Neerl. 1-27 Acta Bot. 26(1), February 1977, p.

On the wood anatomy of the tribe

“Olmedieae” () and the

position of the Olmedia R. & P.

Alberta+M.W. Mennega and Marijke Lanzing-Vinkenborg

Instituut voor Systematische Plantkunde, Utrecht

SUMMARY

The ofthe wood of structure the genera ,, , , Olmedia,

Perebealand; ,considered to in the Olmedieae (cf. Berg 1972) is , belong described.

The in anatomical between the is and it is hard diversity structure genera small, to distinguish

Maquira, and Pseudolmedia from each other. Castilla can be recognized by its thin-

walled and wide-lumined fibres, Helicostylis by its parenchyma distribution, Naucleopsis

its vessels smaller marked difference (usually) by more numerous with a diameter. A more

is shown the with banded instead of by monotypic genus Olmedia apotracheal parenchyma

the aliform confluent-banded of paratracheal to parenchyma the other genera. Septate

fibres, which characteristic for the other - of are genera some Helicostylis excepted - are nearly completely absent in Olmedia. This structural difference is considered as an

in favour of argument the exclusion of Olmedia from the tribe Olmedieae (Berg 1977).

1. INTRODUCTION

The structure of the secondary wood of the Moraceae shows in comparison to

that of many other families a rather uniform pattern. This is particularly true

for of the tribe Olmedieae. Differences found in size most genera are mainly

and numberof vessels, absence or presence of septate fibres, and in the distribu-

tion and quantity of axial parenchyma. Besides the description of the Moraceae

in Metcalfe& Chalk’s Anatomy of theDicotyledons (1950), we have Tippo’s

of the and of the American (1938) account family a treatment genera by

Record & Hess (1940). The latter authors pointed out how unreliable their

material from because the botanists who was, even originating type trees,

proposed the new species were sometimes uncertain as to the genera to which these species should be assigned.

recent time several taxonomic revisions which In were published among Berg's (1972) “Revision of the Neotropical Olmedieae and Brosimeae.”

While Berg studied the of the Olmedieae, the second author in-

vestigated the wood structure of our material of the genera at that time con-

sidered to belong to the Olmedieae. It appeared from her study (unpublished

doctorate thesis) that differences in wood structure between the species,

although small, do enable a certain arrangement into groups. She drew her

conclusions from anatomy independantly from Berg’s taxonomical results.

A subsequent comparison of their respective conclusions showed that the 2 A. M. W. MENNEGA AND M. LANZING-VINKENBORG

of the wood well with the taxonomic arrangement samples agreed remarkably arrangement.

It seemed desirable to wait with publication of the results ofthe anatomical

until additional material in not investigation genera or poorly represented up from table till then would be available. As appears I all seven genera which, according to Berg (1972), constitute the tribe Olmedieae are now represented in a most satisfactory way. Among others the genus Olmediais now represented.

recent Berg out that the Olmedia must be In a paper (1976) points genus excluded from the tribe Olmedieae for various reasons. This necessitated a redefinition for it: C. C. For of the tribe and a new name Berg. practical reasons the present paper still refers to the Olmedieae in the former wide sense.

2. MATERIAL AND METHODS

The material is listed in table 1. In the table the revised names as well as the

which the wood received from scientific names accompanied specimens other

institutes are listed. In those cases where the material was very abundant a selection was made of the samples to be sectioned, the other ones being in-

spected by a x 10 hand lens. All wood specimens are backed by herbarium

identified vouchers which were by Berg.

In the usual way sections and macerations were prepared for studying the

of microscopical characters. For some the specimens softening in hydrofluoric acid to Macerations made previous cutting was necessary. were by placing wood in mixture of of acetic acid and chips overnight a equal parts hydrogen

of °C. peroxide at a temperature 60

In the generic descriptions the averages for the length of vessel members and

fibres the values for the the are calculated from average species, lowest and

highest averages being placed between brackets. The ranges comprise the

lowest and highest valuesencounteredwithinthe species.

3. GENERAL ANATOMICAL CHARACTERS OF THE WOOD

Characteristic for all genera are the scattered vessels with simple perforations and moderate to rather large alternate intervascular pits ranging from 5.5-10

/im; vessel members with short tips or without tips, and a length of about 400-

of 500 /im. The fibre tissue consists libriformwith small simple pits on the radial

walls. Mostly fibres are septate with 1-3 septae per fibre, but in some genera,

like of the fibres are and in Olmedia Helicostylis , only part septate septate

fibres are extremely scarce. In a number of the fibres gelatinous walls are

nearly always present. The majority shows the long spindle form and ranges

from 1000-1700 but shorter fibres with wider and /im long, a body more

abruptly ending in bayonet-like tips are also present. In Castilla the fibres are

wider and thinner-walledthan in the other genera; in Naucleopsis they tend to be slightly shorter than in the other genera. The ratioof fibre to vessel member THE TRIBE 3 WOOD ANATOMY OF “OLMEDIEAE” (MORACEAE) of: scabra name elegans elegans the fallax fallax lancifolia asperifolia asperifolia poeppigiana Distributed under Perebea Perebea Pseudolmedia C. C. H. H. H. H. 23141

83 7946 7948 8162 Xylarium number elsewhere MADw USw SJRw USw MGw USw USw Grosso Grosso Grosso Grosso locality Purus

Mato R. Mato Amaz. Amaz. Amaz. Mato Mato Amapa Amaz. Amaz. Amaz. Amaz. Guiana Guiana Collection Guatemala Panama Nicaragua ------Suriname Suriname - Suriname - - - - Peru Braz. Braz. Peru Braz. Braz. Braz. Braz. Braz. Braz. Fr. Fr. Braz. Braz. Braz. Peru Braz. Braz. Guyana

(F.D.2757) 421 2992 19852 18613 19868 M 17967 2949 3544 11 2507 148 97 1409 5766 165 267M 6809 8075 8521 Lind. 68 7081 8762 c.s. c.s. c.s. 51670 nselaar c.s. 715986 Williams & Collector(s) Englesing Ellenberg Lindeman D Ellenberg Fanshawe US Krukoff Krukoff Prance, Krukoff Krukoff Krukoff Prance, Prance, Lanj. BAFOG BAFOG Pires van Krukoff Krukoff Krukoff Prance, - El.

of cm cm

5 5 cm cm cm cm cm cm cm 16 tree tree cm 25 32 26 8 32 24 Diameter branch, branch, cm cm cm cm cm cm cm cm cm cm the 100 12 14 14 8 10 12 9 over over over over over large over over - - - - 11 20 20

Utrecht wood number 18026 7048 18808 8827 18427 18807 18433 20943 7946 16159 16201 20924 20944 1224 3544 5244 5342 8956 11963 7948 8162 8710 16222 20953 985

Berg Berg Rusby C. C. Material C. Ben. C. E.) et R. of Cerv. (Macbr.) (Macbr.) (P. List Hemsl. Warb. 1. pedunculata pedunculata pedunculata pedunculata pedunculata pedunculata elastica tunu tunu ulei ulei ulei ulei ulei elegans elegans elegans elegans elegans scabra scabra scabra scabra scabra tomentosa Table Castilla Helicostylis 4 A. M. W. MENNEGA AND M. LANZICG-VINKENBORG caloph.) of: obliqua obliqua name calophylla the pedunculata calophylla calophylla Distributed under Olmedia (Olmedioperebea Olmediophaena (Pseudolmedia) Pseudolmedia H. O. O. X-3885 7655 7851 7817 7886 39526 Xylarium number elsewhere USw USw USw USw USw INPAw

Amapa Amaz. Amaz. Amaz. Amaz. Amaz. Amaz. Amaz. Amaz. Amapa Amaz.

- Collection Suriname Suriname Suriname Suriname Suriname Suriname Suriname Suriname Suriname Suriname ------locality Braz. Braz. Guiana Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Peru Venezuela Braz. Braz.

51743) 10963 45735 (Maguire) 15027 (Pires 4076 4756 5299 5356 5357 5469 6712 5 2823 8859 c.s. 1557 8832 6347 6655 8580 6705 7178 8316 661 3726 c.s. Collectors) 51834 (Maas) c.s. Lindeman Lindeman Lindeman Lindeman Lindeman Lindeman Lindeman Schulz Schulz Pires LBB Krukoff Maguire Prance Krukoff Krukoff Krukoff Krukoff Krukoff Krukoff Krukoff Ellenberg Breteler Maguire

of cm cm tree 14 18 Diameter cm cm cm cm cm cm cm cm cm cm cm cm cm cm cm the 2 5 2 3 5 5 5 1 14 10 24 13 20 over over 17 12

Utrecht wood number 3041 3267 3672 3711 3712 3791 4539 4924 6836 9077 11698 16256 16775 19122 19400 16258 7655 7851 16208 7817 7886 8774 11746 17149 18851

Berg

C.

Berg E.) C.

et C. (P. C. (Karst) (continued) Berg C.

1 tomentosa tomentosa tomentosa tomentosa tomentosa tomentosa tomentosa tomentosa tomentosa tomentosa tomentosa tomentosa tomentosa tomentosa tomentosa turbinata calophylla C. calophylla calophylla coriacea coriacea coriacea coriacea coriacea coriacea Table Maquira WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE) 5

Ducke amara of nna name maxima ulei caloneura laurifolia laurifolia xinguana cone concinna the temstroemiflora pallescens Distributed Olmedia Perebea Perebea Acanthosphaera Ogcodeia under Perebea Perebea Perebea Perebea Og. Og. Olmedia

39684 7822 7875 Xylarium number elsewhere USw INPAw USw USw Grosso Grosso Grosso Grosso Grosso Grosso Amaz. Amapa Amapa Mato Amaz. Mato Amaz. Amaz. Amaz. Mato Mato Amaz. Mato Mato ------Collection locality Braz. Suriname Suriname Suriname Suriname Suriname Suriname Suriname Braz. Braz. Suriname Braz. Suriname Braz. Braz. Braz.- Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Suriname

1 2187 281 1083 10795 Lindeman Lindeman 19609 18241 19805 3547 4757 Jonker 51612 51803 5040 3790 8642 & & 1133 1406 6620 6687 6198 1300 1347 1392 5323 222 & 8980 c.s. c.s. c.s. c.s. c.s. (Maas)

Collector(s) Krukoff Stahel Lanjouw Lanjouw Lindeman Lindeman Daniels Schulz Pires Pires LBB Prance Lindeman Krukoff Krukoff Krukoff Krukoff Krukoff Krukoff Krukoff Krukoff Krukoff Prance Prance Lindeman

cm cm cm cm 15 10 14 8 cm tree cm cm cm cm cm cm cm cm cm cm cm cm cm cm Diameter x 7 5 6 4.5 the over 10 14 5.5 17 10 1 over over 10 10 3 28 25 5

222 Utrecht wood number 16210 1644 1936 2318 3268 8597 8855 8949 9634 11210 20921 3424 19224 19340 7822 7875 7549 19277 19304 19331 19899 20901 20938 2776

Berg Berg C. C. C. C. Baill. (Stand.) (Ducke) ex Aubl. (Stand.) (Mildbr.) (continued) Berg Ducke Spruce Berg C, C.

1 costaricana guianensis guianensis guianensis guianensis guianensis guianensis guianensis guianensis guianensis guianensis guianensis sclerophylla C. sclerophylla amara concinna concinna glabra glabra glabra glabra glabra glabra glabra guianensisC. Table Naucleopsis 6 A. M. W. MENNEGA AND M. LANZING-VINKENBORG

of: stipularis ulei inaequalis caloneura ternstroemi- ternstroemi- name ulei

the Distributed Ogcodeia Naucleopsis Ogcodeia flora Palmolmedia Ogcodeia flora ulei ulei Acanthosphaera under Ogcodeia Og. Og.

16629 31959 7974 40

Xylarium number elsewhere Yale USw SJRw SJRw MADw MADw Grosso Grosso Grosso Grosso Grosso Amaz. Mato Amaz. Mato Amaz. Amaz. Mato Mato Mato Amaz. ------Collection locality Suriname Suriname Suriname Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Colombia Peru Braz. Peru Peru Panama

2544 Lanj. 3791 51293 18534 19807 18467 19881 5194 7082 6019 & 4700 c.s. 5485 1458 4544 6839 4969 1306 8271 1301 5196 Donselaar 232 c.s. 370 c.s. c.s. c.s. Williams Williams Williams Collectors) Lindeman Lindeman Maguire Krukoff Krukoff Krukoff Ducke Prance Krukoff Krukoff Ducke Prance Krukoff Prance Prance Krukoff Cuatrecasas Krukoff Dressier V. LI. LI. LI

cm cm cm cm cm cm 14 cm tree cm 16 tree cm cm cm cm cm 7 cm cm cm cm cm cm

Diameter 3 8 4.5 2.5 4.5 6 10,5 4 5 5 5 the 3.5 ca. 13 big 4.5 ca. 14 14 over

Utrecht wood number 1816 4700 12167 17042 19991 19364 19254 2667 20920 7974 19694 18423 20939 19281 20912 20960 16170 18421 18428 19278 18429 18430 21743

Berg Berg Berg C. C. C. C. (Mildbr.) C. C. (Standi.) Pav. Miq. Ducke (Ducke) Ducke et (Ducke) (Stand.) Berg (continued) Berg Ruiz, C. (Warb.) 1 C. temstroemiiflora temstroemiiflora guianensis guianensis guianensis guianensis imitans inaequalis krukovii macrophylla macrophylla mello-barreto mello-barretoi stipularis stipularis C. temstroemiiflora aspera aspera aspera Table C. ulei ulei ulei ulei Olmedia WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE) 7

of:

name tessmannii tessmannii pseudopeltata mollis mollis mollis ulei? ulei? the brosimifolia Distributed Noyera Ogcodeia Ogcodeia under Perebea Perebea Perebea Noyera Noyera Ps.

8128 8241 38402 23637 8080 40090 7880 8096 Xylarium number elsewhere USw USw USw SJRw USw SJRw MADw SJRw USw USw Grosso

Amaz. Amaz. Amaz. Amaz. Amaz. Mato Amaz. Amaz. ------Collection locality Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Suriname Colombia Peru Braz. Braz. Braz. Colombia Colombia Colombia Colombia Colombia Colombia

18525 s.n. 3412 48718 18131 7041 7209 8350 6985 1289 6697 7003 1063 1504. 1510 1538 1543 1557 c.s. 175 c.s. 359 c.s. 211 Williams Bruyn Bruyn Bruyn Bruyn Bruyn Bruyn Collector(s) Krukoff KrukolT Prance KrukolT Ducke Krukoff Krukoff Prance, Cuatracasas Krukoff Krukoff Irwin Stahel LI. Ducke de de de de de de

cm cm tree -16 14 cm Diameter cm cm cm cm cm cm cm cm cm cm cm 4 8 4 5 over shrub 7 the 18 over 24 53 26 26 43 24.5

Utrecht wood number 8128 8241 16989 20795 16183 2668 8080 19268 20918 211 18421 18431 2669 7880 8096 14465 14482 14484 14492 14493 14498

and (Tree.) guian. guian. guian. interned. Huber Karsten Aublet guian, E.) mollis mollis mollis rubra ssp. ssp. ssp. ssp. Tree. (continued) guianensis & mollis Berg (P ssp. ssp, ssp. ssp. C. 1 between acanthogyne guianensis ssp. guianensis guianensis guianensis guianensis mollis ssp. mollis mollis mollis mollis C. xanthochyma xanthochyma laevigata laevigata laevigata laevigata laevigata laevigata laevigata laevigata Table Perebea Pseudolmedia +laevigata 8 A. M. W. MENNEGA AND M. LANZING-VINKENBORG

of: name the mildbreadii Distributed sagotii laevis laevis under Ps. Ps. Ps. Ps,

39639 14901 15351 Xylarium number elsewhere SJRw SJRw SJRw Grosso Grosso Grosso Grosso Mato Amaz. Amaz. Amaz. Amaz. Mato Mato Mato Honduras Honduras Collection - Suriname Suriname Venezuela - - - - - Guatemala locality Braz. Braz. Guyana Braz. Braz. Braz. Braz. Braz. Braz. Bolivia Braz. Braz. Br. Mexico Br.

152 18376 35 18611 5378 5034 18437 18610 119 8383 3929 5188 5217 5312 5440 10738 8524 IN1F c.s. 96a Dept. c.s. c.s. c.s. Brown

S. Collector(s) Krukoff Prance Stahel For. Lindeman Breteler Krukoff Krukoff Krukoff Krukoff Prance Prance Krukoff Krukoff Prance Stevenson Mexico C. -

cm cm tree tree tree 12 10 tree tree tree cm cm cm cm cm cm big

Diameter 1 big 10 over 14 14 10 the 30 over big big big ?

96a 988 Utrecht wood number 16186 20923 3421 11817 19815 19823 19889 19916 20907 20922 2670 16202 20906 20653 18027 - -

Sm. Macbr. Tree. Donn. P.) Griseb.

et only (continued) (Sw.) (R.

1 laevigata laevigata laevis laevis laevis laevis laevis laevis laevis laevis laevis laevis macrophylla macrophylla oxyphyllaria spuria Sections Table +macrophylla +oxyphyllaria +spuria + 9 WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE)

is it between 2-3. In the tissue the length moderately high as ranges ray scarcity of uniseriate rays is characteristic; the multiseriates range from 2-8 cells wide, they mostly consist of numerous rows of low procumbent cells bordered by

short uniseriate the 2.6 often wings, highest rays up to mm high (165 cells), are

dissected fibres. seldom in diagonally by one or two Rays are contact with the

vessels, consequently ray/vessel pitting is rare and restricted to occasional half- bordered round in the cells and oval the pits procumbent larger pits in square and cells. Narrow latex tubes in the of all upright occur rays genera except

Olmedia. Crystals may be present but usually are not abundant; brown or

yellowish contents are generally Parenchyma, in Olmedia is present. except ,

always paratracheal in mostly incomplete, vasicentric, aliform rings, the

confluent banded lateral wings locally confluent or even forming parenchyma.

A narrow terminal band is occasionally present. Strands of 2-4 (8) cells; in the

2-celled strands the individual cells sometimes with long pointed ends, resem-

bling short fibres. Crystals rather scarce, but abundant in long rows of divided

cells in Pseudolmedia oxyphyllaria. Brown contents generally present. Pits

to vessels large, ovalor irregular, mainly in horizontalarrangement. In Olmedia

the parenchyma distribution is different, the parenchyma occurring in wide, regular, apotracheal bands.

4. KEY TO THE WOOD OF THE OLMEDIEAE

It not without hesitation that the this was a key to genera was prepared. In rather homogeneous group of genera it is extremely difficult to find distin-

guishing characters which are sufficiently clearly defined to allow someone not

familiar with the tribe to identify his material.This difficulty is enhanced by

the fact that in large genera represented by much material usually an odd

sample is likely to be present that does not correspond to the general picture of

the Aberrant features genus. are usually found in the parenchyma distribution and the average vessel number. From a detailedinvestigation of the material of Helicostylis tomentosa (table 2) it appeared that stems and branches with a

diameter of less than 5 differ deal in those features from cm a great more

mature stems. Therefore samples from such small-sized stems or branches

cannot be identifiedwith much certainty.

All this applies to the generaHelicostylis, Maquira, Naucleopsis, Perebea and

Pseudolmedia.The other Castilla and genera, Olmedia, are easily distinguished,

mainly on the basis of general texture, and parenchyma distribution and absence of latex tubes and Perebea respectively. Maquira are so much alike that

they can hardly be distinguished by their wood. Though the absence of dark brown contents in the and ray parenchyma cells is often characteristic for

Maquira, as it comes true for the majority of specimens, it is not sufficiently

distinctive for all Neither the samples. is usually abundant presence of dark brown contents in the parenchymatic cells of Pseudolmedia a constant and

reliablefeature that be used for may diagnostic purposes. Taking into account what is pointed out above, the following tentativekey is proposed: 10 A. M. W. MENNEGA AND M. LANZING-VINKENBORG

con- wings aliform wings sometimes short and longer rings wings wings, with rings, aliform aliform with aliform long long aliform short-aliform often often aliform distribution rather rather and seldom vasicentric and and and and lozenge-shaped with with diameter. Parenchyma vasicentric, vasicentric, incomplete vasicentric, vasicentric, vasicentric, vasicentric, complete aliform aliform fluent various f o mm (4-7) (4-8) (6-8) (4-8) (3-6) (3-8) (4-8) (4-7) (5-8) trees Number rays per 6.5 6.5 5.5 of 6 6 5 6 6 7 8 from

of tomentosa 8-10 8-9 8.5 vascular 8 8 Size pits 10 10-12 10 10 10 Helicoslylis [an 90-120 85-95 80-95 60-80 Vessel diameter 120-150 110-150 120-160 100^120 100-120 60 of in c. samples of per mm wood Number (2-13) (3-10) 5(3-7) 7(5-13) 8(4-13) 7(4-12) 10(8-13) 13(9-17)(8-18) 16(14-18) vessels square 7 7 10 between

the cm cm cm cm cm cm cm cm cm cm

Diameter 2 2 stem 5 5 5 3 1 comparison of 14 12 24 c.

A

2. 6836 9077 4539 3267 3791 3711 3672 3041 4924 Table Uw 11698 11 WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE)

1. Parenchyma in straight, moderately broad, regularly spaced, apotracheal

bands; fibres not septate or exceptionally so: latex tubes absent . Olmedia

Parenchyma vasicentric, short to long aliform, or locally or generally

aliform/confluent to confluent banded 2

2. Wood cream-coloured; weight light, c. 0.40; texture course; fibres thin-

walled with a large lumen and a diameter of 20-35 Castilla /xm ....

Wood yellowish brown to middlebrown, the heartwood sometimes darker

brown or with dark streaks; moderately heavy to heavy; fibres thick-walled

thick-walled with diameter of 15-25 to moderately a /xm 3

3. Vessels moderately numerous to numerous, 20-35 per square mm on the

diameter 70-100 fibres all with 2-3 average, (120) //m; septate septae per

fibre Naucleopsis p.p.

Vessels less numerous, 5-15 per square mm on the average, diameter 90-

150 (200) jum ; fibres usually septate but septae often absent in most fibres

in Helicostylis 4

4. Rays 4-6 per mm; vessels 100-200 nm wide, usually over 140 /mi; inter-

short-aliform vascular pits generally 8-10 /mi; parenchyma in complete, rings Helicostylis

Rays 6-9 per mm; vessels less wide, intervascular pits 5-8 /xm 5

5. Parenchyma generally aliform-confluent, forming wavy tangential bands;

often dissected few fibres Pseudolmedia rays diagonally by a (tg. sections) .

Parenchyma vasicentric, aliform, only locally confluent or forming inter-

bands bands in Perebea mollis and Perebea rupted (the more pronounced

xanthochyma) Maquira, Perebea Naucleopsis amara, N. ulei ...... ,

5. DESCRIPTION OF THE GENERA

1. CASTILLA Sessée plate I, fig. 1 2 ,

of three each wood A genus species, represented by one or more samples. (table 1).

General characters

According to Williams(1936), Record & Mell(1924), Record & Hess (1940,

1944) without clear demarcation between sapwood and heartwood; the sap-

wood light yellowish brown; texture coarse; straight-grained; rather light,

vol. weight c. 0.40.

Microscopical characters

few, 3-6 sometimes Vessels: Few to moderately (2-9) per sq. mm, nearly all

but for about radial solitary, generally 50 % in multiples of 2-4 or in longer

dumb-bell shaped rows or in irregular clusters; fairly regularly distributed;

oval rounded, outline or tang, diameter 160-200 (100-300) /xm; perforations

intervascular 8-10 the border thin hard simple; pits /xm, angular; or tyloses

vessel members the occasionally present; on average 420(200-700) /xm long.

Fibres: All in C. elastica and C. with or partly septate, tunu one or two septae,

in the materialof C. uleithe occurrence ofseptae variable; diam. 20-28 (35) /xm; 12 A. M. W. MENNEGA AND M. LANZING-VINKENBORG

walls thin, 2-3 1270 very /im; length on average (600-1900) /an; simple pits on radial walls only. Fibre/vessel member length ratio 3 (2.6-3.7).

1 and 2-6 cells wide. The uniseriates but in Rays: comprising c. 25%. scarce of two samples C. ulei; up to 30 cells (650 fitn) high, consisting of square and upright cells. The multiseriates generally 2-6 cells wide, but 3 cells wide in one

of sample C. uleL the narrow multiseriates usually with long uniseriate wings,

in the samples with the widest rays the wings 2-3 cells high, and the individual

upright cells less high than in the samples with long uniseriate wings; width

from 20-90 1 if which /zm; height up to mm or more, vertically fused, occurs

the of vessels more frequently in narrow type rays, up to 1700/zm; pits to scarce, round and half-bordered to oval, mainly in the square and upright cells. Latex

tubes in all rather in of present species, though scarce some samples C. ulei. In

of C. sclerotic cells Number one sample tunu (Uw 7046) ray present. 4-8 per mm.

Parenchyma: Paratracheal, mostly restricted to narrow vasicentric or short- aliformrings, occasionally confluent into a narrow band, however up to 8 cells

wide bands of present in C. elastica, and in one sample C. ulei nearly all

parenschyma banded, the bands 12-20cells wide (240 /mi), concentric, regular and completely enveloping the vessels. Strands of 2-4 (8) cells. Pits to vessel

large, oval, and elongated.

Diagnostic characters

All samples share the extremely thin-walled, wide fibres, and the resulting rather light weight of the wood.

Remarks

The variation in distribution of the parenchyma and in structure of the rays

shown by the present material, is also apparent from the description given by

Record & Hess (1940). Williams (1936) described the parenchyma of the

sample collected by him near Loreto as paratracheal and sometimes aliform.

2. HELICOSTYLIS Trécul plate I, fig. 3, 4; plate III, fig. 1

In Berg's revision Wood (1972) Helicostylis comprises seven species. spe- cimens from five of those were available. The type species of the genus H.

tomentosa (P. et E.) Rusby is by far the most widespread and is also the best represented in our wood collection. Several ofthe wood specimens studied were originally assigned to other genera like Pseudolmedia and Perebea as appears from table I.

General characters

Sapwood a golden yellowish brown, c. 7-10 cm wide, the heartwood light brown in H. pedunculata to dark brown in H. elegans and H. scabra very .

mediumbrown in the other species; sometimeswith dark brown streaks on the

tangential surfaces, in the cross section showing as irregular light and dark concentric zones; the demarcation between sapwood and heartwood generally

irregular. Texture moderately fine; straight- or roey-grained; moderately

hard and moderately heavy to heavy, vol. weight 0.50-0.86. 13 WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE)

Microscopical characters

Vessels: Mostly moderately few, averages ranging from 5-12.5 (2-16) per sq. of the of turbinata mm, the highest average 12.5 occurring in sample H. (Uw 16528); mostly solitary, for 10-30% in radial multiples of 2-3(5), scattered,

diameter but with a tendency towards a diagonal arrangement. Outline round,

intervascular 8-10 but 120-160 (200) /mi; perforations simple; pits (12) /an, the in one sample of H. scabra (Uw 16222) 4 pm, the borders oval or angular,

enclosed extended and Thin often Vessel slits or confluent. tyloses present. members on average 470 (430-515) /an long, ranging from 300-630pm.

Fibres: the numberof fibres variable for the Libriform, partly septate, septate

same species, in some samples of H. tomentosa and H. scabra nearly all fibres

diam. 16-20 in 20-30 walls non-septate. /an, H. pedunculata /an; occasionally

gelatinous; pits restricted to the radial walls, small; the average length 1230

the 900-1700 member ratio (1140-1380) /an. range pm. Fibre/vessel length

on average 2.6 (2.3-3.0).

cells wide. The uniseriates low, 3-5 Rays: 1- and 2-5 scarce, very mostly

(1-10) cells high, consisting of square and upright cells. The multiseriates

mostly 3-4 cells (40-50 pm) wide, but in H. pedunculata and in H. turbinata and in one sample of H. scabra 50-80 /an wide: in the other sample of H.

scabra and in 2-4 cells wide: the uniseriate H. elegans (20-40) pm wings

generally very low, however, sometimes up to 6 or 8 cells high; vertically fused

300-800 but also of rays present but not abundant, normally height pm, rays

1300-1500pm (70-90 cells) high present; sheath cells locally. Pits to vessels

half-bordered and restricted the scarce, large, irregular, or round, mainly to

and cells. Latex tubes square upright present. Crystals occasionally present; absent brown contents frequent in two samples of H. scabra, or very scarce

in the other Number 5 mm. species. on average (4-7) per broad Parenchyma: Paratracheal, usually as a moderately ring completely

surrounding the pores, and broadened laterally; in H. turbinata often con-

fluentand locally forming long uninterrupted bands 7-8 cells wide, in the other

species also locally confluent, but the bands narrower and less conspicuous;

sometimes a straight narrow band suggesting marginal parenchyma present.

Strands of 2-8 cells, mostly 4 cells. Brown contents often present, crystals

and sclerotic cells occasionally noticed. Pits to vessels large, irregular to oval,

often horizontally arranged.

Diagnostic characters

The medium-sized vessels, the short-aliform vasicentric parenchyma, and the

moderately few rays are characteristic for Helicostylis.

Remarks

Helicostylis is one of the genera of the Olmedieae which can be recognized

well anatomical features. least moderately by its At no great difficulties were

encountered in assigning a large number of Moraceous wood samples backed

herbarium this which identifications by sterile twigs to genus, were later on

confirmed by Berg.

mentionedin their of the the different Record & Mell (1924) description genus 14 A. M. W. MENNEGA AND M. LANZING-VINKENBORG

distribution of the parenchyma in materialof H. latifolia Pitt, as compared to

H. poeppigiana. Tothemthe long and narrow aliform parenchyma ofH. latifolia is of the revision suggestive genus . In Berg’s H. latifolia Pitt, was indeed transferred to Brosimum alicastrum.

3. MAQUIRA Aublet plate III,fig. 3; plate IV, fig. 1

reestablished Aublet’s genus Maquira by Berg (1972) comprises a.o. M. guia-

nensis Aubl. formerly known as Perebea laurifolia Tree., a relatively common

tree in the Guyanas. Furthermore Ducke’s genera Olmedioperebea and

of Olmedia with the of Olmediophaena. as well as all species exception O. aspera R. et P., were assigned by Berg to Maquira. The genus consists of five

of which wood table species, all are represented by samples ( 1).

General characters

Without clear demarcation of sapwood and heartwood; straw-coloured or

in of with dark in the light brown, one sample M. guianensis irregular patches central part of the stem; irregular light and dark concentric zones may be

conspicuous on the polished cross surface of a stem. Texture rather fine;

straight- or roey-grained; moderately hard; heavy, vol. weight 0.70 -0.90.

Microscopical characters

Vessels: Number variable from 6-26 per sq. mm, most numerous in M. coria-

cea but the same value of 11-23 per sg. mm also occurs in some samples of M. guianensis; mostly solitary, for 20-40 % inradial multiplesof2-3, or in clusters;

scattered or rather irregularly distributed without a definite pattern; outline oval to nearly round, tang, diameter 100-160 /tm; perforations simple; inter-

the slits enclosed; thin vascular pits on average 8 (6-11) pm, border angular,

tyloses sometimes present; length of vessel members on the average 500 (390-

580)/mi. range260- 850 /un.

Fibres: Libriform. The fibres septate with one or two septae; diameter 16-26

than the breadth of the but /on. lumen as wide as or narrower walls, in M. sclerophylla the lumen wider; occasionally with gelatinous walls; length on

the average 1225 (1100-1400) /mi, ranging from 800-1700 /mi. Fibre/vessel

memberlength ratio 2.6 (2.1 -3).

Rays: 1-seriate and (2) 3-5 (8)-seriate. The uniseriates very few, mostly about cells of and cells. The 7 high, sometimes 15-17, consisting square upright

multiseriates of variable width even within a species, but generally 3-5 cells

and (40-70 /mi) wide, widest in M. coriacea M. sclerophylla (4-8 cells); the

uniseriate wings usually 1-3 cells high, occasionally much higher; height

variable, mostly 20 60 cells (300-800 pm), but often some rays 100 cells

and those and wide dissected (1200 1400/rm) high present, high rays diagonally

by one or two fibres; vertically fused rays present but scarce; sheath cells rare. Light brown to brown contents often present, crystals scarce. Pits to

half and cells. vessels scarce, rounded, bordered, mainly in the square upright

Latex tubes present. Number on the average 6-8 per mm.

Parenchyma: Paratracheal, generally in wide to rather narrow vasicentric. WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE) 15

short-aliformrings, occasionally confluent into narrow bands, sometimes with a tendency to diagonal alignment ofthe vessels; however, in some specimens of

M. guianensis wings and bands much better developed. Pits to vessels large and irregular. Crystals rare. Strands of 2-4 (8) cells.

Diagnostic characters

The lozenge-shaped parenchyma, occasionally confluent tangentially or diagonally over a short distance is the only constant feature to distinguish species of Maquira from Pseudolmedia, which it closely resembles in most characters. Admittedly distinction between individual specimens may prove

the in the of viz. impossible due to great variability samples a species, M. guianensis. It is even more hazardous to distinguish Maquira from Perebea, while characters of the species here are often completely overlapping.

Remarks

As the reestablished genus Maquira includes so many species formerly as-

to several other related to the itself signed genera, no description genus nor to the majority of its species could be found in the literature.Record & Hess

(1940) referred to the aberrant structure of Perebea calophylla (Yw 23650

Ducke 188) now identified as M. calophylla. which did not fit eitherin Perebea

been Under the nor in Olmedia. where it also had placed. paragraphs treating

Olmedia and Olmedioperebea we also find references to species now placed in

Maquira but since their descriptions are very brief and general a comparison

with data is noticable that drew our hardly possible. In any case it is they attention to the fact that the abundant parenchyma of O. aspera is strongly

differentfromthat ofthe otherspecies they treatedunder Olmedia.

4. NAUCLEOPSIS Miquel plate III, fig. 2

the According to Berg’s (1972) conception of the genus, Naucleopsis comprises

species formerly assigned to Acanthosphaera, Ogcodeia, Palmolmedia, and two

species earlier placed in Perebea: P. concinnaand P. macrophylla. In the present

circumscription thereare 18 species in Naucleopsis, twelve ofwhich are represen-

ted by wood samples.

General characters

Wood uniformly straw-coloured to yellow-brown, heartwood not seen but

according to L. Williams (1936) dark brown in Ogcodeia tamamuri (syn. of

N. glabra). Record & Hess (1940) stated that the heartwood is pale brown and

not clearly differentiatedfrom the sapwood. Texture moderately fine, straight-

grained. Moderately hard and moderately heavy, vol weight c. 0.75.

Microscopical characters

Vessels: to the 20-35 Fairly numerous numerous, on average (16-50) per sq.

mm. but less numerous in N. amara (Uw 19340) and in N. ulei (Uw 18620)

with 13 (10-17) per sq. mm; about 10-20 (40)% in radial rows of 2-4, oc-

casionally in longer rows and then the vessels often of diminishing diameter,

or in irregular groups. Outline round to angular, diameter generally 70-100 pm. in N. amara up to 140 pm ; intervascular pits 5.5-7.5 /un, in some cases 16 A. M. W. MENNEGA AND M. LANZING-VINKENBORG

9 /zm and in N. amara and N. ulei normally 8-10 /zm. Thin tyloses seldom

the 450 300- present. Vessel members on average (400-500) /mi long, range

670 /jm.

Libriform, all fibres with2-3 fibre. Diameter Fibres: nearly septate, septate per

(12) 15-20 /zm, relatively thin-walled, walls 3-4 /an, but thicker (4-6 /an)

the radial in N. glabra; gelatinous walls frequent; pits simple, exclusively on

variable within the 950 walls; length a sample, on average (800 1060) /zm,

the range 430-1400 /zm. Fibre/vessel member length ratio on average 2.1

(1.6-2. 4).

Rays: 1- and 3-4 (6) cells wide. The uniseriates usually scarce, very low, 3-7 cells. The multiseriates in (14) cells high, with square and upright general

3-4 cells wide, the uniseriate wings with square and upright cells mostly not

over two cells but much in one of N. high, occasionally higher sample glabra ,

and width 25-50 20-60 cells in N. imitans. N. stipularis N. ulei; /zm; usually

(700-1500 /zm) high, but some up to 100 cells (2 mm)or. when vertically fused, Cell of the even 3 mm high. Pits to vessels scarce, rounded. dimensions pro- cumbent cells radial section Latex tubes as seen on a often very irregular.

usually present but not abundant. Crystals generally present as well as brown

cell contents. Average number6-8 per mm.

aliform Parenchyma: Paratracheal, vasicentric with narrow and short ex-

tensions, occasionally locally confluent over a short distance, the vasicentric ring often incomplete and absent on the adaxial side of the vessels. In one

specimen of N. guianensis and in N. macrophylla marginal parenchyma is present as a one cell wide band; in N. inaequalis, N. ternstroemiiflora and

N. ulei growth rings are sometimes indicated by a 3-5 cells wide marginal band. Strands of4 (2-8) cells. Pits to vessels large and irregular, or oval and in

horizontal cell rows. Brown contents usually present; crystals rare.

Diagnostic characters

Fairly numerous and rather narrow to moderately wide vessels, except in one

sample of N. amara and of N. ulei with less numerous vessels of a larger size.

Intervascular pits 5.5-7.5 /zm, but larger in N. amara and N. ulei. Parenchyma

inconspicuous, vasicentric, aliform, and confluent. Crystals usually present in

the rays.

Remarks

L. Williams (1936) described the general and the lens characters of three

Macbr, species of Ogcodeia (O. tamamuri N. glabra Spruce ex Pitt., O.

tessmannii Mildbr. N. glabra and O. ulei (Warb.) Macbr. N. ulei(Warb.)

Ducke). He also recorded the numerous vessels, but his statement that paren-

short lines chyma occurs both surrounding the pores and in fine, tangential

could not be confirmedas far as the lines are concerned. Record & Hess (1940)

based their of of description Naucleopsis on one specimen N. macrophylla Miq.

The affinity of the wood with that of Ogcodeia and Noyera was pointed out.

Their description of the genus Ogcodeia, based on one wood sample of each of 6 species, the names of which were not given, matches the description of

well indeed, and also with the N. macrophylla quite agrees general description 17 WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE) of Naucleopsis sensu Berg given by us. Noyera mollis (now Perebea mollis

(Poepp.) Huber) is differentfrom Naucleopsis in the less numerous and wider

vessels. Loureiro & Freitas (1968) described the lens characters of Perebea

concinna (= N. concinna (Standi.) C. C. Berg). They mentioned a pore distri-

value. bution of 7-17 per sq. mm, considerably less than our

5. OLMEDIA Ruiz & Pavon plate II, fig. 3,4; plate IV, 3

Berg (1972) considered the genus as monotypic, Olmediaaspera R. & P. being

the sole species. Specimens formerly considered to belong in this genus were

transferred by him to Pseudolmedia and Helicostylis. Three samples of O.

available for aspera were investigation.

General characters

Wood uniformly cream-coloured; texture rather fine, parenchyma clearly

visible on cross sections; slightly roey-grained. Moderately heavy, vol. weight

0.78.

Microscopical characters

for about radial Vessels: Moderately few, 7.5-10 (4-13) per sq. mm; 50% in multiples of 2-4, seldom in clusters, fairly regularly distributed.Outline round,

diameter generally 100 (70-140) /un; perforations simple; intervascular pits

angular with enclosed slit, 7-10 pm. Vessel members on the average 530 (370-

700)/un long.

Fibres: fibres in Non-septate, exceptionally one or two septate seen macera-

tions. Rather irregularly arranged. Diameter 16-22 pm, walls thin, c. 2 pni;

walls in all radial walls gelatinous present nearly fibres; pits very small, on

only; length on the average 1050 (640-1400) /jm. Fibre/vessel member length ratio 2.- (1.7-2.2).

Rays: 1- and 2-4 (5) cells wide. The uniseriates scarce, consisting of square cells The multiseriates and upright cells, very low, 2-8 high. mostly 2-4 cells

wide, the uniseriate wings often restricted to one or two rows of upright cells,

sheath cells but sometimes much longer; nearly always present, though not

completely enveloping the rays; on tangential section the cell sizes very

sometimes dissected irregular; rays vertically fused or diagonally by a fibre;

width 30-50 cells pm; up to 65 (1700/rm) high. Pits to vessels scarce, rounded and half bordered or large and elongated. Latex tubes not observed. Brown

number cell contents absent, crystals very scarce. Average 7 (5-8) per mm.

Parenchyma: Apotracheal and paratracheal. Chiefly in regularly spaced, cells straight or slightly undulating, 5-8 (12) wide bands, which occasionally border the vessels or envelop them; paratracheal parenchyma usually present

of as an incomplete vasicentric ring. Strands 2-4 cells, on average 530 pm long.

No specific cellcontents observed.

Diagnostic characters

Regularly spaced, moderately straight, broad bands of apotracheal paren-

fibres and absence of chyma, non-septate the latex tubes characterize the wood

of Olmedia. 18 A. M. W. MENNEGA AND M. LANZING-VINKENBORG

Discussion

The wood of Olmedia aspera was first described by Williams (1936) based on specimens collected by himself in Peru. The same material was subsequently studied by Record & Hess (1940) and by ourselves. Williams mentionstrunks of 20-25 cm in diameter, but the material at hand (MADw s.n.) is 5 cm in diameter.Anotherspecimen which we owe to the courtesy of Dr. R. L. Dressier

of of is a piece a stem an arching shrub (Uw 21743). It shows the same features as the Williams’ samples. Like Record & Hess remarked, the wood by its

“lamellated”structure resembles the wood of more than that of the other genera of the Olmedieae. This different wood structure combined with the absence of characteristic of branch abscission all a very type present in other genera of the Olmediaeae - anatomical details of which are described by

- Koek-Noorman & Ter Welle (1976) as well as differences in the floral structure lead Berg (1977) to exclude Olmedia from the tribe.

6. PEREBEA Aublet plate III,fig. 4; plate IV, fig. 2, 4

Berg (1972) widened the conceptof the genusto include several species assigned by previous taxonomists to other genera. Among them is the former genus

Noyera Tree, which became a section of Perebea. Of the eight species only the

the distribution three most common ones with widest are represented by wood

samples.

General characters

The wood is greyish or yellowish brown without a clear demarcation between sapwood and heartwood; straight-grained, texture moderately fine, moderately

light, the vol. weight between 0.70-0.80, much lighter (0.53) in one sample of

P. guianensis.

Microscopical characters

Vessels: Moderately few, on the average 7-8 (2-14) per sq. mm, slightly more numerous in P. guianensis (12-15 per sq. mm); mostly solitary, for about 20-

25 % in radial multiples of 2-4, fairly regularly distributed with a tendency

towards outline oval diameter a diagonal arrangement; or rounded, tang, generally 100-160 (90-200) /an, widest in P. guianensis; perforations simple;

intervascular 7-9 thin pits /an; tyloses occasionally present in P. guianensis and P. mollis; vessel memberlength on the average 460 (370-560) /mi, the total

from240 range to 680/an.

Fibres: All septate, usually with 2-3 septae; arrangement irregular. Diameter mostly 18-20(24) /an, walls 3-4 pm thick; length on the average 1260 (1050-

500-1900 limited to the radial 1460) /an. range /an; pits very small, walls. Fibre/vessel member length ratio: 2.6 (2-3.8).

Rays: Uniseriate and 2-5 cells wide. The uniseriatesconsisting of square and upright cells, rather frequent in P. guianensis and P. xanthochyma, but nearly absent in P. mollis; mostly 4-10 cells high. The multiseriates mostly 2-3 cells

(20—40 pm) wide in P. guianensis. 4-6 cells (50-70 in P. mollis and P. xanthochyma, generally with short uniseriate wings, often partly with longer WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE) 19 wings; height usually 600-800 pm, but up to 1000 and 1250 pm (50-68 cells).

Pits to vessels scarce, round, half bordered, mostly in the square and upright cells. Often contents Latex tubes present. brown in the cells; crystals scarce, noticedin P. mollis and in P. xanthochyma. Average number6.5 (5-8) per mm.

Parenchyma: Paratracheal in generally complete, narrow, aliform vasicentric rings, often confluent in tangential or diagonal direction over a short distance, the sheaths usually widest on the abaxial sides of the vessels, continuous bands

often mollis. Strands of cells. rare, though fairly present in P. 4 (2-8) Pits to vessels large, irregular, mostly horizontal. Brown cell contents often present;

noticed mollis and crystals very scarce, only in P. p.p. P. xanthochyma.

Diagnostic characters

As already stated before, it is very hard to find characters which are specific

for Perebea as and Perebea resembleeach other in all . Maquira closely nearly anatomical features, as they also do in morphological characters.

Remarks

P.Record & Hess (1940) described the wood of Noyera mollis (at present mollis mollis) and their well with ssp description complies very our findings,

the as is to be expected since specimens studied by them and by us are, at least

partly, the same. Under the heading Perebea they treated wood samples of

of viz: castilloides Pitt, 5 species, only 3 which, P. (now P. guianensis ssp. castilloides (Pittier) C. C. Berg), P. chimiqua Macbr. (now P. xanthochyma

Karsten) and P. tessmannii Mildbr. are still considered as Perebea species at present. Of the other two P. concinna Standi, was transferred by Berg to Nau- cleopsis and P. laevigata Standi, from Panama could not be traced. They mentioned the rather aberrant of with structure P. castilloides non-septate fibres and the abundant parenchyma often occurring in wide concentric bands next to the more usual short aliform vasicentric rings. This, however, need not

considered be as a really disturbing fact, since we found in our materialbanded

of mollis rather often. The absence of parenchyma in a sample P. septate fibres

in their material is more unusual as this character was always clearly present in

earlier described our samples. In an publication Record & Mell (1924) a

sample ofPerebea spec. (Yale 4952 = BW 5493) fromSuriname. The herbarium

voucherof number has been identified this now by Berg as P. mollis ssp. rubra.

In their description they mentioned the high, and up to five cells wide rays and

of bands of theoccasional occurrence parenchyma apparently limiting growth rings, but otherwise present in wing-like patches about the pores, and the often fibres. which matches septate A description quite well our general

description given above.

7. PSEUDOLMEDIA Trécul plate II, fig. 1, 2

In Berg’s revision a numberof species formerly consideredtobelong in Pseudol-

media were transferred to other like Helicostylis and as will genera Maquira , be discussed below. 20 A. M. W. MENNEGA AND M, LANZING-VINKENBORG

The material investigated consists of 18 samples belonging to 5 species (table 1)

The genuscomprises 9 species.

General characters

Colour yellowish brown, heartwood not different from the sapwood, except,

according to Record & Hess (1940, 1944), in P. spuria with reddish brown

heartwood not sharply demarcated from the thick grayish or pinkish brown

sapwood. L. Williams (1936) described the heartwood of a large tree of P.

multinervis (now P. laevis) as pinkish or dark brown. The wood is straight or

slightly roey-grained; moderately heavy, vol weight 0.75-0.80, or heavy, like

in P. oxyphyllaria with a vol. weight of 1.0.

Microscopical characters

Vessels: Number variable, on the average 10-15 per sq, mm, in samples of

small-sized trees (Krukoff 6697, 7003) often more numerous: 30 or more per

and then scattered but sometimes sq. mm resembling Naucleopsis; usually irregularly distributed, for about 30-50% in radial multiples of 2-3(4), multi-

ples less frequent in P. laevigata; tang. diam. 100-150 pm: intervascular pits

6-8 /mi; tyloses often present, generally thin, but occasionally very thick and

sclerotic; often with solid white contents in P. laevigata, P. laevis and P. oxy-

vessel member phyllaria; length 350-500 pm.

Fibre tissue; Nearly all fibres septate, except in one sample of P. macrophylla;

fibre Diameter walls thick, in arrangement irregular. 15-20 (24) pm, 3-5 pm

and walls thick and the lumen 3 P. oxyphyllaria P. spuria 6 pm narrow, c. pm:

sometimes a gelatinous layer present, pits small, simple, on the radial walls

about 1100 member only; average length (800-1340) pm. Fibre/vessel length ratio 2.60, range 2.25-3.10.

of and cells, Rays: Uniseriates scarce, short, consisting square upright some-

times also a few procumbent cells present. The multiseriates mostly h-A (2-6)

cells width to wide with uniseriate margins, 1-3 cells high; 30-50 pm and up

70 pm: height variable between 30 and 100 cells or 300 and 800 (1400) /un

the often dissected fibres respectively, large rays by one or two diagonal

(plate II, fig. 2). Pits to vessels large, mainly horizontal. Latex tubes present.

cells with brown rhombic sometimes Many contents, crystals present, very

in and P. Number the 7-8 numerous P. oxyphyllaria spuria. on average (5-11) per mm.

Parenchyma: Paratracheal, vasicentric in complete and incomplete rings, often aliform with long wings in P. laevis and P. laevigata in part of the growth ring,

but mostly confluent into irregular, wavy, narrow, 3-6 cells wide, bands;

strands of in and (2)4-8 cells; crystals present P. laevigata, P. oxyphyllaria

P. spuria.

Diagnostic characters

Paratracheal parenchyma predominantly arranged in wavy irregular bands; rather wide and often often crossed fibres; high rays, obliquely by frequent

of in and cells. occurence crystals rays parenchyma

Discussion

collected Of P. macrophylla two wood samples were available, both by Krukoff, 21 WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE)

the herbarium vouchers of which were identified by Berg. The wood structure

of of them = one (Uw 16202 Krukoff 8524) differs in various characters

from the of the described above: the fibres general structure genus as are

the 3 cells wide hardly septate, rays not over and not dissected by fibres, the

parenchyma is narrow vasicentric instead of aliform confluent.

The most extensive information on the genus to be found in the literature is

Record Hess’s based and & (1940) description chiefly on P. spuria P. oxy- phyllaria. In 3 other species, viz. P. multinervis, P. obliqua and P. scabra, they

different fibre noted a parenchyma distribution and a larger width. In Berg’s revision P. multinervis is placed in synonymy with P. laevis. Material of P.

laevis seen by us, is rather uniform and agrees quite well with the general aspect of Pseudolmedia. However, like stated above, individual variations in

structure to in happen occur occasionally each genus. P. obliqua was transferred

by Berg to Maquira coriacea and P. scabra to Helicostylis scabra. This is in accordance with their neither in in anatomy as Maquira nor Helicostylis (ex-

cept H. turbinata) parenchyma confluent over a long distance occurs.

GENERAL DISCUSSION

From the key and descriptions it becomes clear how much the genera Helicosty-

lis, Maquira, Naucleopsis. Perebea and Pseudolmediahave in common in their

wood structure. As already pointed out in the remarks following each generic

description, many of the incongruities signalized by Record & Hess (1940)

have been eliminated Berg’s of the by (1972) new concepts generic circum-

scriptions. On the other hand some inconsistencies in the general aspect of

of the still remain. Sometimes it is of several most larger genera one out

samples of the same species that shows a slightly differentwood pattern, like

Pseudolmedia 16202 = macrophylla (Uw Krukoff8524) resembling a Helicosty-

of mollis lis or two samples Perebea (Uw 8080, Uw 19268 Krukoff 6985,

Krukoff 1289) the parenchyma distributionofwhich is more like Pseudolmedia.

Or in Naucleopsis, N. amara (Uw 19340 Krukoff 1406) being different from

the other species in its numberof vessels. However, these differences are much

smaller and of a different nature than those between Olmedia and the other

genera, Olmedia being distinguished by the apotracheal banded parenchyma,

the bands being regularly spaced and more or less straight, atype of parenchyma

distributionnot occurring in any other genusof the Olmedieae, but resembling that in Ficus, Malaisia. Pseudomorus, , Sparattosyce and . To

the latter genusOlmedia seems to be related (Berg 1977). Metcalfe & Chalk

(1950) also list Trymatococcus among the genera with banded non-confluent

parenchyma, but this could not be confirmed in our material of the three

of this In Olmedia fibres which is species genus. septate are extremely rare,

another character distinguishing it from the other genera. However, in Helico-

stylis septate fibres are notof a general occurrence and in some specimens like in

H. turbinata they could not be traced at all. Moreover, in the Anatomy of

the is not cited the with fibres. Dicotyledons Helicostylis among genera septate 22 A. M. W. MENNEGA AND M. LANZING-VINKENBORG

indicating that theirpresence is not as evident as it is in the other genera. Thus, the parenchyma distributionin particular is an argument in favour of Berg’s

(1976) recent exclusion of Olmediafrom the tribe.

of his of the of Tippo (1938), in a special paragraph general survey anatomy the family, drew attention to the distribution of septate fibres within the

fibres Moraceae. From a total of 28 species with septate 25 belong to the sub- family Artocarpoideae; in all Olmedieae septate fibres are present. He also found them in two of Olmedia but as he didnot names or numbers species , give of collections it is hard to be certain whether the specimens seen by him really

to what is now considered as Olmedia[- i.e. O. R. & P. or belong aspera only -, not. Another genus that would fit in well with the Olmedieae is the African

in the Olmedieae, which followed genusAntiaris, placed by Bureau (1873) was by Bentham (1880) and more recently by Corner (1962). Fromother paleotro-

Olmedieae, pical genera which Comer tentatively placed in the Sparattosyce

would not fit. Of two more genera cited in this context, viz. and

available. no material was According to Tippo (1938) Mesogyne , possesses septate fibres which fact he stresses as being exceptional in the Dorste-

in included nieae, the tribe which Mesogyne was at that time. As a matter of fact of the beside fibres found in none tribes, Olmedieae, septate were by him,

with the exception, however, of Ficus religiosa in the Ficeae and Prainea in the

Thus the fibres of Conocephaloideae. septate Mesogyne form a strong argument

in favour of Comer’s opinion that Mesogyne is closely related to .

(The wood of Prainea shows a certain resemblance to the Olmedieae not only

in the septate fibres, but also in its aliform-vasicentric parenchyma distribu-

tion. Corner Prainea in the allianceof and Malaisia (1962) placed .

the tribe Brosimeae stressing, however, its own character.) Genera belonging to

show a certain resemblance to the Olmedieae in their parenchyma distribution

but differ other the (Lindeman & Mennega 1963) they in characters, a.o. by

complete or almost complete absence of septate fibres.

In general, the structure of the wood of the Olmedieae may be considered as advanced. Features like scalariform vessel perforations, strongly hetero-

geneous rays, diffuse apotracheal parenchyma (cf. the characters listed by

Tippo 1938), usually considered as primitive structures are not encountered in any of the genera. If septate fibres and a considerable amount of paratracheal

parenchyma are an indication of specialization then Helicostylis has to be

placed at the base of the series and Castilla at the top. The other genera are to be placed somewhere in between. 23\I WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE)

Plate I. Fig. 1,2: Castilla ulei (Uw 8827), cross sect, x 36, tg. sect, x 36; fig, 3, 4: Helicostylis

scabra (Uw 16222),cross sect. X 36, tg. sect, x 36. 24\II A. M. W. MENNEGA AND M. LANZING-VINKENBORG

Plate II. Fig. 1, 2: Pseudolmedia laevis (Uw 11817), cross sect, x 36, tg. sect, x 36; fig. 3, 4:

Olmedia 36. aspera (Uw 18429) cross sect. X 36, tg. sect, x WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE) 25\III

Plate III. Fig. 1: Heliocostylis tomentosa (Uw 6836), cross sect. X 10; fig. 2; (Uw 7549), cross sect, x 10; fig. 3: Maquira coriacea (Uw 17149) cross sect, x 36;

4: Perebea indet. 36. fig. guianensis ssp. (Uw 16183)cross sect, x 26\IV A. M. W. MENNEGA AND M, LANZING-VINKENBORG

Plate lucida coriacea various IV, camera drawings. Fig. 1; Maquira (Uw 7886), ray types,

“e” the 2: Perebea type being most common one; fig. guianensis ssp. guianensis(Uw 8241),

of which “c” and “d” the 3: Olmedia types rays among types are most frequent; fig. aspera

(Uw 21743) elements from macerated wood; fig. 4: Perebea xanthochyma (Uw 18431). 27 WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE)

ACKNOWLEDGEMENTS

Useful discussions comments with Dr. C. C. Berg as well as his suggestions and critical on the manuscript are gratefully acknowledged. Our thanks are also extended to curators of wood collections to who kindly provided us with valuable wood samples, and those who collected material in the them Dr. R. L. who im- field, among Dressier, upon our request

made effort indebted mediately a special to collect material ofOlmedia aspera. We are much to Mr. L. Y. TH. Westra for his careful correction of the English text, and to Messrs. A. Kuiper and T, Schipper for their assistance with photographs and drawings.

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