked. Specific grav- (?): RTlw IND. Col. 20013 (Uw 24437), RTIw

14316 - P. ity 250—450 N per cubic metre. IND. Col. (Uw 24438). papuana

Microscopic features: Vessels diffuse, Becc. Indonesia, Irian Jaya: BW 11785 (Uw

solitary (35—90%) and in short radial multiples 18124), BW 1559 Fokkinga (Uw 24356), BW

4-10 1664 or irregular clusters of 2-4; per sq.mm, Fokkinga (Uw 24357).

round to slightly oval, diameter 105—210 pm, General features: Growth rings absent;

vessel member length 670-695 pm. Perfora- heartwood dark-brown, sharply defined from

tions simple, end walls almost transverse. Inter- the light-brown sapwood. Texture medium to

vascular pits alternate, round and/or polygonal, coarse, grain slightly interlocked. Specific gravi-

6—9 and ty 600—700 N cubic metre P. pm. Vessel-ray vessel-parenchyma pits per (in limpato

larger, elongated, more irregular-shaped, half- 780—980 N per cubic metre).

bordered, the borders reduced. Thin-walled Microscopic features; Vessels diffuse,

tyloses sometimes present, but always scarce. solitary (50-84%) and in short radial multiples

Fibres nonseptate, with small simple pits re- and irregular clusters of 2-4; 2—4 per sq.mm, lAW A Bulletin n.s., Vol. 7 (3), 1986 187

to round slightly oval, diameter 190-220 jam (Uw 20913); Surinam: Oldenburgeret al. 1402

in P. Uw vessel member S. rhodo- (130 jam limpato, 24438), (Uw 17968). trophoides Burger ssp. 350—450 length jam. Perforations simple, end rachis Cuatrec. Colombia, Choco: Cuatrecasas

walls almost transverse. Intervascular pits alter- 14891 (Uw 25362), 15572, type (Uw 25436).

nate, round and polygonal, 9—12 pm. Vessel- General features: Growth rings absent ray and vessel-parenchymapits larger,elongated, or faint; yellowish to light brown, no differ-

more irregular-shaped, half-bordered, the bor- ence between heartwood and sapwood. Tex-

ders often reduced. Thin-walled tyloses com- ture fine, grain straight, sometimes slightly in-

to mon, varying from abundant scarce. Fibres terlocked. Specific gravity 500-1000 N per all or in part septate, several septa per fibre, cubic metre.

with small simple pits restricted to the radial Microscopic features: Vessels diffuse,

walls 2—5 lumina 7—14 walls; pm, pm, partly solitary (20—63%) and in short radial multiples

gelatinous in P. frutescens;, length 1230-1420 and clusters of 2-4 irregular (5);8-21 permm, 3.1—3.5. uniseriate pm; F/V-ratio Rays and round to slightly oval, diameter 68-130 pm, 4—7 vessel 325—475 multiseriate, heterogeneous type 111, per member length jam. Perfora-

mm. Uniseriate rays 13—20%, composed of up- tion simple, end walls almost transverse. Inter-

and right square cells, only occasionally some vascular pits alternate, round and sometimes procumbent cells, height up to 4—7 cells (240 polygonal, 6—12 jam. Vessel-ray and vessel-

420 Multiseriate of half-border- pm). rays composed pro- parenchyma pits larger, elongated,

cumbent cells except for the uniseriate margins ed, the borders often reduced. Thin-walled ty-

of I—3 of loses rows square and/or upright cells; common. Fibres nonseptate, with small

3-5 cells wide, up to 720-1200 pm (in P. fru- simple pits restricted to the radial cell walls;

to tescens up 480 pm) high. Occasionally with walls 2—4 pm, lumina 4—12 pm, gelatinous

some sheath cells. vasicentric-ali- fibres 1080—1460 Parenchyma common; length pm; F/V- form with short to long wings, occasionally -ratio 2.5—3.9. Rays uniseriate and multiseriate,

confluent or short in (wavy) bands. Parenchy- heterogeneous type 11/111, 6—9 per mm. Uni-

ma strands of 4 cells. Vitreous silica seriate of common rays 7—40%, composed upright and

in the the occasion- and few vessels, enveloping tyloses; square occasionally procumbent cells, ally also in the fibres and sporadically in the height up to 6—19 cells (180—600 pm). Multi- axial parenchyma. seriate rays composed of procumbent cells ex-

cept for the uniseriate margins of 1-3 (10)

Sorocea St. Hil. of (Fig. 22) rows square and/or upright cells; 2—5 cells The genus has recently been monographed wide, up to 660—1200 pm high. Parenchyma by and Berg Akkermans (1985), who recognised scanty paratracheal, but predominantly as con- 16 species. The genus Paraclarisia was already centric, sometimes more or less wavy bands en- included Sorocea in by Burger et al. (1962). closing the vessel in part, sometimes also termi- Sorocea is distributed in the from Neotropics, nal bands present, 2—9 cells wide and 2.7—5 Guatemala and and in to Paraguay Argentina, bands per mm. Parenchyma strands of 3—4

Curacao. The shrubs or trees are to 6—20 m cells. Rhombic in the and up crystals square up-

high, and are generally components of the un- right cells from few to abundant, and less fre-

in non-flooded from to derstory moist, forests, quent occasionally absent in the axial paren- level 2000 altitude. cells. Radial latex sea up to m chyma tubes observed in one

Material studied: S. bonplandii (Bail- sample.

W. & W. Boer, Notes. The Ion) Burger, Lanjouw Brazil, wood anatomy of the genus is Santa Catharina: Reitz 15026 Para- One S. (Uw 6409), very homogeneous. species, muriculata. na: Lindeman & H. de Haas 1988 (Uw 13437), can be identified using the average vessel diam- 2842 S. duckei (Uw 13928).- W. Burger. Brazil, eter, being 68—70 jam in this species, and 90—

Amazonas: F. Mello,INPAX-3745 130 in the other (Uw 23664). pm species. S. faustiniana Cuatrec. Colombia, Choco:

Cuatrecasas 15204 (Uw 24928). S. guillimi- Sparattosyce Bureau (Figs. 23, 24)

Gaudich. Brazil, A niana Amazonas: Prance et al. genus comprising two species, restricted to 18248 19825 (Uw 20902), (Uw 20941), Matto New Caledonia and occurring in moderately Grosso: Krukoff 1349 (Uw 19305), 7039 (Uw high forest as trees of up to 8 m high.

8126), 6143, type (Uw 7513), 6223 (Uw 7568). Material studied: S. dioica Bureau. New

S. hirtella Mildbread. Brazil, Acre; Krukoff Caledonia: Sarlin G 153, ex C.T.F.T. 6201 (Uw 5084 (Uw 19745), 5785 (Uw 20188); Peru: 24627), ex SJRw 14688 s.n., (Uw 18425), s.n.,

Schunke V. 4827 (Uw 21086). - S. muriculata exSJRw 14301 (Uw 18424).

Amazonas: Krukoff 6988 General Miq. Brazil, (Uw features: Growth rings absent; 8082), 8520 (Uw 16200);Prance & Berg 18468 light yellow brown, unknown whether dealing 188 IAWA Bulletin Vol. 7 1986 n.s., (3),

with heartwood or sapwood. Texture medium and irregular clusters of 2—5; 3—5 (lO)permm,

to coarse, grain straight. Specific gravity 500- round to slightly oval, diameter 145—205 jam,

540 N cubic vessel member 400-520 Perfora- per metre. length jam.

Microscopic features: Vessels diffuse, tions simple, end walls almost transverse. Inter-

solitary (25-50%) and in short radial multiples vascular pits alternate, round and/orpolygonal,

and occasionally irregular clusters of 2—5; 4-9 7—9 jam. Vessel-ray and vessel-parenchymapits per sq.mm, round to slightly oval, diameter larger, elongated, half-bordered, the borders in

130—160 vessel member 425—530 reduced. Thin-walled but pm, length part tyloses common,

Perforations end walls almost Fibres with small sim- pm. simple, trans- scarce. nonseptate, many

verse. Intervascular pits alternate, round and/or ple pits in the radial walls, occasionally also in polygonal, 9—ll pm. Vessel-ray and vessel-paren- the tangential walls; walls 3-5 jam, lumina 9

the 20 fibres chyma pits larger, elongated, half-bordered, jam, gelatinous occasionally present;

borders often reduced. few thin-walled 1245—1575 2.7—3.7. Very length /am; F/V-ratio

tyloses occasionally present. Fibres septate, Rays uniseriate and multiseriate, heterogeneous

several septa per fibre, but also nonseptate in type 11/111, 4-6 per mm. Uniseriate rays 5—20

the with small restrict- of and same sample, simple pits %, composed upright square cells, height

ed the radial walls 3—4 lumina 7 3—5 cells Multiseriate to walls; pm, up to (180—360 jam).

20 pm, occasionally gelatinous; length 1080— rays composed of procumbent cells except for

1255 pm; F/V-ratio 2.1—2.6. Rays uniseriate the uniseriate margins of I—4 rows of upright

and 3—5 cells to 600— multiseriate, heterogeneous type 11/111, and/or square cells, wide, up

6-7 per mm. Uniseriate rays 8-23%, composed 660 /am high. Occasionally with some sheath

of cells. aliform with square, upright and some procumbent cells, Parenchyma long narrow

height up to 6-13 cells (180-600 pm). Multi- wings, confluent and sometimes in short wavy

seriate rays composed of procumbent cells ex- bands. Parenchyma strands of 2 cells. Rhombic

cept for the uniseriate margins of I—2 (in Uw crystals common in the square and upright

of of and few in the 18425 I—6) rows square and/or upright marginal ray cells, few, very two

3—5 cells 720—1450 from Zaire. Radial latex tubes cells; wide, up to pm high. samples com-

Occasionally with a few sheath cells. Parenchy- mon, the diameter of the tubes large compared

ma in concentric bands, partly enclosing the with the diameter of the ray cells (tangential

vessels, 4—14 cells wide and 1.4—2 per mm, sections).

and some scanty paratracheal strands. Paren- chyma strands of 2—4 cells. Rhombic crystals

abundant in the axial parenchyma cells, and Discussion

few in the ray cells. Vitreous silica in the fibres

and the axial parenchyma of one sample (Uw Wood anatomical variation within genera

18425). Most studied genera are very homogeneous

in their wood anatomical characters. This holds

Treculia Decaisne (Figs. 25, 26) in particular for Artocarpus, Bagassa, Pararto-

of the A recent monograph genus by Berg carpus, Poulsenia, Sparattosyce and Treculia.

(1977b) comprises three species. The genus is Within Batocarpus, slightly more variation is

distributed as trees or shrubs from Senegal to found in the amount and distribution of the

Sudan, Angola, Mozambique, and Madagascar. parenchyma (Figs. 9, 10).

Species available for this study are trees up to In Prainea, the sample of P. limpato deviates

30(—50) m high, from humid to dry primary by its narrow vessels and high specific gravity.

forest The of P. shows forests, swamp or secondary vegetation, sample frutescens relatively to from sea level 1300 m altitude. low multiseriate rays. These few exceptions do

Material studied: T. africana Decaisne. not disturb the impression of Prainea as a natu-

Ivory Coast: Detienne 117 (Uw 24640); Came- ral, well defined genus.

roon: Foury 131 (Uw 25687); Sao Tome: Museu The wood anatomy of Sorocea is also homo-

Ultramar no. 30(Uw 26978); Zaire; Louis 2507 geneous. The fact, that the vessel diameter of

(Uw 24242), Inst. Nat. Agron. 3144, ex MADw S. muriculata is lower than those of the other

17257 is due (Uw 26977). species (see genus description), not to a

General features: Growth rings absent; dry habitat. Only one of the samples studied

light brown, no differences observed between by us was collected on a dry slope in the Sipali-

sapwood and heartwood. Texture medium to wini Savanna in southern Surinam showing that

in coarse, grainstraight. Specific gravity 480—670 the species can endure water stress this area

metre. N per cubic with its long dry season and low annual rainfall.

Microscopic features: Vessels diffuse, However, according to Burger et al. (1962),

solitary (32—50%) and in short radial multiples this species occurs in ‘moist forests and the I AWA Bulletin n.s., Vol. 7 (3), 1986 189

firma and the Varzeas.’ In other cells Anti- mata de terra right ray only. Crystals are lacking in aspects, S. muriculala also completely fits in aropsis, Artocarpus, Batocarpus (mostly), Poul- with the samples of other species, at least two senia and Prainea. In Clarisia, crystals vary not

of which are known from an area with an an- only quantitatively, but also in distribution. nual rainfall of more than 7000 mm (i.e., Rhombic crystals are absent in C. biflora. Few

Choco, near Bajo Calima, Colombia); S. fausti- to very few rhombic crystals occur in C. race-

rhodorachis. three niana and S. trophoides ssp. mosa: they are absent in samples; in one Clarisia much less few rhombic observ- The genus is homogeneous. sample very crystals were

Three species were recognised by Berg (1968). ed in the ray cells; in three samples from Vene-

The wood anatomical characters be used zuela and Colombia few few can to to very crystals identify the individual species quite easily (see occur in the axial parenchyma cells. The occur-

Table 1 on page 185). In 1936 the genera Clari- rence of the crystals in C. ilicifolia follows a

sia and Acanthinophyllum were revised by Lan- comparable pattern: three samples lack crystals; jouw. He concluded that they are congeneric. in one sample very few crystals occur in the ray

A monograph including these two genera was cells; in the three other samples many to few published in 1962 by Burger. He separated the crystals occur in the ray cells and in the axial species with capitate pistillate inflorescences as parenchyma cells. a distinct genus, Acanthinophyllum, from Cla- risia, leaving only C. biflora and C. racemosa in Vitreous silica

Clarisia. In Acanthinophyllum two species, viz. The quantity of vitreous silica is always

A. and A. variable. It surround the ilicifolium spruceanum, were recog- (very) may tyloses, nised. The recent taxonomic discussion but in it fills the most some samples completely up on this problem is by Berg (1968). In his opin- smaller vessels. Fibres and axial parenchyma ion, A. spruceanum, only known from the type and occasionally also some of the ray cells may

collection, cannot be distinguished from the be completely or partly filled.

very variable A. ilicifolium. As a result Berg in- As can be seen in Table 2, silica is always

cluded A. spruceanum in A. ilicifolium. Berg present in Artocarpus, Poulsenia and Prainea ;

also re-evaluated the characters used by Burger it was never found in Antiaropsis, Bagassa, Par-

(1962) to separate A. ilicifolium from Clarisia. artocarpus, Sorocea and Treculia.

His conclusion is that Acanthinophyllum does

not merit generic rank. Consequently, A. ilici- Axial and radial latex tubes folium was transferred again to Clarisia. The A study of the occurrence of axial and radial

wood anatomical differences as observed be- latex tubes in Artocarpus was carried out by

tween the three species thus accommodated in Topper and Koek-Noorman (1980). They con-

Clarisia are substantial (Table 1). Based on cluded that the value of axial latex tubes in

these differences could defend be one a genus taxonomy seems to restricted. Although

rank for C. ilicifolia as suggested by Burger radial latex tubes were recorded for all species

account (1962). Taking into the variation range of Artocarpus studied, in three species both

within other, larger genera, however, we decided specimens with and specimens without radial

to follow the genus concept of Berg (1968). latex tubes were encountered.

The radial latex tubes can be classified into

Crystals two types, differing in relative size:

Rhombic crystals are rather common in the 1. radial latex tubes with a diameter more or studied. The in the genera crystals occur ray less the cells equal to surroundingray as seen cells, and less frequently in the axial paren- in tangential sections; chyma cells. The number of crystals is general- 2. radial latex tubes with a diameter (much) ly known to be variable from sample to sample. larger than the surroundingray cells. Also in the material used for this study, the

amount of crystals varies considerably. How- Tubes of type 2 are easy to detect, even with-

and distribution of out ever, presence crystals, a special staining. However, tubes oftype 1

regardless of quantity, distinguishes(groups of) can only be observed if the contents of the

genera (Table 2). tubes have a colour distinct from that of the

Sorocea Bagassa and are similar in their crys- ray cells and are retained in the sections, or

tal distribution (predominantly in square and when special staining with Sudan III is applied.

cells few upright ray but also in procumbent The results presented here are partly based on

ray cells and axial parenchyma). Parartocarpus observations of unstained sections. Therefore,

and Sparattosyce show crystals in axial paren- further studies, using specific staining tech-

chyma (Sparattosyce also occasionally in ray niques, may eventually reveal the occurrence of Treculia shows in and radial latex of cells), crystals square up- tubes type 1 in more taxa. 190 lAW A Bulletin n.s., Vol. 7 (3), 1986

>

CO 9

*< B B B B u4>Q. >■ >»rayS III in ho III III RaysCJ type Il/III II/III III III 5 ho 111 III ll/III II/III II/III *3 (Il)/IIl/(ho) s

I

1 to vo 6 -o b b b 3.8bo bi bv OJ U) to U) OJ w o 4^ 4-a to to F/V-ratio.g!S13V *>C tt. 3.0 1 1 1 1 1 1 1 1 U) to U)2.3-3.9 -o2.7-3.0 b3.9-4.4 2.6-4.2bv 2.2— b2.0-2.1 3.1-3.5 bl2.S-3.9 2.1-2.I— o2.7-3.7 to to U) to to to U) to to to

stamens. CO

Ul o VO o oo to Ul o Ov ~o U)1230-1420 00 001080-1255 4^ OJ to o o to urticaceous vo VO 9 - -

septate

1

Ol Moreae OJ U) to to 0 VO vo /am)§?é(in sizea> .s«° pit'S. -9 1 1 7—9 9-131 1 1 6-111 6-9 1 6-12 1 1 7-9 9-13 7-10 9-12 ~o 14—15 OV 9-11 the VO vo O Ov VO On VO of genera

U) o Ov tyi U) 4^ VO (10) to 9 & 4(7) 5(10) sq.mm)BE (pern. w frequency§o*(U

CO o o t^i o O 4^ OJ o o characters 00 Ov Ov Ol Ov VO 00 ON Ol Ol O (%)_5S vessels%“ > solitary« ** —S 50 45-801 20-601 40-651 25-511 21—60 35-901 50-841 20-63 25-501 32-501 CO 4^ to 4^ to to U) Ot to to OJ VesselsCD anatomical < _ o Ol o o Ot o Ol o o Ov 4-* -u o Ov 4^ t-o Ot * M ■U /am)B S(in lengthS member-sC 0>E 1 1 -1 960-1500 1 1 1 1 - - 1080-1460 525 - 1170-1950 I - 1245-1575 T- 945-1150 - —1165-1700 - 1370—1810 1410-1415 Wood 325-475 Ol350-630 325-445 295-440 310-485 465—740Ov 670-695-o Ol350-450 J t 425-530to 400-520o to vo to u> U) U) U) U) 4^ Ov 2. Table

Ot o o V/l Ol O O o o o ON o U) to to to to U) to to to /am) (ine= diameterua> +-• sC CTJ -5 108 1 1 1 1 1 1 108 160-310 145-240 125-205 75-2001 170—275 130-220 68-130 130-160 145-205 Ov to O 105-210 U) OJ 9 75 O 68 9

I

Co <* c r> Co CO S' V; 52 c c T3 S' *3 •3 2 o o a & S' Cl s O & £ S' Ol CO o Ço £ r> c S' Co' S“ 2 o o S' 3 •-» c g 1 Cl N Sparattosyce Antiaropsis>5 Arlocarpus•>( Batocarpus Parartocarpus Genus BagassaCl Cl Clarisia Poulsenia Prainea Sorocea Treculia o to to Q 2= ? §■ 8> ■& 3 lAWA Bulletin n.s., Vol. 7 (3), 1986 191

ho: to

*o p)*0 p)TJ ■—/ (p) W aliform; Crystal v) *< vt) p)’o £ J“*3 equal < 1 •—* i 1 (f, 1 | silicas® vitreous£I'> (v, vt,< j-h (f, (v,< (f, /—s (1968); as: parenchyma; J-hf, svt, less or view). Kribs in ata w a p; a) a yp, more y y to crystals& a I p, p, ata p,y (p)s * rhombic'Z •£ 1 1 1 1 uC vasicentric; c/u, ta Cu, (u,'p cu, fibres; v: tubes (tangential

in to ■— 2) / according to to to the f; « " 3 'to (2) (2) tubess latex~ radialI2 +(2) +(1) 1 +(2) 1 1 1 cells + + + -/+( + ++ + III, of ray silica: II, paratracheal; the

toi U) OJ diameter mm)E (pera § “ c types of bands•“ number1 4-51 1 1 2-31 r1-3 1 i 1 z~v\1.4—2 I 4l to 2.7-5 Vitreous scanty 1:

o sp: diameter ON ON NO £ tubes: cells) (in « S 1 1 i 1 1 1 1 -aband Iwidth 1 1 cells. 3-6 2-6 2-9 1 OJ 3- to tO 4-14 4*. heterogeneous the diffuse; latex

1 than s cb + 1 1 + + + (+) i + III: d: /—s Radial parenchyma II, larger wb i 1 1 + (+)3 (+) i (+) i - /“S U ta -- Rays: axial O c i (+)3 1 (+) (+)3 (+) 3(+) (+) i i (+) 3 1 distribution: bands, (much) in *< c B-al i + 1 1 i i a: 3* + (+) (+) + (+) + r—V /—s o* O (+)---- tubes 3 2. C/3 cells; faas i + + + + i i 1 O 3 (+)3 (+)3 present. concentric the Parenchyma distributionC/3 ta ray

»o _(+)_____+ sp 3 1 (+) 1 I 1 1 1 1 C/3 3 (+)3 3 occasionally bands; diameter only. procumbent dCL 1 1 (+)3 1 1 1 1 1 1 1 1 l wavy 2: cells in 1 O' ■o O' ~o O' O» -o NO -O On /-smm)i (perS> &frequencySI brackets; p: <2 7'J 3—6 3-71 4-6 4-7 1 4-6 3—51 4—7 1 6-91 6—71 wb: view); U) u> 4-6 4l 4l 4L U) ON On —+(1) —f,— —+——— cells; 8 ——- 3-10 ——+—— a — S Ol Ol - - U) cells) .S(in72width 4-* - On On On c/1 to» Ul U\ 1 1 - - procumbent 1 1- 1 1 1 1 ray>. “ So j between C/3 multiseriate-g IIE 3-4 3—5 2-5 4-6 3-6 2-3 4—6 vessels. U) to to 3—5 2-5 3-5 3-5 tangential OJ 4L U) OJ to OJ U) confluent; 'C upright ta of Rays (in the >3 c:

o o o o O o O o o and in o o absent; Q O' cells 4L to to to tO pm) (in22height 00 .S .a £ £ On — — — — — £3. wings; >.ray Soif 1 multiseriate■g S —: O' composed ray O o tyloses 600-840 o O O o O 0 o 600-660 square 960-960-1400 600—1200 O 700-1400 600-1560 600-1200 960—1400 480-1200 660-1200 720-1450 o < On Os On O On On NO 4L ON -O long the i.e., in in of present; u: vt: c*3 with c C5 (continued) 5’ 05 £ C +: g S' *§ -3 -5 <■> O 2 s § S' vessels; k. £ S' o C/3 5 c* diameter n i aliform c ■>s 2 £ o 3 ? s s c. homogeneous, in o Antiaropsis Artocarpus>« I Batocarpus Parartocarpus Sparattosyce distribution: & Poulsenia Sorocea Treculia Legend: Table Genus Bagassa Clarisia Prainea1 the o to Q 3= 3= S’ 3 al; v: 192 lAWA Bulletin n.s., Vol, 7 (3), 1986

dis- As far as we know, radial latex tubes are vitreous silica, and latex tubes, parenchyma present in all samples of: tribution and the occurrence of septate fibres

Antiaropsis : type 2, radial, diameter approx. are characters sometimes useful in classification.

28 pm, few; Three genera are considered to be closely al-

Artocarpus : type 1, radial and axial (in part), lied by all taxonomists: Artocarpus, Pararto-

10-20 and Treculia. Jarrett mentions pm, relatively abundant; carpus (1959a)

Bagassa ; type 1, radial and occasionally Prainea as a close ally ofArtocarpus with Parar-

axial, 12-20 pm, few; tocarpus and Treculia in a less closely allied

Parartocarpus : type 2, radial, diameter 20-48 series. We found no particular wood anatomical

features close between pm, usually abundant; to support a relationship

Poulsenia lb- these four their of ; type 2,radial,diameter(10) genera, despite possession

24 pm, few; vasicentric-aliform (to confluent) parenchyma.

Treculia : type 2, radial, diameter 24-48 This pattern is also found, however, in Bagassa

few abundant. and which in the pm, to Poulsenia, were never placed

The genus Clarisia is problematicwith respect same tribe with Artocarpus and allies before.

the of radial latex tubes. The with concentric to presence They are genera parenchyma

absent in C. racemosa and C. biflora. In C. ilici- bands were formerly placed in two different folia three of the four specimens from Surinam groups; Clarisia and Sorocea in subtribe Euarto- show some radial latex tubes of type 1 (diam- carpeae (Engler, 1888) and Moreae (Corner,

12 and latter eter approx. pm). 1962); Antiaropsis Sparattosyce (the Olme- One latex tube of size type 2 was observed in Ficeae according to Engler, 1888) in in Sorocea hirtella (Uw 19745). As none of the dieae (Corner, 1962). The Olmedieae (= Castil-

other Sorocea samples studied showed any leae sensu Berg, 1977a) are characterised by the latex of fibres & tubes, the genus Sorocea is regarded es- presence septate (Mennega Lanzing- latex tubes. fibres also in sentially as a taxon without Finally, Vinkenborg, 1977).Septate occur

latex tubes found in the Bato- and and no were genera Antiaropsis Sparattosyce (this paper)

carpus, Prainea, and Sparattosyce. Olmedia (Ter Welle et al., 1986). Within the

Based on these data, the occurrence of radial Castilleae sensu Berg the parenchyma pattern

to latex tubes may be regarded as a useful charac- varies from aliform-confluent banded (Koek- ter et for generic diagnosis, even if we keep in Noorman al., 1984), and we found no wood mind the possibility that narrow latex tubes anatomical arguments to exclude Anliaropsis,

overlooked and Olmedia from the Olmedieae. (type 1) may have been in some Sparattosyce taxa until now. Prainea is the only genus with septate fibres,

which has never assigned to the Olmedieae.

Identificationof the individual genera

It will be evident from the also All individual genera studied are (highly) foregoing(see homogeneous, except Clarisia, where several Ter Welle et al., 1986), that wood anatomical

to wood anatomical characters vary from species characters within the Moreae can be used

the comments to species, or evenwithin species. identify (groups of) genera, but that

In contrast, the wood anatomy on the inter- on the delimitation of this tribe can only be

generic level is heterogeneous. Most individual made when the family as a whole is taken into

genera may be identified using the following consideration. This will be the subject of a sub- fibres Anti- features: the occurrence of septate ( sequent paper. aropsis, Prainea, Sparattosyce); of rhombic crys- tals ( Bagassa, Clarisia, Parartocarpus, Sorocea, Acknowledgements

Sparattosyce, Treculia), homogeneousrays (Cla- Several persons, in one way or another, con- risia, Parartocarpus), and the parenchyma pat- tributed in the preparation and presentation of

tern: bands in contrast to a vasicentric or ali- this series of five publications on the wood form-confluent distribution (see Table 2). The anatomy of the Moraceae. only problem is found in Clarisia, as the three In the first place we wish to thank the cura-

to tors species appear be variable in all these features of the wood collections all over the world.

with the exception of absence of septate fibres. Owing to their help and willingness in sending

us duplicates and answering our questions con-

Taxonomic affinities cerning collectors and localities, we were able

The 11 genera studied, now united in the to study almost all genera of the Moraceae,

group of Moreae without urticaceous stamens using exclusively well-vouchered material. Spe-

to curators (Berg, 1983), have been assigned toseveral tribes cial thanks are due the of the wood

and subtribes in the past (see Introduction). collections in Madison (U. S.A.), Hamburg and

Next to the presence or absence of crystals. Berlin (F.R.G.), Tervuren (Belgium), Leiden lAWA Bulletin n.s., Vol. 7 (3), 1986 193

and Amsterdam (the Netherlands), Nogent-sur- — 1959b. Ibid. II. A revision of Prainea. J.

Mame (France), and Kew and Oxford (U.K.), Arn. Arbor. 40: 30—37.

who together provided about 85% of the taxa — 1959c. Ibid. III. A revision of Artocarpus not represented in the Utrecht wood collection. subgenus Artocarpus. J. Arn. Arbor. 40:

Drs. L.Y. Th. Westra and G. Schatz kindly cor- 113-155. rected the English text of the manuscripts. Mr. — 1959d. Ibid. III. Ibid, (continued). J. Arn.

E. Elsendoorn, Mr. M.H. Rypkema, and Mr. T. Arbor. 40: 298-326.

Schipper prepared the photographs and plates. — 1959e. Ibid. III. Ibid, (continued). J. Am.

The discussions with Dr. A.M.W. Mennega and Arbor. 40: 327-368.

Dr. C.C. Berg (Utrecht), and Dr. P. Baas (Leiden) — 1960a. Ibid. IV. A revision of Artocarpus have done much to improve the manuscripts. subgenus Pseudojaca. J. Arn. Arbor. 41:

73-109.

References — 1960b. Ibid. IV. Ibid, (continued). J. Am.

Bentham, G. & J.D. Hooker. 1880. GeneraPlan- Arbor, 41: 111-140.

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17; 309-312. Welle. 1984. Systematic wood anatomy of

— 1975. Flora of Suriname V (1). Moraceae: Moraceae (Urticales). I. Tribe Castilleae.

173-299. Brill, Leiden. IAWA Bull. n.s. 5: 183-195.

— 1977a. The Castilleae, a tribe of the Mora- Kribs, D.A. 1968. Commercial foreignwoodson

ceae, renamed and redefined due to the ex- the American market.Dover Pub., New York.

clusion of the type Olmedia from the ‘Ol- Lanjouw, J. 1936. Studies in Moraceae II. The

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— 1977b. Revisions ofAfrican Moraceae (excl. onyms with a discussion of the generic

Dorstenia, , Musanga and Myrianthus). name. Rec. Trav. Bot. Need. 33: 254—276,

Bull. Jard. Bot. Nat. Belg. 47: 267-407. Mello Filho, L.E. & M. Emmerich. 1968. Revi-

— 1983. Dispersal and distribution in the Ur- sao do genero Batocarpus Krst. (Moraceae-

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Ver. no 7: 219-229. Mennega, A.M.W. & M. Lanzing-Vinkenborg.

— & R.W.A.P. Akkermans. 1985. Studies on 1977. On the wood anatomy of the tribe

the flora of the Guianas 14. New taxa and ‘Olmedia’ (Moraceae) and the position of

combinations in Sorocea (Moraceae) and a the genus Olmedia R. & P. Acta Bot. Neeri.

key to its species. Proc. Kon. Ned. Akad. 26: 1-27.

Wet. 88: 381-395. ser.C, Slooten, H.J. van. 1968. Investigacion y desar-

Burger, W.C. 1962. Studies in the New World rollo de zonas forestales selectas de Costa

Moraceae: , Clarisia,Acanthinophyl- Rica. Inst. Interamer. Cienc. Agricolas,

lum. Ann. Missouri Bot. Card. 49: 1—34. Turrialba, Costa Rica.

— J.Lanjouw & J.G. Wessels Boer. 1962. The Topper, S.M.C. & J.Koek-Noorman. 1980,The ,

genus Sorocea St.Hil. (Moraceae).Acta Bot. occurrence of axial latex tubes in the sec-

Neeri. 11: 428-477. ondary xylem of some species of Artocar-

Corner, E.J.H. 1962. The classification of Mo- pus J.R. & G. Forster (Moraceae). IAWA

raceae. Card. Bull. Sing. 19; 187—252. Bull. n.s. 1: 113-119.

— 1976. A new species of Parartocarpus (Mo- Welle, B.J.H. ter, J. Koek-Noorman & S.M.C.

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1888. Engler, A. Moraceae. In: Die natiiriichen my of the Moraceae (Urticales) IV. Genera

Pflanzenfamilien 3 (1) (ed. A. Engler & of the tribe Moreae with urticaceous sta-

K. 66—98. Prantl): Leipzig. mens. IAWA Bull. n.s. 7: 91-128.

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allied genera. I. General considerations. J. Botanical Series, Field Museum of Natural

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