On the wood anatomy of the tribe “Olmedieae” () and the position of the Olmedia R. & P.

Alberta+M.W. MennegaandMarijke Lanzing-Vinkenborg

Instituut voorSystematische Plantkunde,Utrecht

SUMMARY

The structure ofthe wood ofthe Olmedia genera , , , , ,

Perebeaand ,considered to belongin the Olmedieae (cf. Berg 1972) is described.

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

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

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

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

which characteristic for the other - of fibres, are genera some Helicostylis excepted

- are nearly completely absent in Olmedia. This structural difference is considered as an

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

1. INTRODUCTION

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

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

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

and numberof vessels, absence of fibres, and in the distribu- or presence septate tion and quantity ofaxial parenchyma. Besides the description of the Moraceae

have Tippo’s in Metcalfe& Chalk’s Anatomy ofthe Dicotyledons (1950), we

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

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

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

proposed the new species were sometimes uncertain as to the genera to which

these species should be assigned.

In recent time several taxonomic revisions were published among which

Berg’s (1972) “Revision of the Neotropical Olmedieae and Brosimeae.”

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

vestigated the wood structure of our materialof 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 It appeared from her study (unpublished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 arrangement of the wood samples agreed remarkably well with the taxonomic arrangement.

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

until additionalmaterial in not investigation genera or poorly represented up till then would be available. As from table all appears 1 seven genera which, according to Berg (1972), constitute the tribe Olmedieae are now represented in most others the Olmedia is a satisfactory way. Among genus now represented.

In a recent paper Berg (1976) points out that the genus Olmedia must be excluded from the tribe Olmedieae for various reasons. This necessitated a redefinition of the tribe and for a new name it: C. C. Berg. For 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 I. In the table the revised names as well as the scientific wood names which accompanied the specimens received from other institutes are listed. In those cases where the material abundant was very a selection was made of the samples to be sectioned, the other ones being in-

hand spected by a x 10 lens. All wood specimens are backed by herbarium vouchers which were identifiedby Berg.

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

characters. For of the microscopical some specimens softening in hydrofluoric acid previous to cutting was necessary. Macerations were made by placing wood in mixture of of acetic acid and chips overnight a equal parts hydrogen peroxide at a temperature of 60 °C.

In the generic descriptions the averages for the length of vessel members and fibres are calculated from the average values for the species, the lowest and

between brackets. The the highest averages being placed ranges comprise lowest and highest valuesencounteredwithin the species.

3. GENERAL ANATOMICAL CHARACTERS OF THE WOOD

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

/un; vessel memberswith short tips or without tips, and a length of about 400-

The fibre tissue consists of libriformwith small the 500 /un. simple pits on radial walls. Mostly fibres are septate with 1-3 septae per fibre, but in some genera, like of the fibres Helicostylis, only part are septate and in Olmedia 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 1700 but shorter fibres 1000 /un long, with a wider body and more

in the fibres abruptly ending bayonet-like tips are also present. In Castilla are wider and thinner-walled than in the other genera; in Naucleopsis they tend to be slightly shorter than in the other genera. The ratio of fibre to vessel member 3 WOOD ANATOMY OF THE TRIBE “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 Mato Amaz. Amaz. Amaz. Mato Mato Amapa Amaz. Amaz. Amaz. Amaz. R. Guiana Guiana Collection Guatemala Panama Nicaragua ------ Suriname - Suriname - - - - Peru Braz. Braz. Peru Braz. Braz. Braz. Braz. Braz. Braz. Fr. Fr. Braz. Braz. Braz. Peru Braz. Braz.

(F.D.2757) 421 2992 19852 18613 19868 M 17967 3544 1 97 2949 267M 2507 148 1409 5766 6809 8075 8521 Lind. 165 681 7081 8762 c.s. c.s. c.s. 51670 nselaar c.s. 715986 Williams & Collectors) Englesing Ellenberg Krukoff Krukoff Krukoff Krukoff Krukoff Lindeman BAFOG BAFOG D Ellenberg Fanshawe US LI. Prance, Prance, Prance, Lanj. Pires van Krukoff Krukoff Krukoff Prance, -

of cm cm

5 5 cm cm cm cm cm cm cm 25 32 26 16 tree 32 24 tree cm 8 Diameter cm cm cm cm cm cm cm cm cm cm branch, branch, 8 1 9 the 100 12 14 14 10 12 over over over over over large over over - - - - 1 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. elastica tunu tunu elegans elegans elegans elegans elegans pedunculata pedunculata pedunculata pedunculata pedunculata pedunculata scabra scabra scabra scabra scabra tomentosa Table ulei ulei ulei ulei Helicostylis 4 A. M. W. MENNEGA AND M. LANZICG-VINKENBORG

caloph.) obliqua of; 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 locality Suriname Suriname Suriname Suriname Suriname Suriname Suriname Suriname Suriname Braz. Suriname Braz. Guiana Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Peru Venezuela Braz. Braz.

51743) (Maguire) 10963 45735 (Pires

15027 5 4076 4756 5299 5356 5357 5469 6712 2823 8859 c.s. 1557 8832 6347 6655 8580 661 6705 3726 c.s. 7178 8316 51834 (Maas) c.s. Collector(s) berg Lindeman Lindeman Lindeman Lindeman Lindeman Lindeman Lindeman Schulz Schulz Pires LBB Krukoff Maguire Prance Krukoff Krukoff Krukoff Krukoff Krukoff Krukoff Krukoff Ellen 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. E.) Berg 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 laurifolia laurifolia xinguana cone concinna ulei caloneura the temstroemiflora pallescens Distributed Perebea Perebea Perebea Acanthosphaera Ogcodeia under Olmedia 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 Fires Fires LBB Prance Lindeman Krukoff Krukoff Krukoff Krukoff Krukoff Krukoff Krukoff Krukoff Krukoff Prance Prance Lindeman

cm cm cm cm 15 10 8 cm 14 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 10 1 over over 10 10 3 28 25 17 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 guianensis C. 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. Amaz. Amaz. Mato Mato Mato Amaz. - - Mato - - - - 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 Collector(s) Lindeman Lindeman MaguireKrukoff 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 2.5 3 6 8 4.5 5 5 5 4.5 the 3.5 10.5 ca. 13 big 4.5 ca. 4 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. (Mildbr.) C. (Standi.) Pav. Miq. Ducke (Ducke) Ducke et (Ducke) Berg (continued) Berg Ruiz, C. (Warb.) 1 krukovii(Stand.)C.C. C. guianensis guianensis guianensis guianensis imitans inaequalis macrophylla macrophylla mello-barreto mello-barretoi stipularis stipularis lemstroemiiflora C. ternstroemiiflora temstroemiiflora aspera aspera aspera Table C. ulei uiei ulei ulei Olmedia WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE) 7

of:

name tessmannii tessmannii pseudopeltata mollis mollis mollis ulei? ulei? the brosimifolia Distributed Perebea Ogcodeia Ogcodeia under Perebea Perebea Noyera 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. Amaz. Amaz. - - - - - Mato- - - Collection locality Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Braz. Suriname Colombia Peru Braz. Braz. Braz. Colombia Colombia Colombia Colombia Colombia Colombia

18131 18525 s.n. 3412 7041 7209 48718 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) KrukolT Krukoff Irwin Prance KrukolT Ducke Krukoff Krukoff Prance, Stahel Cuatracasas Ducke Krukoff Krukoff LI. de de de de de de

cm cm tree -16 14 cm cm cm cm cm cm cm cm cm cm cm cm

Diameter 8 5 4 4 shrub 7 the over 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 (Tréc.) guian. guian. guian. interned. Huber Karsten Aublet guian. E.) mollis mollis mollis rubra ssp. ssp. ssp. ssp. Tréc. (continued) guianensis & Berg (P mollis 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.

1 152 35 1861 5378 5034 18437 18610 18376 119 8383 3929 5188 5217 5312 5440 10738 8524 INIF 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 Diameter big over 1 over the big 10 30 big big 14 14 big 10 ?

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

Sm. Macbr. Tree. Dorm. P.) Griseb. (continued) et only (R. (Sw.)

I laevigata laevigata laevis laevis laevis laevis laevis laevis laevis laevis laevis laevis macrophylla macrophylla macrophylla oxyphyllaria spuria Sections Table +oxyphyllaria +spuria 4 + WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE) 9

length is moderately high as it ranges between2-3. In the ray tissue the 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 uniseriatewings, the highest rays up to 2.6 mm high (165 cells), are often

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

vessels, consequently ray/vessel pitting is rare and restricted to occasional half- bordered round pits in the procumbent cells and larger oval pits in the 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 lateral wings locally confluent or even forming confluent banded parenchyma.

terminal band is A narrow 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

vessels oval in In Olmedia to large, or irregular, mainly horizontalarrangement. the parenchyma distribution is different, the parenchyma occurring in wide, regular, apotracheal bands.

4. KEY TO THE WOOD OF THE OLMEDIEAE

It was not without hesitation that a key to the genera was prepared. In this 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 genus. Aberrant features are usually found in the parenchyma distribution

the vessel number. and average From a detailed investigation of the material of (table 2) it appeared that stems and branches with a

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

Therefore from mature stems. samples such small-sized stems or branches cannot be identifiedwith much certainty.

All this applies to the genera Helicostylis, Maquira, Naucleopsis, Perebea and

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

mainly on the basis of general texture, and parenchyma distribution and absence of latex tubes respectively. Maquira and Perebea are so much alike that they can hardly be distinguished by their wood. Though the absence of dark brown contents in the ray and parenchyma cells is often characteristic for

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

distinctive for all samples. Neither is the 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 rays Number per of 6 6.5 6.5 6 5 6 6 5.5 7 8 from

of tomentosa 8-10 8-9 8.5 vascular 8 8 Size pits 10 10-12 10 10 10 Helicostylis jun 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 (2-13) 7(3-10) 5(3-7) 7(5-13) (4-13) 7(4-12) 10(8-13) 13(9-17) 10(8-18) 16(14-18) Number 7 vessels square 8 between

Diameter the cm cm cm cm cm cm cm cm cm cm stem 5 5 5 3 2 2 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

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

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

aliform/confluent to confluent banded 2

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

walled with lumen and diameterof 20-35 Castilla a large a pm ....

Wood yellowish brown to middle brown, the heartwood sometimes darker

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

to moderately thick-walled with a diameterof 15-25 /an 3

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

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

fibre Naucleopsis p.p.

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

150 (200) pm: fibres usually septate but septae often absent in most fibres

in Helicostylis 4

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

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

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

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

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

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

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

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

5. DESCRIPTION OF THE GENERA

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

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

General characters

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

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

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

vol. weight c. 0.40.

Microscopical characters

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

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

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

outline oval or rounded, tang, diameter 160-200 (100 300) pm : perforations

simple; intervascular pits 8-10 /an. the border angular; thin or hard tyloses

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

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

in the materialof C. ulei the occurrence ofseptae variable; diam. 20-28 (35) pm ; 12 A. M. W. MENNEGA AND M. LANZING-VINKENBORG walls 2-3 1270 very thin, pm; length on average (600-1900) pm; simple pits on radial walls only. Fibre/vessel memberlength ratio 3 (2.6-3.7).

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

multiseriates upright cells. The generally 2-6 cells wide, but 3 cells wide in one sample of C. ulei, the narrow multiseriates usually with long uniseriate wings, in the with the widest the 2-3 cells and the individual samples rays wings high, upright cells less high than in the samples with long uniseriate wings; width from 20 90 if which //m; height up to 1 mm or more, vertically fused, occurs more in the narrow of to 1700 to vessels frequently type rays, up fan ; pits scarce, round and half-borderedto oval, mainly in the square and upright cells. Latex tubes present in all species, though rather scarce in some samples of C. ulei. In

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

Parenchyma: Paratracheal, mostly restricted to narrow vasicentric or short- aliform confluent into narrow band, however 8 cells rings, occasionally a up to wide bands present in C. elastica, and in one sample of 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 ratherlight weight of the wood.

Remarks

of The variation in distribution of the parenchyma and in structure 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 Trecul plate I, fig. 3 4; plate III, fig. 1 ,

In Berg’s (1972) revision Helicostylis comprises seven species. Wood spe-

from five of those available. The of the cimens were type species genus H.

tomentosa (P. et E.) Rusby is by far the most widespread and is also the best represented in our wood collection. Several of the 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 and brown in H. pedunculata to very dark in H. elegans H. scabra, medium brown in the other species; sometimes with darkbrown streaks on the

in the section tangential surfaces, cross showing as irregular light and dark concentric the between and heartwood zones; demarcation sapwood generally irregular. Texture moderately fine; straight- or roey-grained; moderately

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

Microscopical characters

from 5-12.5 Vessels: Mostly moderately few, averages ranging (2-16) per sq.

of in the of H. turbinata mm, the highest average 12.5 occurring sample (Uw 16528); mostly solitary, for 10-30% in radial multiples of 2-3(5), scattered,

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

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

and confluent. Thin often Vessel slits enclosed or extended tyloses present. members 470 from 300-630 on average (430-515) /rm long, ranging µm. the number of fibres variable for the Fibres: Libriform, partly septate, septate in all fibres same species, some samples of H. tomentosa and H. scabra nearly non-septate, diam. 16-20 pm. in H. pedunculata 20-30pm: walls occasionally

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

the 900-1700 member ratio (1140-1380) /im. range pm. Fibre/vessel length on average 2.6 (2.3-3.0).

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

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

mostly 3-4 cells (40-50 /un) wide, but in H. pedunculata and in H. turbinata

and in one sample of H. scabra 50-80 /un wide; in the other sample of H.

scabra and in 2-4 cells wide; the uniseriate H. elegans (20-40) /un wings

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

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

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

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

square and upright cells. Latex tubes present. Crystals occasionally present;

brown in of scabra absent contents frequent two samples H. or very scarce ,

in theother Number 5 species. on average (4-7) per mm.

Parenchyma: Paratracheal, usually as a moderately broad ring completely

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

Remarks

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

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

encountered in assigning a large number of Moraceous wood samples backed

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

confirmed by Berg.

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

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

of H.poeppigiana. To themthe long and narrow aliform parenchyma H. latifolia

is suggestive of the genus . In Berg’s revision H. latifolia Pitt, was indeed transferred to Brosimum alicastrum.

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

Aublet’s reestablished Berg a.o. M. genus Maquira by (1972) comprises guia- nensis Aubl. formerly known as Perebea laurifolia Tree., a relatively common

tree in the . Furthermore Ducke's genera Olmedioperebea and

Olmediophaena. as well as all species of Olmedia with the exception of O.

consists of five aspera R. et P.. were assigned by Berg to Maquira. The genus

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

General characters

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

light brown, in one sample of M. guianensis with irregular dark patches in the 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 numerous in M. per sq. mm, most coria-

cea but the same value of 11-23 per sg. mm also occurs in some samples of M.

for 20-40 in radial of in guianensis; mostly solitary, % multiples 2-3, or clusters;

rather distributed without definite outline scattered or irregularly a pattern; oval to nearly round, tang, diameter 100-160pm ; 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) pm. range 260 850/urn.

Fibres: Libriform. The fibres with diameter 16-26 septate one or two septae; pm. lumen as wide as or narrower than the breadth of the walls, but in M.

the lumen with sclerophylla wider; occasionally gelatinous walls; length on

the 1225 1400) from 800-1700 average (1100 /un, ranging pm. Fibre/vessel memberlength ratio2.6 (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

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

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

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

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

(1200-1400 pm) high present, and those high and wide rays dissected diagonally

by one or two fibres; vertically fused rays present but scarce; sheath cells

often rare. Light brown to brown contents present, crystals scarce. Pits to

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

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

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

into sometimeswith short-aliformrings, occasionally confluent narrow bands,

of the in of a tendency to diagonal alignment vessels; however, some specimens

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

of from Pseudolmedia which it resembles in most species Maquira , closely

characters. Admittedly distinction between individual specimens may prove

impossible due to the great variability in the samples of a species, viz. M.

from guianensis. It is even more hazardous to distinguish Maquira Perebea,

while characters of the species here are often completely overlapping.

Remarks

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

signed to several other genera, no description related to the genus itself 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 =

which did fiteither Ducke 188) now identified as M. calophylla, not in Perebea

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

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

but since their brief and Maquira descriptions are very general a comparison drew with our data is hardly possible. In any case it is noticable that they

attention the fact the of is to that abundant parenchyma O. aspera strongly differentfrom that ofthe other species they treated underOlmedia.

4. NAUCLEOPSIS Miquel plate III, fig. 2

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

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

species earlier placed in Perebea: P. concinna and P. macrophylla. In thepresent

there 18 in ofwhich circumscription are species Naucleopsis, twelve arerepresen-

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 hardand moderately heavy, vol weight c. 0.75.

Microscopical characters

Vessels: Fairly numerous to numerous, on the average 20 35 (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, in Outline round or irregular groups. to angular, diameter generally 70-100

in to 140 intervascular /mi, N. amara up fim; pits 5.5-7.5 /an, in some cases 16 A. M. W. MENNEGA AND M. LANZING-VINKENBORG

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

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

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

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

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

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

member ratio the 2.1 range 430-1400 pm. Fibre/vessel length on average (1.6-2.4).

low, Rays: 1- and 3-4 (6) cells wide. The uniseriates usually scarce, very 3-7

(14) cells high, with square and upright cells. The multiseriates in general cells 3-4 cells wide, the uniseriate wings with square and upright mostly not

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

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

(700-1500 |xm) high, but some up to 100 cells (2 mm) or, when vertically fused,

vessels rounded. Cell dimensions of the even 3 mm high. Pits to scarce, pro-

cumbent cells as seen on a radial section often very irregular. Latex tubes

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

cell contents. Average number6-8 per mm.

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

tensions. occasionally locally confluent over a short distance, the vasicentric

of the vessels. ring often incomplete and absent on the adaxial side In one

specimen of N. guianensis and in N. macrophylla marginal parenchyma is

cell wide in N. N. and present as a one band; inaequalis, ternstroemiiflora

sometimes indicated 3-5 cells wide N. ulei growth rings are by a marginal

band. Strands of 4 (2-8) cells. Pits to vessels large and irregular, or oval and in

horizontal rows. Brown cell 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 pm. 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

species of Ogcodeia (O. tamamuri Macbr. = 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-

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

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

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

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

of the based on one wood of each Their description genus Ogcodeia, sample

the of which matches the of of 6 species, names were not given, description

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

is different from in the less and wider (Poepp.) Huber) Naucleopsis numerous vessels. Loureiro & Freitas (1968) described the lens characters of Perebea

mentioned concinna (= N. concinna (Standi.) C. C. Berg). They a pore distri- bution of 7-17 less than value. per sq. mm, considerably our

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

Berg (1972) considered the genus as monotypic, Olmedia aspera 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. aspera were available for investigation.

General characters

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

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

0.78.

Microscopical characters

Vessels: Moderately few, 7.5-10 (4-13) per sq. mm; for about 50% in radial multiples of 2-4, seldom in clusters, fairly regularly distributed.Outline round, diameter generally 100 (70-140) pm; perforations simple; intervascular pits angular with enclosed slit, 7-10 pm. Vessel members on the average 530 (370-

700) pm long.

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

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

walls in all radial walls gelatinous present nearly fibres; pits very small, on only; length on the average 1050 (640 1400) pm. 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

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

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

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

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

width 30-50 cells ; up to 65 (1700 pm) high. Pits to vessels scarce, rounded

and half bordered or large and elongated. Latex tubes not observed. Brown

cell contents absent, crystals very scarce. Average number 7 (5-8) per mm. Parenchyma: Apotracheal and paratracheal. Chiefly in regularly spaced,

straight or slightly undulating, 5-8 (12) cells wide bands, which occasionally

border the vessels or envelop them; paratracheal parenchyma usually present

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

No specific cellcontents observed.

Diagnostic characters

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

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

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

Discussion

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

of the Olmedieae. This different wood structure combined with the genera absence of a very characteristic type of branch abscission present in all other

of the Olmediaeae - anatomical details of which described genera are 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) widenedthe concept ofthe genusto include several species assigned by previous taxonomists to other genera. Among them is the former genus

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

three most common ones with the widest distribution 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 a diagonal arrangement; outline oval or rounded, tang, diameter

generally 100 160 (90 200) pm, widest in P. guianensis; perforations simple;

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

from 240 680 range to pm.

Fibres; All septate, usually with 2-3 septae; arrangement irregular. Diameter

mostly 18-20 (24)pm, walls 3-4 pm thick; length on the average 1260 (1050-

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

Pays; Uniseriate and 2-5 cells wide. The uniseriates consisting 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 pan) wide in P. guianensis, 4-6 cells (50-70 pm) 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 vessels round, half in the and to scarce, bordered, mostly square upright cells. Latex tubes present. Often brown contents in the cells; crystals scarce, noticedin P. mollis and in P. xanthochyma. Average number6.5 (5-8) per mm.

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

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

Diagnostic characters

As already stated before, it is very hard to find characters which are specific for Perebea, as Maquira and Perebeaclosely resembleeach other in nearly all anatomical features, as they also do in morphological characters.

Remarks

described the wood of mollis Record & Hess (1940) Noyera (at present p.

mollis) and their well mollis ssp. description complies very with our findings, as is to be expected since the specimens studied by them and by us are, at least

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

of castilloides 5 species, only 3 which, viz: P. Pitt, (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

Of the other Standi, transferred present. two P. concinna was by Berg to Nau- cleopsis and P. laevigata Standi, from Panama could not be traced. They mentioned the rather aberrant structure of P. castilloides with 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 be considered as a really disturbing fact, since we found in our material banded parenchyma in a sample of P. mollisrather often. The absence of septate fibres in theirmaterial 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 voucher of this number has now been identified by Berg as P. mollis ssp. rubra.

their mentioned the and five cells wide In description they high, up to rays and the occasional occurrence of bands of parenchyma apparently limiting growth

otherwise rings, but present in wing-like patches about the pores, and the often fibres. A which matches well septate description quite our general description givenabove.

7. PSEUDOLMEDIA Trecul plate II. fig. I. 2

In Berg's revision a numberof species formerly considered to belong in Pseudol- media transferred to other were genera like Helicostylis and Maquira, as will be discussed below. 20 A. M. W. MENNEGA AND M. LANZING-VINKENBORG

The materialinvestigated 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, the 10-15 in of on average per sq. mm, samples

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

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

less in P. diam. 100-150 intervascular ples frequent laevigata; tang. /im; pits often thin, but thick and 6-8 /im; tyloses present, generally occasionally very

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

vessel member 350-500 phyllaria; length pm. Fibre tissue: Nearly all fibres septate, except in one sample of P. macrophylla;

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

P. oxyphyllaria and P. spuria walls 6 pm thick and the lumennarrow, c. 3 pm;

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

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

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

times also a few procumbent cells present. The multiseriates mostly 3-4 (2-6)

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

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

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

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

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

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

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

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

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

P. spuria.

Diagnostic characters

bands; Paratracheal parenchyma predominantly arranged in wavy irregular

rather wide and often high rays, often obliquely crossed by fibres; frequent

of in occurence crystals rays and parenchyma cells.

Discussion

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

the herbarium vouchers of which identified The wood were by Berg. structure of one of them (Uw 16202 = Krukoff 8524) differs in various characters

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

the not cells wide hardly septate, rays over 3 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 (1940) description chiefly based on P. spuria and P. oxy- phyllaria. In 3 other species, viz. P. multinervis, P. obliqua and P. scabra, they

different distribution and fibre noted a parenchyma a larger width. In Berg’s

revision P. multinervis is placed in synonymy with P. laevis. Material of P.

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

in each structure happen to occur occasionally genus. P. obliqua was transferred by Berg to Maquira coriacea and P. scabra to Helicostylis scabra. This is in accordance with their anatomy as neither in Maquira nor in Helicostylis (ex-

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

GENERAL DISCUSSION

From the key and descriptions it becomes clear how much the genera Helicosty- lis, Maquira, Naucleopsis, Perebea and Pseudolmedia have 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 by (1972) new concepts the generic circum-

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

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

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

Pseudolmediamacrophylla (Uw 16202 Krukoff8524) resembling a Helicosty-

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

Krukoff 1289) the parenchyma distributionof which is more like Pseudolmedia.

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

the other in its numberof vessels. these species However, 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 and less of being regularly spaced more or straight, a type parenchyma

distributionnot occurring in any other genusof the Olmedieae, but resembling

that in Ficus, Malaisia, Pseudomorus, , Sparattosyce and . To

the latter Olmedia to be related Metcalfe genus seems (Berg 1977). & 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

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

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

of like stylis septate fibres are not a general occurrence and in some specimens in

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

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

that their is evident it is in the other indicating presence not as as genera. Thus,

the parenchyma distribution in particular is an argument in favour of Berg’s

(1976) recent exclusion of Olmedia from 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

Moraceae. From a total of 28 species with septate fibres 25 belong to the sub- family Artocarpoideae; in all Olmedieae septate fibres are present. He also found them in two species of iOlmedia but as he did not give names or numbers ,

hard be whether the him of collections it is to certain specimens seen by really

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

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

pical genera which Corner tentatively placed in the Olmedieae, Sparattosyce

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

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

the tribe in which included that time. nieae, Mesogyne was at As a matter of

fact in of the beside fibres found 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 form Conocephaloideae. septate Mesogyne 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 (1962) placed Praineain the alliance of and Malaisia ,

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

show a certain resemblance to the Olmedieaein their parenchyma distribution

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

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-

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

Tippo 1938), usually considered as primitive structures are not encountered in

of the If fibres and considerable of any genera. septate a amount paratracheal

parenchyma are an indication of specialization then Helicostylis has to be

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

Plate I. 2: Fig. 1, Castilla ulei cross (Uw 8827), sect. x 36, tg. sect. x 36; fig. 3, 4:Helicostylis scabra (Uw 16222), cross sect. x 36, tg. sect. x 36. 24\II V 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 sect. aspera (Uw 18429)cross sect. x 36, tg. x 36. “OLMEDIEAE" 25\III WOOD ANATOMY OF THE TRIBE (MORACEAE)

Plate III. Fig. 1: Heliocostylis tomentosa (Uw 6836), cross sect. X 10; fig. 2: Naucleopsis

glabra (Uw 7549), cross sect. x 10; fig. 3: Maquira coriacea (Uw 17149) cross sect. x 36;

Perebea indet. 36. fig. 4: 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,

Perebea type “e” being the most common one; fig. 2: 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 maceratedwood; fig. 4: Perebea xanthochyma (Uw 18431). WOOD ANATOMY OF THE TRIBE “OLMEDIEAE” (MORACEAE) 27

ACKNOWLEDGEMENTS

Useful discussions with Dr. C. C. Berg as well as his suggestions and critical comments on

the manuscript are gratefully acknowledged. Our thanks are also extended to curators of

wood collections who kindly provided us with valuable wood samples, and to those who collected in the Dr. material field, among them R. L. Dressier, who upon our request im-

made effort collect material ofOlmedia We much indebted mediately a special to aspera. are

Mr. L. Y. TH. Westra for his careful correction of the and to English text, to Messrs. A.

Kuiper and T. Schipper for their assistance with photographs and drawings.

REFERENCES

Bentham, G. (1880): Urticaceae. In Bentham & Hooker, Genera Plantarum 3 (1): 341-395. C. C. Olmedieae, Flora Berg, (1972): Brosimeae (Moraceae). Neotropica. Monographno. 7. Hafner Publ. Comp. New York.

— The of (1977): Castilleae, a tribe the Moraceae, renamed and redefined due to the ex-

of the Olmedia from the Acta Bol. 26: 73-82. clusion type genus “Olmedieae”. Need.

Bureau, E. (1873): Artocarpaceae. In De Candolle, Prodromus Systematis Naturalis Regni Vegetabilis 17: 280-288.

Corner, E. J. H. (1962): The classification of Moraceae. Gard. Bull. Singapore 19: 187-252.

Koek-Noorman, J. & B. J. H. ter Welle (1976): The anatomy of branch abscission layers

in Perebea mollis and Naucleopsis guianensis(Castilleae, Moraceae). Leiden Bot. Series 3 (in press).

Lindeman, J. C. & A. M. W. Mennega (1963): Bomenboek voor Suriname, ’s Lands Bos-

beheer. Paramaribo.

Loureiro,A. & M. Freitas da Silva (1968): Catalogo das Madeiras da Amazonia. Belem. Metcalfe,C. R. &L. Chalk (1950): Anatomyof the Dicotyledons. Clarendon Press. Oxford.

Record, S. J. & R. W. Hess American Woods ofthe Moraceae. Woods (1940): family Prop. ,

no. 61: 11-54.

— & — (1944): Timbers of the New World. Yale University Press, New Haven.

— & C. D. Mell (1924): Timbers of Tropical America. Yale University Press, New Haven.

Tippo, O. (1938) Comparative anatomy ofthe Moraceae and their allies. Bot. Gaz. 100; 1-99.

Williams,LI. (1936): Woods ofNortheastern Peru. Field Mus. Nat. Hist. Bot. Ser. 15: 71-95.