183

The systematic wood anatomy of the (Urticales)

I. Tribe Castilleaen

by

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

Institute ofSystematic Botany, University ofUtrecht, Heidelberglaan 2, 3508 TC Utrecht,

The Netherlands

Summary

The wood anatomy of the tribe The main aim of the present research project

sensu is described. Similarities differ- Berg and is to find an answer to the following questions:

ences are discussed in relation to his What is the anatomical in concepts variability range taxa of the of the tribe. The wood anato- of Moraceae as compared to closely related

mical variation does not enable to distinguish taxa? — To what extent can the anatomical in-

between . , and formation be used to contribute to our under-

Antiaris, , , and standing of patterns of relationship, and aid in

Naucleopsis can be recognised on the basis of generic and tribal delimitation?

slight differences. However, no reasons are A thorough treatment of the wood anatomy

found to question the delimitation of the Cas- of the tribe Olmedieae was published by Men-

tilleae sensu Berg on the basis of their wood nega and Lanzing-Vinkenborg (1977). One of

anatomy. their conclusions was that the exclusion of Ol-

Key words: Wood anatomy, systematics, media (Berg, 1977a) was justified. They did

Moraceae, Castilleae. not discuss the relation of the Olmedieae (later

renamed Castilleae) to the other Moraceae. Be-

Introduction sides, they did not study the African genera The first present paper is the one of a series and Mesogyne, also placed in the Cas- with the dealing systematic wood anatomy of tilleae by Berg (1973). Therefore we decided to

Moraceae. It includes wood anatomical include the Castilleae in of the descrip- our survey tions of the of the tribe Castilleae. This genera wood anatomy of Moraceae. We intend to pre-

project follows in logical the work of sent a series of in each of which the sequence papers, C.C. taxonomist of the who had Berg, family, genera of one tribe are described and their

experienced problems in delimitingand defining wood anatomical similarities and differences

the family as a whole, as well as the individual are discussed in relation to ideas about delimi-

and tribes. The wood and leaf genera anatomy tation and relationships as brought forward by,

of the Cecropiaceae, a segregate of the Mora- i.a.. Corner (1962, and his unpublished Flora

in relation the of this Malesiana ceae, to taxonomy group account of 1972) and Berg (1972,

has been discussed in Bonsen already a paper by 1977a, b, 1983). We will follow the most re-

and Ter Welle (1983). Their study on the cent on sys- views ofBerg the tribal arrangement, as tematic of the indicated him anatomy Urticaceae, closely re- by in his paper on distribution

lated to the Moraceae, is in the press (1984). and dispersal in the Urticales (1983).

Most wood anatomical literature Mora- We intend on to publish separate papers on the

ceae is rather difficult to taxonomi- tribes ofthe Moraceae the folio interpret in wing sequence; cally, due to the often dubious identifications I. Castilleae (Antiaris, Castilla, Helicostylis,

and the nomenclature! frequent changes. Fur- Maquira, Mesogyne, , Perebea,

thermore, descriptions are nearly always based Pseudolmedia );

few and relate on samples only to individual II. Dorstenieae i Bosqueiopsis, , Dor-

genera or commercially important timbers. stenia, Helianthostylis, , Tryma-

Numerous accounts in the literature were com- of tococcus; and Utsetela no

pared with our findings. Here we only refer to material is available);

literature elaborate including wood anatomical III. Ficeae ( );

descriptions based on identified mate- IV. Moreae characterised the reliably p.p., by presence rial, and not mentioned elsewhere in the text: of ‘urticaceous’ stamens (, Cal- Janssonius, 1934; Koek-Noorman & ter Welle, pidochlamys, Cardiogyne, Chlorophora, 1976; Record & 1938. Hess, 1940;Tippo, s.l. inch Plecospermum and Cudra-

1 This project was made possible by a grant ofBION-ZWO (14.45-01). 1984 184 lAWA Bulletin n.s., Vol. 5 (3),

Pa- the established nia, Maillardia, Malaisia, Militia, , Finally, specific gravity was

chytrophe, s.l., s.l. incl. in the usual way. Measuring of the moisture

mate- content of the considered unneces- Olmedia,i; of Ampalis and Fatouia no samples was

rial is available); sary as all samples were stored in a room with

the absence central and air of V. Moreae p.p., characterised by heating an humidity c. 40%

of ‘urticaceous’ stamens ( , Arto- during at least several weeks. It is the general

Clarisia, Parar- experience that this results in a relative mois- carpus, , , of tocarpus, , Prainea, , Spa- ture content approx. 12%.

Treculia; of Hullettia material rattosyce, no is available).

intend to discuss In a concluding paper we Generic descriptions

the taxonomic value of the various wood ana-

tomical features the tribal level and to on com- Antiaris Lesch. (Figs. 1, 2) the individual tribes. pare From the 17 , described since the

was established, Corner (1962) reduced

the number to four, but he already pointed out

Material and Methods that there were no essential differences between

combined all Most wood samples studied are backed by these species. Berg (1977b) spe-

herbarium vouchers which were identified by cies in Antiaris toxicaria Lesch., lowering the

C.C. Berg, the taxonomist of the family. Most rank of these four species to subspecies. The

to 40-60 samples were taken from the trunk of the tree; genus Antiaris (trees, up m tall, or

sometimes, however, the origin was unknown. shrubs) occurs in the Palaeotropics in primary

institu- and forests in wet and Many wood samples were provided by secondary dry habitats,

all the world. The from level 1500 altitude. tional wood collections over sea up to m

Utrecht wood collection served as a first basis Material studied: A. toxicaria Lesch.

for this study. In the generic descriptions, only Uganda: B. T. Styles 201 (Uw 19390), For.

8-1950 Zaire: C. Donis413 those samples are cited of which sections were Dept. (Uw 18387):

made. The general features of the wood are (Uw 24211), J. Louis 11201 (Uw 18780); West

Africa: F.P.R.L. 209 A East Afri- often based upon the study of more wood (Uw 18386);

samples not cited here. Sections and macera- ca: H.E. Desch-F.R.I.C.S. (Uw 24305); Malay-

to standard tech- sia: For. 9-1954 tions were prepared according Dept. (Uw 18388); Indonesia,

niques and embedded in Canada balsam and in Java: Koorders 8316t/127S7B (Uw 24417),

glycerin respectively. The sections were stained 1127g/22746B (Uw 24418), 1898m/13415B

with safranin and the macerations in astrablue. (Uw 24430); Indonesia, Irian Jaya: BW 11519

Besides, unstained sections were also studied. (Uw 20481), BW 11916 (Uw 18169), Fokkinga

The terminology proposed by the Commit- 1573 (Uw 24306); Philippines: Bur. of For.,

tee on Nomenclature of the IAWA (1964) is Manila P.l. 21092b.f. (Uw 24431).

followed. As for the quantitative data: vessel General features: Growth rings faint

diameters were measured in tangential direc- or absent; colour light brown, no difference be-

tion including the walls and averages are based tween sapwood and heartwood. Texture coarse;

Vessel is based interlocked. 270-530 N on 25 measurements. frequency grain Specific gravity

25 In the for both cubic on counts. descriptions, per metre.

characters minimum and maximum averages Microscopic features: Vessels diffuse,

found of each in towards a in samples genus are provided. some samples tending diagonal ar- in The percentage of solitary vessels was calculated rangement; solitary (45—65%) and short ra-

after examiningan area with at least 100 spores. dial multiples or irregular clusters of 2—6; 3—

10 to diameter 145— Clusters and multiples were regarded as 2, 3, 4, per sq.mm, round oval,

425-575 etc. vessels, depending on the number ofvessels 230 pun, vessel member length pun.

Vessel member and fibre Perforations end walls transverse. Inter- per group. length simple,

length are based on at least 100 measurements vascular pits alternate,round, oval or polygonal,

and minimum and maximum 9-13 Vessel-ray and vessel-parenchyma per sample, again, pun. half-bordered. sample averages found in the genera are pro- pits larger and irregularly shaped,

vided. Additionally, the averages were used to Thin-walled tyloses occasionally present in

calculate the ratio of fibre length/vessel mem- samples from Asia. Fibres septate with small

ber length, in the descriptions referred to as simple pits restricted to the radial walls; walls

wall thick- 1—3 lumina 10-30 often F/V-ratio. For the fibres, maximum pun, pun; gelatinous;

and maximum lumen diameter 1030-1390 1.8-3.1. ness are given. length pun; F/V-ratio

The percentage of uniseriate rays is based on Rays uniseriate and multiseriate, 4—7 per mm. Uniseriate of counts ofat least 100 rays. rays 2—15%, composed procum- 185 lAWA Bulletin n.s., Vol. 5 (3), 1984

bent and few with small restrict- square or upright cells, height up or partly septate simple pits

10 cells Multiseriate ed to the radial walls 1—3 lumina to (430 pun). rays com- walls; pun, 2 of cells uniseriate 15—25 posed procumbent , margins pun, occasionally gelatinous; length

of 1—2 of cells 1000—1200 2.0—2.9. (4) rows square and/or upright pun; F/V-ratio Rays uni-

and occasionally few sheath cells; 4—7 cells seriate and multiseriate,4—7 per mm. Uniseriate

1100 of wide and up to pun high. Parenchyma pa- rays 0—25%, composed procumbent and up-

vasicentric-aliform with short 6—10 cells ratracheal, wings, right cells, height up to (150—500 occasionally confluent, parenchyma strands of pun). Multiseriate rays composed of procum-

2 3—5 (8) cells. Radial latex tubes present in all bent cells with uniseriate of 1—3 rows , margins

Rhombic in of cells and few sheath samples. crystals few, marginal ray square and/or upright cells and axial in cells in 3—6 cells 800— parenchyma some samples some rays; wide, up to

from Asia. 1300 the pun high. Parenchyma paratracheal,

Note: Rhombic and elongated crystals are pattern varying from almost restricted to vasi- extremely abundant in the axial parenchyma centric or aliform with short wings to confluent-

and cells of the ofKoorders the bands 20 cells wide and 1—2 marginal ray sample banded, up to

1127g/22746B. per mm. Parenchyma strands of 2—4 (8) cells.

Radial latex tubes present in all samples. Vitre-

Castilla Sesse Cervantes silica in (Figs. 3, 4) ous present in some vessels in two samples.

Berg (1972) monographed the genus and dis- Note: Castilla ulei (Williams 1802) devia-

three C. elastica tes to tinguished species: Cerv., C. in the high uniseriate rays (up 30 cells,

tunu and C. ulei Warb. The Cas- 650 Hemsl., genus pun) and the narrow multiseriate rays (up

tilla occurs in Central America, the coastal re- to 3 cells wide) with uniseriate margins of 3—6

gion of NW. South America, and the Amazon rows of upright cells.

basin. Castilla elastica has been introduced and

naturalised in most tropical countries. Castilla Helicostylis Trecul (Fig. 13) consists of about 30-40 This trees, m tall, growing Neotropical genus was monographedby

in tropical rainforest up to 850 m altitude. Berg in 1972. He distinguished seven species,

Material studied: C. elastica Cerv. Gua- distributed mainly in the Amazon basin and

temala; McClay & Clara 30 (Uw 24222); Co- the coastal regions of Venezuela and the Guia-

Chocó: Cuatrecasas 14218 lombia, (Uw25161), nas. One species (H. tovarensis) occurs up to

Cuatrecasas 15217(Uw25289);Surinam:Linde- Costa Rica. Some species, originally described

man 229 (Uw 23350). — C. turn Hemsl. Pana- under Helicostylis, proved to belong to the

ma: Pittier s.n. (US 715986, 7048); UW Ecua- genus Brosimum. Helicostylis is a genus of trees,

dor: 424 to Berg (Uw 23615); Nicaragua: Engle- up 20-30 m tall, of the tropical lowland

sing 97 (Uw 18808). — C. ulei Warb. Brazil, rainforest.

Amazonas: Prance & Berg 19852 (Uw 20943); Material studied; H. elegans (Macbr.)

Brazil, Mato Grosso: Krukoff 1409 (Uw 18427); C.C. Berg. Brazil, Mato Grosso: Prance & Berg

Brazil, Rio Purus: Krukoff 5766 (Uw 18807); 18613 (Uw 20924), Prance & Berg 19868 (Uw

Peru: Ellenberg2949(Uw 8827), Williams 1802 20944); Brazil, Amazonas: Krukoff 6809 (Uw

(Uw 18433). 7946), 8075 (Uw 16159), 8521 (Uw 16201). -

General features; Growth faint rings H. pedunculata R. Ben. Surinam: van Donselaar

or absent; colour light yellowish brown, no de- 2992 (Uw 11963), Lanjouw & Lindeman 421 marcation between sapwood and heartwood. (Uw 1224), Lindeman 3544 (Uw 2315); French

Texture coarse, grain straight. Specific gravity Guiana: BAFOG 165 M (Uw 5244), 267 M (Uw

310—480 N per cubic metre. 5342); Brazil, Amapa: Pires 51670 (Uw 8956).

Microscopic features: Vessels diffuse, — H. scabra (Macbr.) C.C. Berg. Brazil, Amazo- solitary (30—70%) and in short radial multiples nas: Krukoff 6811 (Uw 7948), 7081 (Uw or irregular clusters of 2-6; 4-8 per sq.mm, 8162), Prance & Berg 17967 (Uw 20953). - H. round to oval, diameter 135—215 pun, vessel tomentosa (P. & E.) Rusby. Guyana: Fanshawe member Perforations length 350—560pun. sim- 148, For. Dept. 2757 (Uw 985), Maguire et al. ple, end walls transverse. Intervascular pits al- 45735 (Uw 16775); Surinam; Lindeman 4756

oval to 8-11 5356 5357 ternate, round, polygonal, pun. (Uw 3276), (Uw 3711), (Uw 3712),

Vessel-ray and vessel-parenchyma pits larger 6712 (Uw 4539), LBB (Maas) 10963 (Uw

and irregularly shaped, half-bordered. Thin- 11698); Brazil, Mato Grosso: Krukoff 1557

walled tyloses occasionally present. Fibres all (Uw 19400), 8859 (Uw 16256), Prance & Maas

2 ‘Procumbent’ refers to ratio very strongly procumbent cells with a length ; height of over 3;

‘weakly procumbent’ is used for length : height ratios of 2-3 : 1 of at least part ofthe procum-

bent ray cells. lAWA Bulletin Vol. 5 1984 186 n.s., (3),

15027 (Uw 19122);Brazil, Amapa: Fires 51834 Maquira Aublet (Fig. 7)

(Uw 9077). - H. tovarensis (Klotzsch & Kar- This Neotropical genus was revised by Berg

sten) C.C. Berg. Colombia, Choc6: Cuatrecasas in 1972. Species of this genus were formerly

14424 (Uw 24955). - H. turbinata C.C. Berg. assigned to i.a. Olmedia and Perebea. Berg dis-

Brazil, Amazonas: Krukoff 8832 (Uw 16258), tinguishes five species, all being medium-sized

type. or small trees of the tropical rainforest, up to

General features: Growth rings absent; 20-50 m tall, distributed in northern South

sapwood golden yellowish brown, heartwood America with one species up to Nicaragua.

light to dark brown, sometimes with dark brown Material studied: M. calophylla (P. &

C.C. streaks on the tangentialsurfaces. The demarca- E.) Berg. Brazil, Amazonas: Krukoff 6347

tion between sapwood and heartwood general- (Uw 7655), 6655 (Uw 7851), 8580 (Uw 16208),

ly irregular. Texture moderately fine, grain 8642 (Uw 16210). - M. coriacea (Karst.) C.C.

straight. Specific gravity 590—1070 N per cu- Berg. Brazil, Amazonas: INPA-Manaus X-3885

bic metre. (Uw 18851), Krukoff 6615 (Uw 7817), 6705

Microscopic features: Vessels diffuse (Uw 7886); Brazil, Amapa: Pires 51743 (Uw

but with a tendency towards a diagonalarrange- 9012); Venezuela: Breteler 3726 (Uw 11746).

ment; mostly solitary (70—90%) and in short — M. costaricana (Standi.)C.C. Berg. Colombia, radial of round Choco: Cuatrecasas 14135 — M. multiples 2-4;5-12 per sq.mm, (Uw 25382).

90—200 vessel member Surinam: Linde- to oval, diameter pm, guianensis Aublet. Lanjouw &

400—550 Perforations 2187 length pm. simple, end man (Uw 1644), 2811 (Uw 1936), LBB

walls almost transverse; intervascular pits alter- (Maas) 10795 (Uw 11210); Schulz 8980 (Uw

nate, round, oval or polygonal, 8—lOpun. Ves- 8855); Stahel 222 (Uw222); Brazil, Amazonas: sel-ray and vessel-parenchyma pits larger and ir- Pires 51803 (Uw 9634), Prance & Berg 18609 regularly shaped, half-bordered. Thin-walled (Uw 20921); Brazil, Amapa: Pires, Rodrigues

often Fibres all & Irwin 51612 — M. tyloses present. or partly sep- (Uw 8949). sclerophylla tate, in some specimens non-septate;with small (Ducke) C.C. Berg. Surinam: Lindeman 5040 simple pits restricted to the radial walls; walls (Uw 3424); Brazil, Para: Krukoff 1133 (Uw

2—5 lumina 7—14 in pm, pun; some samples 19224).

1140—1585 General features: Growth partly gelatinous; length pun; F/V- rings faint ratio 2.4—2.9. Rays uniseriate and multiseriate, or absent; colour (light) brown, no demarca-

4—7 per mm. Uniseriate rays 0—20%, generally tions between heartwood and sapwood. Tex-

composed of few rows ofprocumbent cells and ture medium, grain slightly interlocked. Speci-

few of fic 610—1000 N cubic to many rows upright cells, height up to gravity per metre.

8 cells Multiseriate entire- features: Vessels diffuse (150—350pun). rays Microscopic

1 ly composed of (weakly) procumbent cells with a slight tendency towards a diagonal ar- except for the uniseriate margins of 1—2 (5) rangement, solitary (40-90%) and in short ra-

of cells and few dial of 5-20 round rows square and/or upright multiples 2—4, per sq.mm,

sheath cells in of the 2-5 cells diameter 90—160 vessel some rays; (6) to oval, pun, member

310—900 400—640 Perforations wide, up to pun high. Parenchyma pa- length pun. simple, end ratracheal, vasicentric aliform with narrow to walls almost transverse. Intervascular pits alter-

broad sometimes towards uni- oval 7—9 Vessel- wings; tending nate, round, to polygonal, pun.

lateral arrangement abaxially, sometimes con- ray and vessel-parenchyma pits larger and irre-

sometimes terminal half-bordered. Thin-walled fluent; parenchyma pres- gularly shaped, ty-

ent. — Parenchyma strands of 4 (2 8) cells. loses often present. Fibres all or partly septate

Rhombic crystals in variable amounts in rays with small simple pits restricted to the radial

and axial Vitreous silica walls 3—5 lumina 5—9 occasion- parenchyma. present walls; pm, pun,

in the also in the 1015—1600 occasionally fibres, rarely ally gelatinous; length pun; F/V- axial parenchyma. Radial latex tubes always ratio 2—3. Rays uniseriate and multiseriate,

albeit sometimes Sclero- 6—8 Uniseriate present, sporadically. per mm. rays 0—10%, composed

tic cells noticed in the of and few occasionally rays and/or upright procumbent cells, height up

axial parenchyma. to 3-6 cells (150-300 Multiseriate pun). rays H. turbinata deviates in its 1 Note: F/V- composed of procumbent cells except for the ratio due to the fibres (3.9), relatively long uniseriate margins of 1—3 rows and square

(1585 pun) and short vessel members (400 pun). and/or upright cells and few sheath cells; 3—5

The is often confluent and cells 300— 1200 parenchyma locally wide, up to pun high. Parenchy- forms to 7—8 cells variable: wide to long parenchyma bands, up ma paratracheal, narrow vasi-

wide. Furthermore, the multiseriate rays are centric, aliform with short wings to confluent

frequently up to 7 cells wide and no diagnonal

vessel was found. (text continued arrangement on page 191) lAWA Bulletin n.s., Vol. 5 (3), 1984 187

1. — Antiaris Uw 24430. 2. Uw — Fig. toxicaria, Fig. Ibid., 18780. Fig. 3. Castilla elastica, Uw

25289. — 4. — Fig. C. ulei, Uw 20943. All x 360. 188 IAWA Bulletin n.s., Vol. 5 (3), 1984

Uw 5. Perebea 18431. — P. — Fig. xanthochyma, Fig. 6. guianensis. Uw 16183. Fig. 7. Maquira

— Uw 9012. 8. Pseudolmedia — coriacea, Fig. laevis, Uw 11817, All x 360. lAWA Bulletin n.s., Vol. 5 (3), 1984 189

9. — Fig. ulei Uw 18428. 10-12. N. — Naucleopsis , Fig. imitans, Uw 19991. All x 360. 190 lAWA Bulletin n.s., Vol. 5 (3), 1984

13. — Fig. Helicostylis turbinta, Uw 16258. Fig. 14-15. Mesogyne insignis, Uw 25797. — Fig.

16. Ibid., Uw 25686. — All x 360. IAWA Bulletin n.s., Vol. 5 (3), 1984 191

and in narrow bands, sometimes wavy, strands Naucleopsis Miquel (Figs. 9—12)

of 3-4 (2-8) cells. Radial latex tubes generally In this Neotropical genus Berg (1972) com- present, though sometimes very infrequent. bined two Perebea species with the species for-

Rhombic crystals occasionally present in the merly assigned to Acanthosphaera, Ogcodeia

and in the axial Vitre- and Palmolmedia. The consists of small rays rarely parenchyma. genus ous silica sometimes present in the axial paren- to medium-sized trees of the tropical lowland

chyma, rarely also in the vessels and fibres. and is distributed from Honduras to Rio de

Note: Wood anatomically, the individual Janeiro (Brazil). species show only part ofthe variation indicated Material studied: N. amara Ducke. Bra-

above. As a genus, however, Maquira is very zil, Mato Grosso: Krukoff 1406 (Uw 19340). — heterogeneous in, at least, the parenchyma pat- N. concinna (Standi.) C.C. Berg. Brazil, Ama- tern, the ray structure, the presence and fre- zonas. Krukoff 6620 (Uw 7822), 6687 (Uw

quency of crystals and silica, and the diameter 7875). — N. glabra Spruce ex Baillon. Brazil,

and density of the vessels. The wood structure Acre: Krukoff 5323 (Uw 19899); Brazil, Ama- shows an overlap with that found in Helicosty- zonas: Berg et al. 754 (Uw 24080), Krukoff lis, Perebea, and Pseudolmedia. 1300 (Uw 19277), 1392 (Uw 19331), 6198

(Uw 7549), 8271 (Uw 16170), Prance & Berg

Mesogyne Engler (Figs. 14 — 16) 18241 (Uw 20901), 19805 (Uw 20938); Brazil,

An African genus, monotypic according to Mato Grosso: Krukoff 1347 (Uw 19304). - N.

Berg (1977b). It consists of shrubs or trees, up guianensis (Mildbr.) C.C, Berg. Surinam; van to 15 (occasionally 40) m tall, of the lower Donselaar 3791 (Uw 12167), Lindeman 3790 stories of the tropical rainforest, between 500 (Uw 2776). — N. imitans (Ducke) C.C. Berg.

1300 m eastern and altitude, occurring in Tan- Brazil, Acre: Krukoff 5485 (Uw 19991). — N.

zania and Sao Tome. inaequalis (Ducke) C.C. Berg. Brazil, Mato Gros-

Material studied: M. insignis Engler. so: Krukoff 1458 (Uw 19364). — N. krukovii

Tanzania: Holst 2290 (Uw 25797), Scheffer 22 (Standi.) C.C. Berg. Brazil, Amazonas: Krukoff

(Uw 25795), Schlieben 158 (Uw 25686). 4544 (Uw 19524). — N. macrophylla Miq. Bra-

General features: Growth rings absent; zil, Amazonas: Ducke 232 (Uw 2667), Maguire colour light brown, no demarcation between et al. 56684 (Uw 16478), Prance & Berg 18543 heartwood and sapwood. Texture fine, grain in- (Uw 20920). - N. mello-barettoi (Standi.) C.C.

terlocked. Specific gravity 700-750 N per cubic Berg. Brazil, Amazonas: Krukoff 6839 (Uw metre. 7974). -N. stipularis Ducke. Brazil, Amazonas:

features: Vessels Microscopic diffuse, Ducke 370 (Uw 18423). - N. ternstroemiflora solitary (60—80%) and in short radial multiples (Mildbr.) C.C. Berg. Brazil, Amazonas: Krukoff

of 13—20 1306 - 2—4, per sq.mm, weakly angular (to (Uw 19281). N. ulei (Warb.) Ducke.

diameter 85—105 vessel member Amazonas; Krukoff round), pm, Brazil, 1301 (Uw 19278); length 375—505 pm. Perforations simple, end Peru: Williams 5194 (Uw 18428). almost Intervascular walls transverse. pits alter- General features: Growth rings faint nate, round or polygonal, 7— lOpun. Vessel-ray or absent; colour light brown, no demarcation and vessel-parenchyma pits larger and irregular- between heartwood and sapwood. Texture fine

sometimes ly shaped, half-bordered, apertures to medium; grain straight. Specific gravity be- coalescent. Thin-walled Fibres 570 and 990 N tyloses present. tween per cubic metre. septate with simple pits restricted to the radial Microscopic features: Vessels diffuse walls; walls 2—4 lumina 9-12 with pm, pun; occa- a tendency towards a diagonal arrange- 1125—1280 sionally gelatinous; length pm; ment, solitary (20-80%) and in short radial

2.4—3.4. uniseriate and multi- of 20—45 F/V-ratio Rays multiples 2—4; per sq.mm, angular, 8-11 Uniseriate seriate, per mm. rays 15-45 round or oval, diameter 60-110 pm, vessel

%, composed of few rows of weakly member 355—535 Perforations procum- length pm. sim- 2 bent cells and many rows of upright cells, ple, end walls almost transverse. Intervascular height to 11 cells (500 Multiseriate oval 6—9 up pun). pits alternate,round, to polygonal, pm. of rays composed (weakly) procumbent cells Vessel-ray and vessel-parenchyma pits larger for the uniseriate of 1—6 except margins rows and irregularly shaped, half-bordered. Thin-

of square and cells and few sheath cells walled Fibres upright tyloses occasionally present. sep- in some 2—4 cells wide, 700 tate rays; up to pun with small simple pits restricted to the ra- high. abundant; dial walls 2-4 lumina 5-10 Parenchyma paratracheal long walls; pirn, pirn; oc-

aliform-confluent to and 1000—1250 banded, apotracheal casionally gelatinous; length pm; in diffuse strands. some Parenchyma strands of F/V-ratio 2.1—2.9. Rays uniseriate and multi-

4 cells. Vitreous silica in fibres 6—8 Uniseriate (1—9) scarce, seriate, per mm. rays 0—15%, and axial of 2 parenchyma. composed procumbent and/or square to 192 lAWA Bulletin n.s., Vol. 5 (3), 1984

to 4—7 cells 420—550 Perforations end upright cells, height up (200-350 length pun. simple,

Multiseriate of walls almost transverse. Intervascular alter- pm), rays composed (weakly) pits

cells for the uniseriate oval 7—9 Vessel- procumbent except mar- nate, round, or polygonal, pun. gins of 1—6 rows of square and/or upright cells ray and vessel-parenchyma pits larger and irre- and few sheath cells; 2-4 (5) cells wide, and up gularly shaped, half-bordered. Thin-walled tylo-

600—1500 sometimes Fibres with to pm high, vertically ses occasionally present. septate

fused. Parenchymaparatracheal, vasicentric ali- small simple pits restricted to the radial walls;

the vasi- walls 2—4 lumina 7—18 form, occasionally locally confluent, pun, pim; occasionally

centric ring often incomplete and lacking on gelatinous; length 1050-1600 pun; F/V-ratio

the adaxial strands of 2-4 — cells. 1.9—3.4 uniseriate and side; (1 8) jam. Rays multiseriate, dia- Latex tubes present, generally with a small 5—7 per mm. Uniseriate rays 0—17%, composed 2 meter in comparison to the surrounding ray of procumbent and upright cells, height up to

cells. Rhombic crystals present in the ray cells 5—7 cells (150—340pan). Multiseriate rays com- of about 50% of the samples, rarely also in the posed of procumbent cells except for the uni- parenchyma tissue. Vitreous silica occasionally seriate margins of 1(—3) rows of square (up-

present in the fibres and parenchyma. right) cells and few sheath cells; 3-6 cells wide

Note: The samples ofN. stipularis, N. ama- and up to 500—1000 pm high. Parenchyma pa-

not ra and N. ulei (Williams 5194) do complete- ratracheal in generally complete, narrow, ali-

ly fit the generic description. N. amara deviates form vasicentric rings, often confluent in tan-

in the diameter (150 pm) and number (IS per gential or diagonal direction over a short dis-

fibre sq.mm) of the vessels and in its length tance, the sheaths usually widest on the abaxial

N. shows abundant side of the strands of 4 cells. Ra- (1450pm). stipularis paren- vessels; (2—8)

chyma in wavy paratracheal bands and ray mar- dial latex tubes often present, some axial latex

gins of up to 8 cells high. Both samples of N. tubes present in one sample (cf. Topper & ulei differ from each other in the vessel dia- Koek-Noorman, 1980). Rhombic crystals ab-

and the values the in in meter ray height: are among sent or scarce ray cells; one sample abun-

lowest ( Williams 5194) and highest (Krukoff dant in rays and axial parenchyma.Vitreoussilica

1301) values found in the genus. scarce in one sample, in both vessels and fibres. Note: One sample of P. guianensis (Kru-

in Perebea Aublet (Figs. 5, 6) koff 7209) deviates its narrow-aliform paren-

without confluent bands. The Ever since this Neotropical genus was estab- chyma, two sec-

the delimitation has been the tions of Perebea as defined by Berg, cannot be lished, generic ,

subject of discussion. In his monograph, Berg distinguished by their wood anatomy in the

(1972) distinguished eight species in two sec- available samples.

tions. The genus consists of small to medium

sized trees or shrubs. Representatives are found Pseudolmedia Trecul (Fig. 8)

in tropical lowland forest in tropical South Berg revised the genus and reduced the num-

Panama. to nine. trans- America and ber of species Some species were

Material studied: P. Aublet ferred other Pseudol- guianensis to genera (Berg, 1972). Brazil: ssp. guianensis. Irwin 48718 (Uw 16989), media occurs in the northern part of South

Krukoff 8350 (Uw 16183); Brazil, Amazonas: America, Central America and the Greater An-

Krukoff 7041 (Uw 8128), Krukoff 7209 (Uw tilles, as small or medium-sized to large trees of

8241). — P. mollis (P. & E.) Huber ssp. mollis. various habitats.

Brazil: Ducke 175 (Uw 2668); Brazil, Amazo- Material studied: P. laevigata Trecul.

nas: Krukoff 1289 (Uw 19268), 6985 (Uw Guyana: A.C. Smith 3633 (Uw 21706); Brazil,

8080). — P. mollis ssp. rubra (Trecul) C.C. Berg. Rio Solimoes: Ducke 359 (SJRw 40090, Uw

Surinam: Stahel 211 (Uw 211). — P. xantho- 2669); Brazil, Amazonas: Krukoff 6697 (Uw

chyma Karsten. Peru: Williams 3412 (Uw 7880), 8383 (Uw 16186), Prance & Berg 18611

18431). (Uw 20923); Colombia: J. de Bruijn 1543 (Uw

General features: Growth rings faint 14493). — P. laevis (R. & P.) Macbr. Guyana:

or absent; colour (light) brown, no demarca- Fanshawe 5378 (Uw 988); Brazil, Amazonas: tion between heartwood and sapwood. Texture Krukoff 5188 (Uw 19815); Brazil, Acre: Kru-

medium; grain straight. Specific gravity 530— koff 5217 (Uw 19823), 5312 (Uw 19889);

830 N per cubic metre. Venezuela: Breteler 3929 (Uw 11817). - P.

Microscopic features: Vessels diffuse macrophylla Trecul. Brazil, Amazonas: Prance

with a tendency towards a diagonal arrange- & Berg 18376 (Uw 20906); Bolivia; Krukoff

ment, solitary (40-80%) and in short radial 10738 (SJRw 39639, Uw 2670). P. oxyphyl-

of 5-15 round laria Donn. Smith. Mexico: I.N.I.F. XI52 multiples 2-4; per sq.mm, to (Uw

diameter 95-160 vessel member Belize: Stevenson 119 oval, jam, 20653); (SJRw 14901). IAWA Bulletin n.s., Vol. 5 (3), 1984 193

— P. Cuatrecasas. rigida (Klotzsch & Karsten) rattosyce, formerly included in the tribe Olme-

Ecuador: Maas, Berg & ter Welle 2918 (Uw dieae (Corner, 1962) were excluded by Berg

23575). — P. spuria (Swartz) Griseb. Guatema- and transferred to the Moreae. la: & Clara 31 Belize: The McClay (Uw 18027); eight genera of the tribe Castilleae are

Brown 35 similar in features show (SJRw 15351), many and considerable

General features: Growth rings absent; overlap, the differences being in quantitative colour demarcation between features far (light) brown, no only. As as we can judgenow, char- heartwood and sapwood. Texture fine to me- acters like vessel member and fibre length, pit

dium; grain straight or slightly interlocked. size and arrangement, ray structure, and occur-

metre. Specific gravity 710—940 N per cubic rence of crystals are of no or very little diag-

Microscopic features: Vessels diffuse, nostic or taxonomic value in this group. In short solitary (30-80%) and in radial multiples Table 1 the characters with at least some diag-

of 8—25 to 2—4; per sq.mm, round oval, dia- nostic value are listed: frequency and diameter meter vessel member 350— 70—165 p,m, length of the pores, the (tendency towards a) diagonal

540 Perforations end walls almost vessel the of either pm. simple, arrangement, presence intervascular transverse; pits alternate, round, weakly or strongly procumbent ray cells’, the

oval to 7—10 polygonal, pm. Vessel-ray and parenchyma distribution, the presence or ab-

vessel-parenchyma pits larger and irregularly sence of radial latex tubes, and the specific shaped, half-bordered. Thin-walled tyloses of- gravity. For these characters, in particular for

sclerotic in of P. ten present, some samples diagonal vessel arrangement and presence of

Fibres all with small laevigata. or partly septate, latex tubes, a careful examination of large areas

simple pits restricted to the radial walls; walls is required. Notwithstanding the high degree of

3—4 lumina 4—9 sometimes /xm, /am; gelatinous; similarity and overlap, some genera can be

1000-1600 2,4-3.4. combination length /am; F/V-ratio recognised by a of features.

uniseriate and 5—8 The Rays multiseriate, per mm. samples of Antiaris studied are all very Uniseriate rays 0-15%, composed of procum- similar. Only one sample is remarkable in the

bent and abundance of square, rarely upright cells, height up crystals. There are no wood ana-

to 3—7 cells Multiseriate tomical features which (150-600 /am). rays contradict the combina- 2 of cells with with- tion of all composed procumbent , or species into one species, as Berg

out 1 —2 rows of The square (rarely upright)margin- (1977b) proposed. genus can be recognised

al cells; 3—5 cells wide and to 370—1000 its low up /am by very specific gravity, a low number to high. Parenchyma paratracheal, confluent of relatively large pores per sq.mm, and paren- aliform wavy banded, occasionally with short chyma only in aliform (confluent)arrangement.

to strands of 3-6 cells. Radial long wings; (11) These features are also found in other Castil-

latex tubes are present, but infrequent and in- leae, but not in this combination. The wood

conspicuous. Axial latex tubes present in one anatomical features thus do not contradict in-

sample of iP. spuria. Rhombic crystals clusion of Antiaris in this tribe. Corner’s present sug- in cells and in axial that Pseudolmedia marginal ray parenchyma gestion (1962) might be one cells. Vitreous silica in most samples in vessels, of the nearest allies ol Antiaris, is not confirm- fibres axial or parenchyma. ed, neither is his idea, that Mesogyne and Anti-

Note: P. shows spuria rhombic crystals in aris might be congeneric (see below). procumbent as as well square and upright ray Castilla shares the low specific gravity (al- cells. ready mentioned by Mennega and Lanzing-

Vinkenborg, 1977) and the low number of

relatively large vessels with Antiaris, but differs

Discussion in its more abundant parenchyma, partly ar- In his of the tribe monograph Olraedieae, ranged in bands, comparable with those found

Berg (1972) presented a detailed of the in and Like in Meso- survey Mesogyne Maquira p.p.

taxonomic history of the tribe. In a subsequent gyne and Pseudolmedia, no trace of diagonal excluded Olmedia from paper, Berg the tribe vessel arrangement was found.

and renamed it Castilleae (1977a). As already Mesogyne and Naucleopsis both show rela- indicated in the Introduction, Mennega and vessels. At least of tively narrow part the pro-

2 studied cumbent cells Lanzing-Vinkenborg (1977) thoroughly ray are weakly procumbent . the six Neotropical genera now assigned to the Mesogyne is characterised abundant by paren- tribe Castilleae. For detailed discussion of the a chyma and the highest number ofrays per mm; wood of these refer to their anatomy taxa we latex tubes are lacking. A remarkable feature in paper. Berg (1977a) also included two African most specimens of Naucleopsis studied is the

genera, Antiaris and in the Castilleae. unilateral the Mesogyne, arrangement of paratracheal pa- The and Palaeotropical genera Antiaropsis Spa- renchyma. This feature was not found in N. sti- 194 lAWA Bulletin n.s., Vol. 5 (3), 1984

) (N/m gravity specific 3 270-530 310-480 590-1070 610-1000 700-750700-750 570-990 530-830 710-940 between cells;

a) a) a) a) a) a - u. - u, crystals rhombic u.a Oh(u.a) (p,u,a) (p,u,a) (p,u,a)(P,u. (p,u,a) u, (P,u, (p. (p, parenchyma

r r r r - r (ax) (ax) tubes latex r(ax) r(ax) r r axial

in

a: arrangement unilateral (+)(+) (+) + cells; bands (+) (+) ray diagonalor wavy + + Parenchyma diagonal (+)--- +-(+)--- - (+) (+)+(+)--- (+)--+(+)-- +- - - + + + confluent (+) + (+) (+) + procumbent Castilleae. mm) (per(per frequency 4-7 6-8 6-8 5-7 5-8 4-7 4-7 4-7 6-8 8-11 6-8 5-7 5-8 in

p: the cells) (in 4-7 3-6 3-5 2-4 2-4 3-63-6 3-5 cells; of width ray multiseriate 4-7 2-5 3-5 rayray

cells procumbent weakly - - ++ - + + - - features ys 2 squaresquare RaysRa anatomical cells marginal of rows 1-2(4) 1-3 1-2(5)1-2(5) 1-3 1-6 1-6 l(-3)l(-3) 1-2 and/or

15 25 20 1010 IS15 17 IS15 upright rays uniseriate % 15-45 < < < < < < < wood in

uu:

Some 9 pan) (in diameter lumenlumen 7-14 5-9 9-12 5-10 7-18 4-9 Fibres 10-30 12-25 4 1. direction, given. Table is arrangement diagonal (+) - (+) + - + + - axial

in

ax;ax; averages pun) (in size pit 9-13- 8-10- 3-6310-480 -8-10- 12-25 7-9- 3-5 7-10--9-12 6-96-9 7-9-- 7-10- direction; sample of Vessels pan) (in(in diameter 145-230 135-215 90-200 90-160 85-105 60-11060-110 95-160 70-16570-165 radial range in present. 185. the r: page sq.mm) (per frequency 3-10 4-8 5-12 5-20 13-20 20-45 5-15 8-258-25 page features absent; on

—: occasionally footnote quantitative present; see

Mesogyne Naucleopsis PseudolmediaPseudolmedia : Antiaris Castilla Helicostylis Maquira Perebea brackets: For +: 2 +: 2 lAWA Bulletin n.s., Vol. 5 (3), 1984 195

pularis where the parenchyma is arranged in — 1977b. Revisions of African Moraceae (ex- wavy irregular bands, including the vessels. cluding , Ficus, Musanga and My-

Helicostylis,with the exception ofthe sample rianthus). Bull. Jard. Bot. Nat. Belg. 47: of H. turbinata, is characterised by its short- 267-407.

aliform parenchyma, occasionally tending to- — 1983. Dispersal and distribution in the Urti-

wards unilateral and the cales - outline. In: a arrangement, pres- an Dispersal and distri- ence ofweakly procumbent ray cells. bution (ed. K. Kubitzki). Publ. Comp, Paul

As already indicated by Mennegaand Lanzing- Parey, Hamburg, Berlin.

Vinkenborg (1977), and as may appear after a Bonsen, K.J. & B.J.H. ter Welle. 1983. Compa-

Table the glance at I, remaining genera Maqui- rative wood and leaf anatomy of the Cecro- Perebea Pseudolmedia ra, and are extremely piaceae. Bull. Mus, natn. Hist, nat., Paris,

difficult on their wood structure. sect. to separate 4e ser., 5, B, Adansonia no 2: 151 — 177.

The absence of vessel in — — diagonal arrangement & 1984. Systematic wood anatomy and the of Pseudolmedia alone samples can hardly affinities of the Urticaceae. Bot. Jahrb. (in

be considered as a sufficiently reliable diagnos- press). tic and distinguishing feature. Berg (1972) dis- Corner, E.J.H. 1962. The classification of Mo-

tinguished sections in each Maquira, Perebea raceae. Card. Bull. Sing. 19: 187—252. and Pseudolmedia. In the wood anatomy these IAWA Committee, 1964. Multilingualglossary sections cannot be recognised. As will be evi- of terms used in wood anatomy. Konkordia,

not to dent from the above, we are able suggest Winterthur.

any other arrangement, in genera or subgenera, Janssonius, H.H. 1934. Micrographiedes Holzes than the one already proposed by Berg. der auf Java vorkommenden Baumarten.VI. In conclusion, in the material described in Brill, Leiden.

to this paper, we find no reason question the Koek-Noorman, J. & B.J.H. ter Welle. 1976. The

delimitation of the tribe Castilleae sensu Berg. anatomy of branch abscission layers in The genera Antiaris, Castilla, Helicostylis, Me- Perebea mollis and Naucleopsis guianensis sogyne, and Naucleopsis can be recognised. (Castilleae, Moraceae). In: Wood structure

Mesogyne most is the deviating genus. How- in biological and technological research there ever, are no features that are restricted to (eds. P. Baas, A.J. Bolton & D.M. Catling): this All shared genus only. are by at least one 196—203. Leiden Bot. Ser. 3. Leiden Univ.

other genus. Press, The Hague.

Maquira, Perebea and Pseudolmedia are hard- Mennega, A.M.W. & M. Lanzing-Vinkenborg.

and similar Heli- ly separable they are highly to 1977. On the wood anatomy of the tribe

costylis. ‘Olmedieae’ (Moraceae) and the position of

the genus Olmedia R. & P. Acta Bot. Neerl.

26: 1-27.

References Record, S. J. & R. W. Hess. 1940. American

Berg, C.C. 1972. Olmedieae, Brosimeae (Mora- woods of the family Moraceae. Trop. Woods ceae). Flora Neotropica. Monograph no 7. 61: 11-54. Hafner Publ. Comp., New York. Tippo, O. 1938. Comparative anatomy of the

— 1973. Some remarks on the classification Moraceae and their allies. Bot. Gaz. 100:

and delimitation of Moraceae. Meded. Bot. 1-99.

Mus. & Utrecht: 386. Herb,, Topper, S.M.C. & J, Koek-Noorman. 1980. The

— 1977a. The Castilleae, a tribe of the Mora- of axial latex occurrence tubes in the sec-

renamed and redefined to ceae, due the ex- ondary xylem of some species of Artocar- clusion of the Olmedia from the type genus pus J. R. & G. Forster (Moraceae). IAWA ‘Olmedieae’. Acta Bot. Neerl. 26: 73—82. Bull, n.s. 1: 113-119.