317

The systematic wood anatomy of the (Urticales)

II. Tribe Dorstenieaeri

by

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

Institute of Systematic Botany, University ofUtrecht, Heidelberglaan2, 3508 TC Utrecht,

The Netherlands

Summary

The wood of the tribe Dorstenieae in anatomy some genera and in some features, connec-

sensu Berg is described. Similarities and differ- tions with the ‘dump-like’ tribe Moreae (Berg,

discussed relation to his ences are in concepts 1977). The tribe consists of eight genera. In

of the of the tribe. Wood anatomi- our the taxonomy study Neotropical genera , cally the tribe Dorstenieae is fairly homogene- Helianthostylis and Trymatococcus and the

Dorstenia in the African ous, deviatingmost juvenilistic genera Bosqueiopsis and

of composition its rays, and the small diameter are represented as well as some African species

and high frequency of its vessels. of . Of Bosqueiopsis the, partly Neotropical, genus differs from the other in the of genera presence Scyphosyce and Utsetela, no samples were

fibre pits in the radial and tangential walls. available.

Helianthostylis and Trymatococcus are highly

similar. shows Brosimum a variation range ex-

ceeding that of the entire tribe. Nevertheless, Generic descriptions individual species of Brosimum can often not

be distinguished. Bosqueiopsis De Wild. & Th. Dur. (Figs. 1, 2).

The This African genera Brosimum, Helianthostylis, Try- genus, monographed by Berg matococcus, and Trilepisium are closely related. (1977), is monotypic since the four species de- The tribe Dorstenieae be from can separated scribed before, are united in his revision. The

the tribe Castilleae on wood anatomical charac- genus occurs in Zaire and southern Congo, as

ters such as: or absence of fi- trees 35 tall in and old second- presence septate up to m primary bres; distribution of the vessels; forest 500 altitude. In Tan- pattern para- are up to m eastern tracheal unilateral and parenchyma or not, ray zania and Mozambique as tall shrubs and tree- composition. lets to 6 in forests up m, thickets, open or words: wood Bos- Key Systematic anatomy. young secondary forest, often on sandy soils,

queiopsis, Brosimum, Dorstenia, Heliantho- from sea level to 500 m altitude (Berg, 1977).

stylis, Trilepisium, Trymatococcus. Material studied: B. gilletii De Wild.

& Th. Dur. East Africa: Schlieben 423 (Uw 15590).

Introduction General features: Growth rings faint; This paper is part of a series, in which the colour light brown; texture fine, grain straight. wood of the Moraceae anatomy is described in Specific gravity 800 N per cubic metre. detail and discussed relation in tothe taxonomy Microscopical features: Vessels diffuse, of the family. For an outline of the project as solitary (30%) and in short radial multiples and well as for the chapter Material and clusters of 60 Methods, irregular 2—6, per sq.mm, round refer to first we our contribution,the treatment to oval, diameter 45 pm, vessel member length of the tribe Castilleae (Koek-Noorman et al., 325 fun. Perforations simple, end walls almost

1984). In the tribe Dorstenieae, Berg (1973, transverse. Intervascular pits alternate, round

1977) included the tribe Brosimeae and the and size 6—9 oval, pm. Vessel-ray and vessel-

Dorstenia be the sole genus (considered to genus parenchyma pits larger and irregularly shaped, of the tribe Dorstenieae by Comer, 1962), be- half-bordered. Thin-walled tyloses abundant. cause of an intermediate position of one spe- Fibres with small non-septate simple pits in ra- cies of Dorstenia. The Dorstenieae, ‘clearly de- dial walls and less frequent in tangential walls; limited against the at Castilleae ..show, least walls 2—4 pm, gelatinous fibres marking the

1 This project was made possible by a grant of BION-ZWO (14.45-01). 318 lAWA Bulletin n.s., Vol. 5 (4), 1984

A gravity specificOC'3 8. 830 occasion- 520-700 750—1020750-1020 830 800800 570-800 880—1320880-1320 690-760690-760 700-800 670-790 1040-1200 440-650 occasion- brackets:

a 2 a) & (a) (a) a u o Bo (a) rr crystals3 rhombicjB •s - - » - r(a) rr, (r)(r) r,r r rr(a) (r.a) ft rrrr (r, between

absent;absent; CO X4> CO + + + + + + + + tubes-D latexJ9 radial3T3 - - + + —:-: present;

e»00 * | - wings■S arc-shapedT3 •s6 CO - - - - (+) - + (+) (+) - Brosimum. +: arrangement0)sg> c§§ unilateral*3 43 f3 + + + + + ++ (+)(+) (+) (+) (+) of parenchyma; regular + + + Parenchyma bandsband regular - + - + (+)+(+)+_ - - (+)(+) + (+)(+) (+) species axial

> * C

a: features cells; ray T) T T COtyloses1_o - - t (t) t - (0(t) T (t)(t) (t.T)(t, anatomical marginalmarginal

in

3. ) wood /an)/*■*» g •S(in asize.§ pit'5L 7—137-13 5-7 5-6 9 7-9 4—64-6 3-*3^1 9—119-11 9-11 5-9 8-13 abundantabundant

Some rr:

1. Vessels -- +—— -+ —— ——+ 700—800—— - - cells; 1 -+ •S

in

(30) r:

s e a* mm)/*"■ e sq. (per

t:

costaricanum gaudichaudii thick-walled; present. acutifolium alicastrumalicastmm costaricanum gaudichaudii glaziovii guianense lactescens parinarioides potabile rubescens utile

B. B. B. B. B. B. B. B. B. B. B. T: ally lAWA Bulletin n.s., Vol. 5 (4), 1984 319

lumina 4—8 830 growth rings, pm; length pm; rubescens Taub. Surinam: BBS 62 (Uw 659),

F/V-ratio 2.6. Rays uniseriate and multiseriate, Stahel 170a(Uw 170a);French Guiana; BAFOG

9 per mm. Uniseriate rays 13%, composed of 349M (Uw 5423); Colombia, Choco: Cuatreca-

procumbent, square and upright cells, up to sas 14030 (Uw 25316); Panama: USw 4489

4—5 (1-9) cells and 190(320) Multiseriate (Uw 11076). — B. utile B. Pitt. pm. (H. K.) ssp. occi-

rays composed of procumbent cells except for dentale C.C. Berg. Colombia, Choco: Cuatreca-

the uniseriate marginsof 1 (3) rows of square and sas 14291 (Uw 25174), 15595 (Uw 25345),

upright cells, and few sheath cells; 2-4 cells 16084 (Uw 25208). - B. utile (H.B.K.) Pitt.

to 560 wide, up pm high. Parenchyma para- ssp. ovatifolium (Ducke) C.C. Berg. Brazil, to tracheal, vasicentric aliform confluent, often Amazonas: Krukoff 6656 (Uw 7852). - B. utile

strands of 4 cells. forming wavy bands; (2—6) (H.B.K.) Pitt. ssp. utile. Colombia, Choco: van

Rooden, Topper & ter Welle 526 (Uw 25607).

Brosimum Schwartz (Figs. 3—12) General features: The woods of the

Berg (1972) recognised Brosimum sensu lato species ofBrosimum are variable in very appear- Piratinera Aublet (including Brosimopsis Moore, ance (Table 1). Highly remarkable are the dif- Ferolia and Aublet) and distinguished between ferences in colour and specific gravity. The

two the basis of dark subgenera on some morpholo- heaviest, reddish brown woods are found

gical features. Brosimum is a genus of trees, of- in B. guianense and B. rubescens commercially

ten of considerable with in known height, most species as ‘letterwood’ and ‘satine rubane’. Most

evergreen, mainly tropical rainforests. Brosi- other species are of considerably lower weight

mum gaudichaudiiis a common species of the (see Table 1) and light brown. Growth rings ab-

cerrados region of central South Ameri- sent. Texture fine campos to medium. Grain straight or

ca. The genus is distributed from Mexico and the interlocked.

Greater Cuba Jamaica Antilles, and to southern Microscopical features. Vessels diffuse, Brazil, with the highest concentration of spe- solitary (20—70%) and in short radial multiples

cies in the Amazon Basin. Some and clusters of 3—16 species, mainly irregular 2—4, (30) per sq. B. and B. valuable guianense rubescens, produce mm, round to oval, diameter 70—185 pm. Ves-

timbers (letterwood, satine rubane). sel member 400—600 Perforations length pm. Material studied: B. Huber acutifolium simple, end walls almost transverse. Intervascu-

ssp. Surinam: & Linde- lar acutifolium. Lanjouw pits alternate, round, oval or polygonal, 3—

man 2945 — B. Huber 13 (Uw 1978). acutifolium pm. Vessel-ray and vessel-parenchyma pits obovatum ssp. (Ducke)C.C. Berg. Guyana: A.C. variable, irregular in size and shape, half-bor- Smith 2617 (Uw 21528); Brazil, Amazonas; dered. Thin-walled tyloses often present; some- Krukoff 5378 6355 times thick-walled (Uw 19921), (Uw 7659). tyloses occur. Fibres non-

— B. alicastrum Sw. alicastrum. Jamaica: with small ssp. septate simple pits restricted to the

U.S. Nat. Herb. 5914 Guatemala: radial (Uw 8318); walls; walls 2-3 (4) pm, lumina 3-16

Peten 23112 (Uw 18018). — B. alicastrum Sw. (19) pm, rarely gelatinous; length 980—1600

bolivarense C.C. Guyana: A.C. Smith 2.0—3.4. ssp. Berg. pm, F/V-ratio Rays uniseriate and

3464 (Uw 21660); Venezuela: Breteler 3972 5—9 multiseriate, per mm. Uniseriate rays 0—

(Uw 12193); Peru: Uw 18000 MADw). - of (via 20%, composed upright cells only or inter-

B. costaricanum Liebm. Costa Rica; lica CCO- with mingled some rows of procumbent cells,

24 (Uw 20677). — B gaudichaudii Tree. Brazil: 3—6 cells (120—300pm) high. Multiseriate rays

al. 56136 — Maguire et (Uw 16368). B. glaziovii composed of procumbent cells with uniseriate

Taub. Brazil, Parana: Lindeman& deHaas 2060 of 1—2 of margins (7) rows square and upright 2201 B. (Uw 13491), (Uw 13588). guianense cells; 2—4 (7) cells to 370—610 wide, pp pm Huber. Surinam: BBS 55 (Aubl.) (Uw 652), high. Parenchyma paratracheal, vasicentric ali-

Stahel 3a 219 For. (Uw3a), (Uw 219); Guyana: form with narrow long wings, which are some-

Dept. 3366 3424 (Uw 986), (Uw 987); Brazil, times arc-shaped, occasionally confluent, some-

Amazonas: Krukoff 5477 — B. times (Uw 19984). banded; mostly unilateral;strands of 2—5 lactescens (S. Moore) C.C. A.C. Berg. Guyana: (1 —8) cells. Rhombic crystals varying from ab- Smith 2876 (Uw 21589); Venezuela: Breteler sent to in extremely abundant; marginal ray 3934 (Uw 12179), 3965 3970 (Uw 12190), cells, rarely in axial parenchyma. Radial latex

(Uw 12192); Amazonas: Krukoff 5377 tubes Brazil, common, although in variable amounts

8420 — (Uw 19920), (Uw 16193). B. parinarioi- and with different diameters.

des Ducke A.C.Smith Note: Silica ssp.parinarioides..Guyana: was found in B. lactescens

2901 (Uw 21593); Surinam: Linderaan 4461 Smith (A.C. 2876). Latex tubes are lacking in

Pulle 316 - (Uw 3120), (Uw 13846). B. pota- part of the samples of B. glaziovii and B.guia- bile Ducke. Amazonas: J. X1200 Brazil, Chagas nense.

Krukoff 6685 - (Uw 10536), (Uw 7873). B. (text continued on page 325) 320 lAWA Bulletin n.s., Vol. 5 (4), 1984

Fig. 1. Bosqueiopsis gilletii, Uw 15590. — Fig. 2. Ibid. — Fig. 3. Brosimum gaudichaudii, Uw

16368. — Fig. 4. B. guianense, Uw 652. lAWA Bulletin n.s., Vol. 5 (4), 1984 321

5. Brosimum Uw 19920. — Fig. lactescens, Fig. 6. B. parinarioides, Uw 3120. — Fig. 7. B. pota- bile, Uw 10536. — Fig. 8. B. utile, Uw 7852. 322 lAWA Bulletin n.s., Vol. 5 (4), 1984

Fig. 9. Brosimum guianense, Uw 19984. — Fig. 10. B. utile, Uw 7852. — Fig. 11. B. alicastrum.

Uw 12193. - Fig. 12. B. utile.Uw 25607. lAWA Bulletin n.s., Vol. 5 (4), 1984 323

13. Dorstenia — Fig. africana, Uw 25827. Fig. 14. D. involute, Uw 25826. — Fig. 15. Heliantho- stylis sprucei, Uw 23663. — Fig. 16. Ibid., Uw 24625. 324 lAWA Bulletin n.s., Vol. 5 (4), 1984

Fig. 17. Trilepisiummadagascariense, Uw 18391. — Fig. 18. Ibid. — Fig. 19. Trymatococcus ama- zonicus, Uw 2671. — Fig. 20. T. oligandrus, Uw 21199. lAWA Bulletin n.s., Vol. 5 (4), 1984 325

Dorstenia L. (Figs. 13, 14) Note: In the wood structure, a number of This is the second of the Mora- differences large genus between the samples of the two

ceae, with representatives in tropical America sections can be found: in D. africana and D.

and West Africa, to Zaire and kameruniana Nothodorstenia the up Angola. Berg (sect. ) paren- united the (1977) genus Craterogyne Lanjouw chyma is arranged in regular concentric bands; (the African species formerly assigned to Try- radial latex tubes and tyloses are present. In D. matococcus) and Dorstenia L. In this delimita- involuta and D. turbinata (sect. Eudorstenia)the

two African tion, sections of the genus are narrow and irregular vasicentric to aliform pat- The other African woody. Dorstenia species are tern is found; latex tubes and tyloses are absent. partly herbaceous with tuberous subter- ranean parts and succulent stems found in the Helianthostylis Baillon (Figs. 15, 16)

drier outside the This considered regions rainforest;partly they Neotropical genus was as a herbaceous to suffrutescent are plants and monotypical genus by Berg (1972). It consists shrubs found the rainforests. The in American of shrubs or trees up to 15m tall in the tropi-

species are all herbaceous. cal rainforest and secondary growths in the Material studied: D. africana (Baillon) Amazon Basin of Brazil, from the Rio Tapajoz

C.C. Cameroon: Berg. Hyman & Weerdenburg (Para) to Colombia. Of the species H. steyer- 303 328 (Uw 25827), (Uw 25831); Leeuwen- markii, described by Berg afterwards, no wood

9780 — berg (Uw 25777). D. involuta Hijman samples are available.

& Berg. Cameroon: Hijman & Weerdenburg 302 Material studied H. sprucei Baillon.

(Uw 25826). — D. kameruniana Engl. Zaire: Brazil, Amazonas: Ducke 180 (Uw 18422),

Louis 3078 (Uw 24224). — D. turbinata Engl. Prance & Berg 18780 (Uw 20927), Rodrigues Cameroon: Hijman & Weerdenburg 320 (Uw & Coelho 7336 (Uw 23663); Brazil, Para:

25829). Prance et al. 25509 (Uw 24081); Colombia:

General features: Growth rings ab- Rio Calima: Cuatrecasas 16303 (SJRw43019A, sent; colour light brown to cream-coloured, Uw 24625). without heartwood (all diameters 1—2 cm). General features: Growth rings absent;

Texture colour fine, grain straight. Specific gravity yellowish white, heartwood absent or 900—1200 N per cubic metre. indistinguishable from the sapwood. Texture features: Microscopical Vessels diffuse, medium, grain straight. Specific gravity 750— and in short solitary (30—70%) multiples of 850 N per cubic metre. 20—40 2—4," per sq.mm, angular, sometimes Microscopical features: Vessels diffuse, diameter 30—45 round, pm, vessel member solitary (30-60%) and in short radial multiples length 450—600 Perforations end of 15—25 round pm. simple, 2—4, per sq.mm, to oval, dia- walls almost transverse. Intervascular pits alter- meter 60—90 (130) pm, vessel member length

round, oval to polygonal, 4—6 Vessel- 340—350 Perforations nate, pm. pm. simple, end walls al- and ray vessel-parenchyma pits larger, tending most transverse. Intervascular pits alternate, towards a scalariform half-border- oval 7—9 Pits between arrangement, round, or polygonal, pm. ed. Thin-walled tyloses occasionally present. vessels and procumbent ray cells comparable Fibres with small non-septate pits restricted to with the intervascular pits in size; vessel-paren- the radial walls 3—4 walls; pm, often gelatinous; chyma pits and pits between vessels and upright lumina 9—11 960-1350 pm; length pm; F/V- ray cells larger and irregularly shaped, occasion- ratio 2.1—2.8. uniseriate and Rays multiseriate, ally tending to scalariform arrangement, half-

12—16 Uniseriate per mm. rays 50—80%, com- bordered. Thin-walled tyloses occasionally posed of 4-8 (-20) rows of upright occa- and present. Fibres non-septate with small pits re- sionally cells, to stricted to the radial walls 2—3 square up 420—645(—1550) walls; pm, oc- high. Multiseriate composed ofupright pm rays casionally gelatinous; lumina 5-10 pm; length and cells with uniseriate occasionally square 800—900 pm; F/V-ratio 2.3—2.6. Rays uni- margins of 4—13 rows of upright 2—3 cells seriate and 7-11 Uniser- cells; multiseriate, per mm. of the multiseriate wide; height parts strongly iate rays 10—25%, composed of few rows of within from 2—50 cells varying one sample strongly and weakly procumbent cells and (100—1500 pm). Parenchyma arrangement many rows of upright cells, 3—6 (10) cells varying from regular concentric bands (2-6 (165—280 pm) high. Multiseriate rays entirely cells 5—8 wide, per mm) to irregular, narrow composed of strongly and weakly procumbent vasicentric to aliform of patches parenchyma cells with uniseriate margins of 1—2 (8) rows tissue, occasionally apotracheal; strands of of square and/or upright cells; 2—3 (4) cells

2—4 (1—7) cells. Rhombic crystals in 230—550 scanty wide, up to pm high. Parenchyma pa- cells. Radial latex tubes marginal ray narrow, ratracheal, often unilateral, aliform with long

in of the present part samples. narrow wings, occasionally confluent to banded; 326 lAWA Bulletin n.s., Vol. 5 (4), 1984

strands of 2—3 cells. Radial latex tubes cells and axial Vitreous silica (1—4) ray parenchyma.

present but few and narrow. scarce, observed in vessels, fibres, and axial pa-

Note. One sample (Prance et al. 25509) is renchyma. Radial latex tubes present or absent,

slightly deviating in the presence of abundant see also the note below.

vitreous silica in the its to Note. the material studied is fibres, ray height(up Although very

1000 pm) and the ray margins (not over 1 cell homogeneous,some differences correlated with row high). According to Berg (1972), collec- the geographical distribution occur. In samples

tions from the same area (Para) are slightly de- from Zimbabwe, Tanzania and South Africa

in features well. the vessel diameter is 80—100 and latex viating some morphological as pm,

tubes are absent. In the other samples, from Ni-

and Trilepisium Thouars (Figs. 17, 18) geria, Uganda, Zaire, latex tubes are always

this is variable the vessel diameter varies from 135— Although genus highly (Berg, present;

the lack of clear distin- 150 1977) morphological pm. guishing characters forced Berg to unite all

in formerly described species one species: T. Trymatococcus P. & E. (Figs. 19, 20)

madagascariense DC. It occurs in tropical In the most recent revision (Berg, 1972), in

Africa, Madagascar and the Seychelles in prim- Trymatococcus three species are distinguished.

and forest from level The consists of shrubs and ary secondary sea up to genus trees (up to

2000 m altitude. 30 m tall) and occurs from the upper Amazon

Material studied: T. madagascariense Basin to the Guyanas.

DC. Nigeria: F.H.I. 7174 (Uw 18391); South Material studied: T. amazonicus P.&E.

Africa: Visser s.n. (Uw 25778); Tanzania;: Surinam: Lindeman 3773 (Uw 2759), 4735 F.P.R.L. 20039 (Uw 18382);Uganda: F.P.R.L. (Uw 3258), 4928 (Uw 3358), 5042 (Uw 3425),

23233 (Uw 18384), 23235 (Uw 18392);Zaire: 6442 (Uw 4422), Maas (LBB) 10832 (Uw 11241);

Vermoesen 1660 (Uw 18768), Dechamps 136 Peru: Williams 3767 (SJRw 18304); Brazil,

(Uw 24223); Zimbabwe: Swynnerton 16-1910 Amazonas: Ducke 174 (Uw 2671), Krukoff

(Uw 18393). 6092 (Uw 7474), Prance & Maas 13725 (Uw General features: Growth rings absent 18990), 14012 (Uw 19050), Prance & Berg

or faint; colour light brown, no demarcation 17972 (Uw 20880). - T. oligandrus (R. Ben.)

between sapwood and heartwood; texture fine Lanjouw. Surinam: Stahel 366 (Uw 366), Lan-

to medium, grain slightly interlocked. Specific jouw& Lindeman 2177 (Uw 1640);Guyana: Fan-

500-720 N cubic shawe 116 French Guiana: gravity per metre. (Uw 989); Maas et al.

Microscopical features. Vessels diffuse, 2195 (Uw 21199). — T. paraensis Ducke. Suri-

solitary (25—30%) and in radial multiples and nam: Lanjouw & Lindeman 2297 (Uw 1697).

irregular clusters of 2—5, 8—13 per sq.mm, General features ; Growthrings absent, round to oval, diameter 80—150 pm, vessel rarely faint; colour fight brown, heartwood

390—450 member length pm. Perforations probably present in one sample, dark brown.

transverse. simple, end walls almost Intervas- Texture fine to medium,grainstraight or slight-

to cular pits alternate, round, oval polygonal, ly interlocked. Specific gravity 600—850 N per

6—7 (9) pm. Vessel-ray and vessel-parenchyma cubic metre. pits larger and slightly irregular, the pits be- Microscopical features: Vessels diffuse,

tween vessels and procumbent ray cells com- solitary (50—70%) and in short radial multiples

in size with the intervascular half- and clusters of 7—9 parable pits, irregular 2—4; per sq.mm,

bordered. Thin-walled tyloses occasionally round to oval, diameter 95—110 pm, vessel

Fibres with small member 370-465 Perforations sim- present. non-septate simple length pm. pits restricted to the radial walls;walls 2—3 pm, ple, end walls almost transverse. Intervascular

lumina 7-14 900—1180 round 7-11 Pits be- pm; length pm; F/V- pits alternate, to oval, pm. ratio 2.0-2.9. Rays uniseriate and multiseriate, tween vessels and procumbentray cells compa-

6-8 per mm. Uniseriate rays 5—25%, composed rable with the intervascular pits in size; pits be-

of upright and square cells, up to 160-275 pm tween vessels and upright cells and vessel-paren-

(5-6 cells) high. Multiseriate rays composed of chyma pits irregular,round to elongate,tending

(weakly) procumbent cells, with uniseriate towards a scalariform pattern, half-bordered.

of 1—3 of margins rows square/upright cells; Thin-walled tyloses often present. Fibres non-

2—4 cells wide, up to 290-540 pm high. Paren- septate with small simple pits restricted to the

chyma paratracheal, in continuous but often radial walls; walls 2—3 pm, occasionally partly 2—4 cells 5—13 1000—1200 wavy bands, wide, partly or com- gelatinous;lumina pm, length pletely including the vessels; strands of 2—4 pm; F/V-ratio 2.6—2.8. Rays uniseriate and

cells. Rhombic abundant in the 8—10 Uniseriate crystals margin- multiseriate, per mm. rays

al also in of ray cells, occasionally procumbent 10—30%, composed square and upright cells, lAWA Bulletin n.s., Vol. 5 (4), 1984 327

o o to On to 00 g gravitygravity specificspecific 800 1 i OO 10-1200 50-850 00-720 440-1320 900-1200 750-850 500-720 8600-850 j. on O'

1 )'w' + crystals rhombic 1 +/- 'g'u 1 + g(u) .+ (

s-￿ + + + tubes latex radialradial 1 + (+) + (+)+ +

0»

'w' + + 3 unilateral 1 (+) 1 + (+)2 + unilateral N- Z

N—' 3* + (+)+ 1 (+)+ ++ (+) bandsbands /*“s /—s /*“S o

3 'w' + + 1 -1 + i n wavy/diagonal (+)

Parenchyma v—￿ to confluent ++ (+)+ 1 (+)+ 1 2(+) -o confluent

O' o VO oo mm) (per frequency 9VO 1 1 5-9 12-16 7-111 6-8 1 8-10 (Moraceae). "J oo

OJ cells)cells)of (number width 2—42—4 1 3-6 2-A 2—4 I2-4(7) to tO Dorstenieae present.3 *a v< present ± ± in » Rays cellscellsprocumbentprocumbent weakly 50 1 3 |. ON 'w' OJ /*N U) occasionally features /“«V (3)to I cells marginal ofrowsrows to to 1 4-13i 1-3 i=- 1 1-2(7)1 1-2(8)I 1-4(6) •&. ?•

« brackets:s On o o 8 O cr diagnostic to rays uniseriate percentage 13 1 r 1 3 uniseriate percentage 0-20 10-25 5-20i 50-80O O o10-30 O Or « On between? Some 5T (A O' ° oo 6» cr pan)pm) (in diameter lumenlumen I' 2. Fibres 4—8 3-16 5-10 7-14 fit 9-11 5-13 a u> NO -J cells; 3 On o« o Table ray to n> ■O 1 1 pm)ptm) (in(in size pitpit 6—96-9 1 4-6 7-9 6-7 3-13 ON 7-11 c -j ——— to - square - _-___ 2-3 - -+ £ o' On O o u> On O 00 CL on VO and/or On n> pm)/mi) (in diameter 45 1 L 30-45 60-90 r ( i 70-185o § 80-150o On95-110 u> M -o oo VO f Vesselsn 3. uprightXi < C

o S' On in US 5 U) u:C 60 O' 1 1 1 sq.mm)(per(per frequency O' 7-9 3-16(30) 020-40 ON15-25 8-13 -J 3 U) absent;s—: a —: Bosqueiopsis BrosimumBrosimum DonteniaDorstenia Helianthostylis Trilepisium TrymatococcusTrymatococcm present;+:.present; +: lAWA Bulletin Vol. 5 1984 328 n.s., (4),

Multiseri- confluent to banded found in the up to 6-9 cells (210-265 pm) high. parenchyma

of cells with Neotropical of Trymatococ- ate rays composed procumbent remaining species

uniseriate margins of 1—4 (6) rows of square cus, the relatively low number and the small

to of of not and upright cells; 2—4 cells wide, up 300- diameter the samples Dorstenia do

900 often a to return to the former in- pm high. Parenchyma paratracheal, justify suggestion unilateral, narrowly aliform with long wings, clusion of Nothodorstenia in Trymatococcus. occasionally confluent-banded;in some samples Of the other genera studied, the Neotropical

terminal parenchyma present; strands of 2—4 genera Helianthostylis and Trymatococcus are

cells. Rhombic crystals occasionally present in highly similar, the main difference being found

the marginal ray cells. Radial latex tubes present in in vessel frequency (Table 2).

all samples. Trilepisium (Africa) deviates from Helian- thostylis and Trymatococcus in the parenchy-

Discussion ma which is arranged in continuous, but often

The representatives of this tribe, seen in the wavy bands and the abundant occurrence of

of whole of the Mora- the rhombic in the perspective the family crystals ray margins.

ceae, constitute a wood anatomically fairly ho- Already at first view, Bosqueiopsis differs

there from the ofthe because mogeneous group, although are some diag- other genera Brosimeae,

nostic differences between individual genera. of the pattern formed by the small and numer-

The Dorstenieae are characterised by a regu- ous vessels, all included in confluent to wavy

lar arrangement of (small to) moderately sized parenchyma (Table 2; Fig, 1). Another remark-

vessels: non-septate libriform fibres; rays of able difference is found in the presence of fibre

Kribs (1968) type heterogeneous II, rarely I, pits on radial and tangential walls, whereas in

to i.e. uniseriate rays composed of upright and all other Brosimeae they are restricted the

multiseriate with 2-6-seriate radial fibre walls. square cells; rays

parts composed of procumbent cells, accom- In the samples of Brosimum, we found for

panied by short (to longer) uniseriate margins some features a range of variation that exceeds

of upright and square cells; parenchyma ali- the variation found in the other Dorstenieae

form-confluent to banded and at least partly (size of the intervascular pits, parenchyma pat-

when in unilateral arrangement. Crystals, pres- tern, specific gravity, the abundance of crystals,

ent, are found nearly exclusively in the marginal etc., see Table 1). Nevertheless, several species

ray cells. are difficult to separate. In itself, this is not

taxo- The samples of Dorstenia, a mainly herba- surprising, if we consider the frequent

Table nomic in the delimitation and ceous genus, are most deviating (see 2). changes generic

The vessels are narrower and more frequent subdivision within Brosimum (for a historical

than in the other The are see No fully cor- genera. rays composed survey, Berg, 1972). features,

of and cells and the related with the subdivision proposed Berg square upright only, per- by found. Neither wood anatomical fea- centage of uniseriate rays is high. It may be are were

questioned whether these features indicate an tures found, justifying the proposal of another

isolated position of Dorstenia, or if these fea- grouping ofspecies. Inconclusion that within tures are due to the small diameter of the sam- we may say the tribe

ples: all woody species of Dorstenia are (sub- Dorstenieae, the wood anatomy suggests an ex-

centric a paedomorphic position ofDorstenia. Bosqueiopsis, shrubs. The ray type suggests too,

structure (Carlquist, 1974: 394-399 and the differs in some noteworthy aspects from the

references given there). As already indicated in other representatives studied. The close rela-

the generic description, some differences are tionships of the other genera is confirmed.

found between the two represented sections of Berg (1977) stated that the Castilleae and

his the genus. Remarkable are the parenchyma pat- Dorstenieae in circumscription are clearly

terns: regular concentric bands in the samples distinguished. If we compare both tribes (see

of section Nothodorstenia, and narrowly ali- Koek-Noorman et al., 1984),we too find some

form, often irregular patches in section Eudor- features and tendencies which separate the two

from tribes. stenia. Both patterns deviate the arrange-

ment found in the other genera ofDorstenieae,

in particular the regular bands of section Notho- Castilleae:

dorstenia. The species of this section belonged — fibres septate, at least partly,

— often less formerly to the genus Craterogyne, established vessels in more or diagonal arrange-

by Lanjouw (1935) to accommodate the Afri- ment,

can species of Trymatococcus. Although the — parenchyma mostly surrounding the vessels,

could — uniseriate of bands found in Nothodorstenia be rays partly composed procum-

thought to be related, for instance, the aliform- bent cells (Kribs heterogeneousIII, rarely II). lAWA Bulletin n.s., Vol. 5 (4), 1984 329

Dorstenieae; — 1973. Some remarks on the classification

— fibres not septate, and delimitation of Moraceae. Meded. Bot.

— vessels regularly distributed Mus. & Herb. Univ. Utrecht: 386.

— parenchyma mostly tending towards unila- — 1977. Revisions of African Moraceae (ex-

teral arrangement, cluding Dorstenia, , Musanga and My-

— uniseriate rays composed of upright and rianthus). Bull. Jard. Bot. Nat. Belg. 47:

cells 267-407. square (Kribs heterogeneousII, rarely I). Carlquist, S. 1974. Island Biology. Columbia

Three species formerly assigned to Brosi- Univ. Press, New York.

transferred the E. J. 1962. classification of mum, were to genera placed in Corner, H. The

Castilleae: mello-barettoi, Pseudol- Moraceae. Card. Bull. Sing. 19: 187—252. media oxyphyllaria and spuria. Koek-Noorman, J., S.M.C. Topper & B.J.H. ter

found In Naucleopsis mello-barettoi we a uni- Welle. 1984. The systematic wood anatomy lateral parenchyma arrangement, but in all of Moraceae (Urticales) -1. Tribe Castilleae.

structure other respects the wood anatomical IAWA Bull. n.s. 5: 183-195.

to of the three species support their transfer Kribs, A. 1968. Commercial foreign woods on

the Castilleae. the American market. Dover Publ. Inc., New

York.

References Lanjouw, J. 1935. Studies in Moraceae, I. The

Berg, C.C. 1972. Olmedieae, Brosimeae (Mora- genera Trymatococcus Poepp. et Endl. and

ceae). Flora Neotropica. Monograph no 7. Craterogyne Lanj. Rec. Trav. Bot. Neeri. 32:

Hafner Publ. Comp., New York. 262-278.