Pollen of Sarawakodendron (Celastraceae) and
some related genera,
with notes on techniques
Ding Hou
Summary
A for of small of material is described 1. simple technique acetolysis quantities polliniferous (herbarium) and notes on pollen photomicrography are presented.
2. Pollen grains of Sarawakodendron and six related genera, consisting of twenty-nine mostly Malesian species, have been examined and recorded.
3. The result of pollen study on Kokoona and Lophopetalum agrees with the generic delimitation based on gross morphology. least four have been found in the examination of all the 4. At pollen types genus Lophopetalumon species involved.
The of Sarawakodendron shows resemblance that of the related 5. pollen a great to genera Xylonymus and Kokoona.
6. The pollenofHedraianthera and Brassiantha resembles that of Sarawakodendron,Kokoona , and Xylonymus in aperture configuration, but differs in sculpture and shows in this respect similarity to the pollen of the African Salacighia.
In Kokoona of reticulate correlated with anther characters. This 7. coarseness sculpture appears genus also be from its can easily distinguished Lophopetalum by single pollen grains.
8. Parallels are found between the pollen types in Lophopetalum and those in Hippocratea (sens. str.).
Introduction
has been shown of the Sarawakodendron closely allied to Salacia Hippocrateaceae on one hand andto KokoonaandLophopetalum ofthe Celastraceae on the other by its gross morphol- link between these related families which ogy. It represents an important two closely have Celastraceae already been treated as one family, the (Hou 1963, 1964, 1967; Robson examined 1965). The pollen grains of Sarawakodendron were already by myself, and it be if those would interesting of its related genera, e.g. Salacia, Kokoona, Xylonymus, could also be studied for Hedraianthera, Brassiantha, and Lophopetalum, etc., comparison.
A. C. Smith and I. W. Bailey (1941), in connection with their studies on Brassiantha of A. C. Sm., stated that the pollen of the Celastraceae (sensustr.) exhibits a similar range that of variability in size, form, and structure as the Hippocrateaceae. Erdtman (1952) also mentionedthat similar in more or less pollen grains occur these two families. Later, other in addition to the publications concerning pollen of these families by workers,
Van Campo and Halle (1959) made a comprehensive study of African representatives of the with the of in of Hippocrateaceae very interesting finding grains polyads consisting their has with four tetrads in Hippocratea (sensu Hallé 1962); work been illustrated forty plates of beautiful drawings and photographs facilitating comparison. Recently,
Waanders, Skvarla, and Pyle (1968) studied the fine structure of pollen walls of four
American Hippocrateaceae; they recorded polyads consisting of 4 tetrads in Hippocratea and volubilis L. (erroneously ascribed to 'Stehle Quentin').
97 98 BLUMEA VOL. XVII, No. I, 1969.
the So far, only pollen grains of a few representatives of the Celastraceae (sensu str.) connection with taxonomic work for the Flora I have been studied. In my Malesiana,
could collect polliniferous samples of all the species accepted by me. of familiarize In order to acquire some reading knowledge pollen morphology and
myself with its terminology and techniques for preparations and photomicrography, I those mentioned started to work on the pollen of genera above. I have not studied pollen African ofSalacia and have relied for comparisons on the descriptions and figures ofWest
species published by Van Campo & Halle (1959). of of I am fully aware of the limitations using simple equipment and my restricted for confine ability, being a taxonomist, studying pollen morphology. So I just myself
to the description of those features of pollen grains which have taxonomic significance of information available. and to the presentation the of In the course of this study, literature on techniques pollen preparations and
photomicrography has been reviewed. After having tested rather simple equipment, I and made have prepared a working procedure for do-it-yourself acetolysis some notes those on photomicrography. I hope that this information will be of use to interested in
such techniques.
DO-IT-YOURSELF ACETOLYSIS
For pollen preparations the fundamentalprinciples of the well known acetolysis method have devised by Erdtman (1943, 1952, i960) have been followed (cf. Traverse 1965). I the small tried to reach goal by using simple equipment, quantities of chemicals and, above all, a minimumof pollen bearing (herbarium) material, as in the 'micro-method' and applied and described by Punt (1962) some others before him (cf. Faegri & Iversen have 1964, p. 79). By trial and error, satisfactory microscopic slides been obtained. My
working procedure is as follows:
or anther evena of soften the material (1) Boil a flower, a flower-bud, an (or part it) to for dissection.
the material in in few (2) Put boiled a watch-glass (about 7 cm diam.), containing a under the binocular. If the anthers drops of (distilled) water, are big enough, remove from small the pollen grains them by using a pair of fine forceps and a needle. For very and break the material into For anthers, take one or more of them pieces. type or scant
sometimes obtain and still be able save the material, one may even enough grains to empty anther(s) or flower(s). the thenadd about fresh solution (3) Let the pollen in watch-glass dry, 1\ cc acetolysis of of and of (a mixture 9 parts acetic anhydride 1 part conc. sulphuric acid, having mea-
a with scales marked down to 0.1 cc). sured each acid with respective pipette, 5 cc, kinds be used. the which (4) Two of heating apparatus can Try one is more convenient
to you.
with (a) Alcohol lamp or Bunsenburner. Hold the watch-glass containing pollen grains rather low flame of alcohol Bunsen- a pair of coarse forceps rotating over a an lamp or dark brown. burner until the solutionturns brown or Keep the solution below boiling point. Then proceed to 5). the (b) Electric tea-warmer. In order to standardize acetolysis process, an electric tea- has been used. has switch for and hot and warmer It a two-way warm (60 W) (120 W), is with marked scale from connected to themain by an Electric EnergyRegulator a 0—10.
For rather with a metal thermometer fixed obtaining a even temperature, a copper plate, to it, is placed on the warmer. (Fig. 1). Pollen Sarawakodendron and related Ding Hou: of (Celastraceae) some genera 99
Fig. 1. Heating apparatus and slide mounting card.
Place the watch-glass containing pollen material on thewarmer. Turn on the Regulator the scale of with the switch of the warmer on 'hot'. Set the Regulator to let the tempera- about ture gradually rise from room-temperature to 100° C in 15 minutes. The glass the should be rotated once and again. After the temperature has reached 100° C, adjust the Regulator to keep temperature at that degree for about three minutes and then turn
it off.
(5) Remove the watch-glass from the flame or warmer for cooling and then place it
under the binocular to examine the grains which would appear brown or dark brown
colour ofthembecomes after If take ofthe solution (the lighter washing). necessary, a drop
from the glass, prepare a slide, and thenexamine the grains under the microscope. With the some experience, one may judge the conditionof acetolysis by taking a look at colour of the solution left in the glass.
(6) Add some alcohol (about 95 %) to the solution in the watch-glass with an eye-
it is wash acid dropper and rotate for washing. It unnecessary to with glacial acetic After few (Erdtman, i960)! a minutes, the grains gather together in the center of the
and at the a ofsolution will surface. a glass same time rim appear on its upper inner Use of soft for off the rim and the solution in the piece paper wiping drawing away glass as
much as possible. Wash the pollen at least once more.
of alcohol and mixture (7) Add a few more drops one or two drops of a of 1 part of and ofalcohol the and glycerine 1 part (about 95 %) to solution rotate it. (8) After the alcohol has evaporated, the grains will either stick to the watch-glass coated of the with a layer glycerine or gather together in the glycerine in center. They are ready to be transferred to a slide for mounting. If the and theircolour dark which grains are over-acetolyzed appears too may obscure 100 BLUMEA VOL. XVII, No. i, 1969.
features for observation, add a few more drops of the mixture of glycerine-alcohol to and with dish. After of the the watch-glass cover it a Petri one day or more, the colour
willbe should grains faded justlike bleached ones. One realize, however, that theoxidation does clear the exines, but at the same timewill also cause the grains to swell (cf. Faegri &
Iversen, 1964, p. 79).
(9) Attach a small piece ofglycerine jelly to a needleand touch the grains in the watch- glass.
Glycerine jelly has been used as mounting medium. For convenience, it is melted in a thin about Petri dish (about 6 cm in diam.) to form a layer ij mm thick and then cut with a razor blade into strips each about i| mm wide. Whenever needed, use a needle to cut off a piece of the size desired. besides If there are enough acetolyzed grains, using glycerine jelly, sometimes one may
use as medium. This kind of has been try to methyl-green glycerine jelly mounting jelly prepared according to the procedure of Wodehouse (1935) (cf. Staining Procedures,
i960) by adding saturated methyl-green solution (in alcohol 50 %) gradually, drop by
drop, to the melted glycerine jelly.
For staining the grains, I have triedto use safranin (1 %; prepared with alcohol or water). of safranin the the Just add a few drops to washed, acetolyzed grains in watch-glass. After a few minutes, wash them with alcohol or water and then follow the procedure for mounting.
card. card Erdtman I made Mounting Prepare a mounting (Fig. i) as designed by (i960). alterations of it for convenience. of some minor my personal Cut a piece L-shaped card- wide and with each Glue this 'L' cardboard board, 2| cm outer edge io|- cm long. to a of 11 X 11 cm with one arm along the top and the other on the left side. Draw a vertical
line from inner of the mark it on the card 42 mm the edge left arm and on one '+' at
and respectively. From the inner edge of the draw three hori- 13, 39, 65 mm, top arm,
zontal lines at and for the ofthree slides. Place a 26, 52, 78 mm position (round) cover slip
on each '+' and mark its size and shape on the card. Place slide the card and then the of with (10) a on mounting put piece jelly grains on the slide above the marked
in for If (11) Put some paraffin (melting point about 55° C. a small beaker melting. the and the the warmer is used, turn on Regulator adjust it to keep paraffin in a melting
state.
slide the flame of the it the (12) Hold the over lamp or burner, or put on warmer.
When thejelly on the slide is melting, stir it with a warm needleif necessary to make the
less grains more or evenly distributed. Usually some air bubbles appear in the jelly con- be able rid of the taining the grains. However, with some experience, one may to get bubbles by using a needle, carefully taking them out one by one under the binocular.
18 the (13) Put a cover slip (nr o, round, mm in diam.) on jelly above the marked the card. Hold the slide the flame the position on over or place it on warmer. Use a the lower end should be the short pipette (the pore at not too small) to put melting
the of the its the at the paraffin near edge cover slip, adjusting quantity by finger upper
end, so that just enough paraffin will run under the cover slip around the jelly. order Sometimes, in to have the grains in various positions to facilitate studying or
photographing them from all sides, especially grains with unequal polar and equatorial
axes, or for avoiding too thin mounts to prevent both drying-out and crushing, a granule of clay has been used beside the jelly by Punt (1962), or small cover-slip splitters have beenadded to the preparation by Faegri and Iversen (1964). However, I place two pieces of human hair in appropriate thickness one on each side of the jelly parallel to the long Ding Hou: Pollen Sarawakodendron and related of (Celastraceae) some genera 101
of axis the slide; put on a cover slip and then seal it in the same way as the one without the be slide hair. There would no difficulty in using such a for making observations
under oil immersion.
about (14) Put the sealed slide upside-down on a rectangular frame, 22 X X 5 cm,
in the form of a box without lid and bottom as designed by Erdtman (i960), for the
surface the paraffin to harden, so the grains may come to lie close to the of cover slip. The of the size jelly on a final slide should not be too large; it is convenient to have it about 5 mm in diameter. Finally, use a razor blade to remove the excess paraffin.
size (15) Label the slide, e.g. by using a square gummed paper of suitable and record on the collector's and it necessary information, i.e. scientific names, locality, name number,
source of origin, etc. Stick the label on the left side of the slide.
If two or can be at the The well necessary, more pollen samples prepared same time. when blue filter will acetolyzed grains examined under the microscope through a appear
light brown with good contrast.
NOTES ON PHOTOMICROGRAPHY OF POLLEN
find the A beginner can fundamentalprinciples and usefulinformation on photomicro-
graphy in the concise Kodak & Scientific Data Book P-2: Photography Through The
consult 'Photo- Microscope, 4th ed. 1966. Furthermore, one should Samuelsson's (1965)
of and fossilized On vacation Stockholm in the micrography recent pollen'. my trip to of summer 1968, I had the pleasure meeting Mr K. E. Samuelsson, Paleobotanical De- partment, Swedish Museum of Natural History, Stockholm, and gratefully received helpful information and advice concerning the photomicrography ofpollen. He showed
me some original plates of photographs and illustrations to be used in his forthcoming will book on photomicrography, in Swedish, which be translated into English by Mrs A.
of Scotland. Some notes my experience in photomicrography of pollen grains follow.
Optical Equipment & Light Source. My equipment for photographing pollen is rather of the trinocular simple. It consists following apparatus: (1) Olympus microscope (Model
with oculars & and oil-immersion E) together 'P' 7 x 10 X, achromatic objective, attachment Model 100 X, N.A. 1.25; (2) Olympus photomicrographic (camera), PM-6 for 35 mm film; and (3) Leitz microscopic lamp, 6V—5A. This is and beautifully designed camera very easy to operate has a shock-proof shutter.
most Furthermore, in addition to the observation viewer, it has a convenient and handy
device, window for which also be used for a exposure meter, can projection. Film and Filter. black-and-white fine For photographs, an extremely grain, high
contrast and roll for has been resolving power 3 5 mm film, example, Agfa Agepan FF, chosen. In the achromatic order to obtain optimum resolution with objectives, a green
filter (Kodak Wratten filter nr 58) has always been used.
Exposure Determination.The pollen grain usually occupies only a part of the field under the would be ideal if could microscope. It one measure its image brightness directly on
the film plane. There are fight meters specially designed for photomicrography. I have been Brummelen. using a handy, self-built meter, designed by my colleague Dr J. van and suitable for the It is very sensitive making spot readings, measuring only image
brightness of the desired part of a pollen grain! Determining the optimum exposure is thus made easy and simple. than the cable release For exposures longer one second, Prontor Ultra-slow-speed
(2 to 32 seconds) has been used. 102 BLUMEA VOL. XVII, No. I, 1969.
of After a series test exposures and with some practice, satisfactorily exposed films and good results have been obtained. Detailed data for each photomicrograph taken have been recorded for future reference.
The final of Developing and Printing. quality the photographs depends largely on and in other experience in developing printing, words, on suitable darkroom techniques.
For developing the film, a proper combinationoffilmand developer should be selected.
The film has been Rodinal for — exposed Agfa Agepan FF developed in Agfa (1:20) 4 minutes 20° C with half minute. at agitation every a For printing, normal procedure and techniques have been followed.
Therequired magnification ofenlargements is conveniently obtained by photographing with combinations the stage micrometer various ofoculars and objective. During enlarg- ing, the desired negative of the micrometeris placed in the enlarger for adjustment, with ruler the reach the a linear on easel, to magnification needed.
My work on enlarging has been facilitated by using an automatic electronic darkroom
meter (Lightmaster Darkroom Computer by E. Wallner, Augsburg, W. Germany) for
measuring image brightness and contrast.
Mounting Photographs and Preparing Plates for Publication. For helpful instruction and
detailed information concerning this subject, one should consult Palmer's (1965) 'Prepa- ration of plates for paleontologie publication'. The and used have been retouched negatives glossy photographs not except sometimes
for getting rid of dirty spots in the background. Only photographs showing the desired
details and good contrast should be chosen.
Photomicrographs showing general shapes and features have usually been enlarged to
X 1000, and those showing detailed characters up to X 2000. They are arranged in
sequence to show the various levels of focus. Their orientation has been in accordance with the principles for palynograms and illustrations used by Erdtman (1952).
Since the rather been photographs of pollen grains are small, rubber cement has used them sheet of stiff white for to glue on a paper. When using this kind ofwet medium the be mounting, prepared plate is clean and flat and the photographs can easily removed for rearrangement if necessary. the information of the of Following my colleague Dr P. W. Leenhouts, plates photo-
have been the exact which tobe micrographs prepared in size in they are going reproduced. For best the results in publication, photographs have been made slightly more contrasty
than because the is reduced in desired, contrast slightly the half-tone process.
Photographing Pollen in Colour. For photographing pollen in colour, the same optical
equipment and light source as used for black-and-white have been used. In connection with the coil-filament II Professional tungsten hght source (6V, lamp), Kodachrome has been chosen. When the the transformer Film, Type A, 35 mm, voltage scale on is set on about 5.8, a Kodak hght balance filter (nr 82B) has been used.
MEASUREMENTS
The size of pollen grains varies slightly according to the methods of preparation and media used The used here mounting (cf. Praglowski, 1959; Anderson, 1965). procedure for is still in acetolysis an experimental stage and has not yet been standardized, so the measurements of pollen may not be comparable. Furthermore, the measurements usually were taken several months after preparation. In order to complete the descriptions, some the variation of measurements of the grains, showing range of together with the average
= about ten grains placed in brackets, have been included. P = length of polar axis; E Ding Hou: Pollen of Sarawakodendron (Celastraceae) and some related genera 103
equatorial diameter; t = side of polar triangle or distance between two colpi ends (cf.
& Van Campo Halle, 1959, f. 2;Punt, 1962, f. B 3).
MATERIALS
All the polliniferous samples were taken from the collections preserved in the Rijks-
herbarium, Leiden (L), Netherlands, except two. The specimen of Lophopetalum flori- bundumResearch was gratefully received from the Forest Institute, Kepong, Malaya. of Prod. One slide Lophopetalum multinervium was on loan from the Kon. Shell Explor.
Lab. (KSEPL), Rijswijk, Netherlands.
The field numbers of Malesian collections belonging to series have been cited under
serial abbreviations listed below. In order to facilitate those who are not familiar with been mentioned brackets the these series, the collector's name has usually in following
series number.
bb.: Bosbouwproefstation (Neth. Ind. For. Serv., Bogor)
BW: Boswezen Nieuw Guinea, Manokwari, New Guinea
CF: Conservator of Forests, Kepong, Malaya
FRI: Forest Research Institute, Kepong, Malaya
KEP: Forest Research Institute, Kepong, Malaya
NGF: New Guinea For. Dept., Lae
S: Sarawak For. Dept, Kuching
SAN: For. Dept, Sandakan, Sabah
SF: Singapore field numbers, Bot. Gard., Singapore
POLLEN DESCRIPTIONS, OBSERVATIONS, AND NOTES
The terminology used in the descriptions follows chiefly Erdtman (1952) and Faegri &
Iversen (1964). The concise descriptions in this paper have been supplemented with photomicrographs.
For literature, synonyms, geographical distributions, and other taxonomical matters of the and treated in I the reader revision of genera species the following, refer to my
Celastraceae (Hou, 1963 & 1964) in the Flora Malesiana. of and related The sequence genera species is one of convenience. However, genera
and/or species have been placed, if possible, close to each other.
I. SARAWAKODENDRON Ding Hou
Sarawakodendron filamentosum Ding Hou — Pl. 1: A—G. sub- Pollen grains single, isopolar, tricolporate. Shape suboblate, spheroidal, rarely
outline P = = prolate to prolate, in polar view circular; 20.0 fx (27.2 fx) 37.5 fx; E 22.5 fx
broadened at slightly bordered; (27.8 fx) 32.5 fx. Colpi fairly long, markedly equator,
t = Ora (3.9 X (6.9 as wide as 5.0 [x . slightly lalongate, 3.0 [x fx) 4.5 fx 6.0 /x fx) 7.5 fx,
wider than bordered. Nexine thick on meso- colpi, occasionally colpi, slightly c. 1.5 fx circular Sexine thick colpia, much thinner over a area surrounding ora. up to 2 fj, on
towards reticulate. Baculae <1 in mesocolpia, gradually thinning apertures, finely fx
round. lumina 11 in diam., fairly even diam., Muri simplibacul: te, <1 fx wide, up to 0.5 the in size and shape, ± isodiametric but finer in a zone surrounding apertures.
BORNEO. Hou Sarawak: Ding 333. 104 BLUMEA VOL. XVII, No. i, 1969.
Notes. By identifying flowering material with the key to the genera of Celastraceae in
theFlora Malesiana (Hou, 1963), Sarawakodendron would come next to Salacia; with the key for fruiting material it would come close to Brassiantha and Xylonymus.
the affinities of Dr Smith October Regarding Sarawakodendron, A. C. wrote me, 11,
1967: 'Your new genus Sarawakodendron appears to be very well marked, from the de-
and and have doubtthat correct. scription plate, I no your disposition is In some ways it of but of differs in seems suggestive Brassiantha, course many details'. several As to gross morphology, the present genus is allied to genera, e.g., Solatia,
Xylonymus, Hedraianthera, Brassiantha, Kokoona, Lophopetalum, etc. It is therefore ofinterest
that the pollen grains of Sarawakodendron are closely similar to those of Xylonymus and
Kokoona both in apertureconfiguration and sculpture. They differ slightly in sculpture from Hedraianthera and Brassiantha, and from Lophopetalum in being single.
II. XYLONYMUS Kalkman
Xylonymus versteeghii Kalkman — Pl. 1: H—K.
Pollen grains single, isopolar, tricolporate. Shape suboblate, outline in polar view
rounded = = triangular; P 22.5 ft (23.8 ft) 25.0 ft; E 27.5 ft (28.8 fi) 30.0 fi. Colpi fairly
short, slightly broadened at slightly bordered; t = Ora subcircular, c. equator, 8.3 ft. wide bordered. thick. 5 fx in diam., as as colpi, slightly Nexine 0.5 ft Sexine 1 fi thick, towards reticulate. Baculae in round. Muri thinning apertures, finely <0.5 ft, diam., lumina and simplibaculate, < x /< wide, up to x ft in diam., fairly even in size shape,
± isodiametric, but finer in a zone surrounding the apertures.
NEW GUINEA: BW 4686 (Chr. Versteegh).
There rather anthers of detached flower the Notes. were only two young a on type
specimen and one ofthem was still attached to the disk. I used the loose one. After boiling,
the anther has been saved. Four slides the grains were removed and empty pollen were prepared.
the of allied In characters phyllotaxy, leaves, and capsular fruits, the present genus is
to Sarawakodendron and this is shown by the close similarity between their pollen grains.
III. HEDRAIANTHERA F. v. M.
Hedraianthera porphyropetalum F. v. M. — Pl. 1: L—N.
Pollen grains single, isopolar, tricolporate. Shape subspheroidal to prolate, outline in
subcircular; = (35.7 = (32.5 Colpi polar view P 32.5 ft ft) 37.5 ft; E 27.5 ft ft) 37.5 ft. medium broadened = circular long, at equator; t 8.8 ft. Ora or slightly lalongate, c. wide thick. towards 7.5 [i in diam., as as colpi. Nexine 1 fi Sexine 2 fjt thick, thinning apertures, irregularly rugulate-reticulate. Baculae indistinct. Muri irregular in width, meandering, lumina irregular, partly connected.
AUSTRALIA. Queensland: Brass 20018.
Notes. The Hedraianthera is allied Queensland endemic monotypic genus closely to Brassiantha from distinct. New Guinea in gross morphology but is taxonomically quite
Its seed is unique so far known in the Celastraceae in having a most peculiar caterpillar- like thickening, evidently the aril, along the raphe (cf. Hou, 1964). Ding Hou: Pollen of Sarawakodendron (Celastraceae) and some related genera 105
I Fertile material of H. porphyropetalum is very scant. From a few specimens available,
fruits. not received could examine only fragments offlowers or Fortunately, long ago, we
and two flowers and fruit two very well prepared preserved specimens, bearing one the best have collected L. Brass 20018 & respectively, ones I ever seen; they were by J. (nr
20229) in 1948. the From two of the threeanthers available, I made two pollen slides and saved empty anthers. Its pollen grains resemble those of Brassiantha and of the African Salacighia (cf. of the Van Campo & Halle, 1959) in the peculiar rugulose sculpture exine. However, of the in and its apertures are general type foundalso Sarawakodendron, Xylonomus, some Kokoona species.
XV. BRASSIANTHA A. C. Smith
— Smith — Pl. Brassiantha pentamera A. C. 2: A G. Pollen grains single, isopolar, tricolporate. Shape prolate, outline in polar view sub-
= = E 20.0 (25.5 Colpi long, triangular; P 30.0[x (32.3 /x) 35.0 fx; fx fx) 27.5 fx. narrow,
an t = narrow-lalongate, dis- and slitlike, sunken in elongated depression; 8.3 [X. Ora
bordered sides. Nexine 1 thick at mesocolpium, thickened on polar tinctly on polar fx sides Sexine much reduced absent in —8 wide of ora. up to 2 fx thick, or a 3 fx elongated Muri depressed zone bordering the colpi. Baculae irregular. very irregularly ± isolated, connected. meandering, lumina very irregular, partly
NEW GUINEA: Brass 8889, 8954; NGF 9587 (K. J. White).
detailed studies the Notes. Smith and Bailey (1941) made on morphology, anatomy,
I its and taxonomie affinities of the interesting genus.Brassiantha. Later, pointed out affinity
with the Australian It is also related and genus Hedraianthera (Hou, 1964). to Xylonymus
mentioned under the latter in the Sarawakodendron as already present paper.
They examined the pollen of Brassiantha and stated that it clearly within the range of
As far its distinct in the Celas- variability of Hippocrateaceae. as known, pollen grains are
traceae. They resemble those of Australian Hedraianthera and African Salacighia (cf. Van
Campo & Halle, 1959) in the exine characters, but differ from them by the general
shape and rather unique apertures.
V. KOKOONA Thwaites
Pollen grains single, isopolar, tricolporate, rarely tetracolporate. Shape varying between oblate and prolate, outline in polar view subtriangular to circular. Colpi long, usually broadened at equator, margins distinct, sometimes slightly costate. Ora suborbicular,
as wide as wider than annulate. Nexine —1 thick. usually or colpi, sometimes 0.5 pi reticulate towards coarsely to finely or even Sexine 1—1.5 pi thick, thinning apertures,
and to 1 round, either tectate-psilate. Baculae varying in size distinctness, lip pi in diam., reticulate and arranged in a pattern or irregularly grouped supporting an apparently Muri continuous, smooth tectum (K. ovatolanceolata and K. reflexa). simplibaculate, lumina rather finer towards
Notes. The sexine of Kokoona pollen varies from finely or coarsely reticulate to nearly be or completely psilate-tectate and it is of interest that this phenomenon appears to
correlated with anther characters.
of the Malesian In the key to the species Kokoona, based on anther characters, species have been grouped under two headings: 106 BLUMEA VOL. XVII, No. I, 1969.
i) Anthers with distinctly prolonged connective — K. ochracea, K. littoralis, K. coriacea,
K. sessilis.
Anthers without with obscure or short ovatolan- 2) or very prolonged connective — K.
ceolata, K. reflexa.
The extra-Malesian K. be added the first species zeylanica can to group and K. filiformis the far to second. So there are eight species recorded for this genus.
Those to the first reticulate species belonging group possess grains (lumen size usually while those the second have < 1 /x), in probably completely psilate-tectate ones (lumen
size> 1 fx). The difference is gradual, however.
The of characteristic and apertures some species appear highly those of K. filiformis and Sarawakodendron are similar to each other (cf. Pi. 4 G—H and I E).
of is in that of The pollen Kokoona also general appearance comparable to Sarawako- dendron Of and Xylonymus. interest, however, are the sharp differences which exist between of the former and Kokoona and Lophopetalum: the pollen grains are single those of the latter in tetrads occur or polyads; they also differ in aperture construction.
The character for the pollen provides anadditional separation of these two genera, which of them has met difficulties in the past. Now I may say that my delimitation based on
is gross morphology (cf. Hou, 1963) supported by wood anatomy (Balan Menon, 1964) and palynology.
1. Kokoona ochracea (Elmer) Merr. — Pl. 2: H—J.
Pollen lumina to diam. = grains suboblate, coarsely reticulate, up 2 /r in P 25.0 fi
(26.5 = t = fi) 27.5 fi. E 30.0fi (30.5 fi) 32.5 fi. 7.5 fi (8.8 fi) 10.0 fi.
PHILIPPINES. Palawan: A. D. E. Elmer 12881.
INDONESIAN BORNEO: Kostermans 5143.
Note. Erdtman's (1952) record of the pollen of this species is the first published infor- of based mation on the palynology the genus. I saw his original sketch and notes of it,
on Elmer 21328 from Borneo.
littoralis — Pl. 2. Kokoona Laws. 3: A—F. Pollen grains suboblate, tetracolporate grains occasionally observed, coarsely reticu-
diam. = = lumina 2 in P (29.8 E t — late, fi 27.5 ft fi) 35.0 /i. 35.0 fi (40.0 fi) 42.5 fi.
10 (10.5 [i /x) 12.5 [X.
MALAY PENINSULA: SF H. 30177 (E. J. Corner). Sarawak: Hou & BORNEO. Ding 534 550; S 9537 (E. F. Brunig).
Note. As stated in my revision of the Celastraceae, the leaves of K. littoralis are variable
consistent in shape, texture, and size, but the floral characters are rather homogeneous and and well (Hou, 1963). Its pollen grains as far as examined, are rather uniform, normal, developed.
— Pl. K 3. Kokoona coriacea King 2: —M.
Pollen to lumina to grains prolate spheroidal subprolate, coarsely reticulate, up 2 fi
in diam. P = E = t = c. 32.5 fi (34.5 fi) 37.5 fi. 30.0 fi. 15 fi.
MALAY PENINSULA: Dr King's collector 4226.
is Note. This species known only from the type collection. The pollen preparation was
made from one anther. Ding Hou: Pollen of Sarawakodendron (Celastraceae) and some related genera 107
4. Kokoona sessilis Ding Hou — Pl. 3: G.
Pollen oblate lumina diam. = u grains spheroidal, finely reticulate, up to I /t in E 30.0
t = (6.0 (32.5 fx) 33.5 fx. 5.0 fx fx) 7.5 fx.
MALAY PENINSULA: SF 36296; FRI3373 (F.S.P. Ng).
Note. in flower cited above and in In addition to the type, two specimens, one as one Ulu fruit(FRI 5372), were collected by F.S.P.Ng, from Kelantan, in 1967. Unfortunately, both flowering specimens have rather young buds. The grains appear not well developed flat. Most of in view and few of themhave been and are very the grains are polar only a observed in equatorial view.
Kokoona Thwaites Hook. t. 5. zeylanica in Kew J. Bot. 5 (1853) 379, 6. — Pl. 4: A.
Pollen lumina in diam. Four grains: P = grains spheroidal, finely reticulate, 0.5 fx
(35.6 E = (34.4 t = 32.5 fx fx) 37.5 fx. 32.5 [X fx) 35.0 fx. 7.5 fx.
CEYLON: Thwaites C.P. 2384.
There flower-buds the available. The Note. are only very young on isotype specimen
colpi of the grain appear very narrow and the ora are transversely elongated.
6. ovatolanceolata Ridl. — Pl. Kokoona 4: B—C.
Pollen to P = grains spheroidal prolate spheroidal, nearly psilate-tectate. 30 fx (35.6 u)
= = (8.1 ) 10.0 40.0 /x.E 30.0 /x (35.0 fx) 37.5 /x.t 7.5 fx [x 11.
BORNBO. Indonesian: bb. — Sarawak: A. R. Anderson 32393. J. 7910.
Note. As far the sexine of the is the is as pollen grains concerned, present species probably tectate-psilate.
Kokoona reflexa — Pl. 7. (Laws.) Ding Hou 3: H—M.
Pollen = grains usually suboblate, rarely subprolate, nearly psilate-tectate. P 32.5 /.<
(35-0 E = t = 10.0 (11.3 fx) 37-5 ft- 27.5 fx (33.8 fx) 37.5 fx. fx fx) 12.5 /x.
1P. SUMATRA: Achmad 943; bb. 34E. 634 & 633 (Thorenaar). INDONESIAN BORNEO: Kostermans 6386.
Note. In the slides prepared from the above cited collections, there are usually more
suboblate than subprolate grains.
8. Kokoona filiformis (Laws.) C. E. C. Fischer, Kew Bull. (1927) 311. (basionym:
—I. Lophopetalum filiforme Laws.) — Pl. 4: D Pollen grains usually tetracolporate, rarely tricolporate. Tetracolporate grains: 4-angular
in polar view, suboblate, probably nearly psilate-tectate; P = 27.5 fx (30.5 /x) 32.5 fx;
= wider than the bordered. E 37.$ fx (40.8 fx) 42.5 fx; ora colpi, Tricolporate grains:
(39.4 in diam. in view; two in equatorial view observed; 37.5 fx u) 40.0 fx polar only grains
P = 27.5—30.0 fx; E = 37.5—40.0 fx; t = c. 12.5 /x.
THAILAND: Kerr 12730.
There of Note. was only one flowering specimen this species available. In the pollen than the slides prepared from one flower, the tetracolporate forms are more common
tricolporate ones. 108 BLUMEA VOL. XVII, No. i, 1969.
VI. LOPHOPETALUM Wight ex Arnott
tricol- Pollen grains in tetrads or polyads (8 or 16 grains, in one species), triporate or of of porate in one species, position apertures variable. Shape single grains spherical. and ill defined. Colpi, whenpresent, long fairly narrow. Ora circular, sometimes Nexine
thick. to sometimes covered 0.5—0.7 fx Sexine 1.4—2.1 fx thick, finely coarsely reticulate, with in Baculae in size and large verrucae two species. varying distinctness, up to 0.5 fx diam. lumina rather isodiametric. in Muri simplibaculate, < 1 fx wide, uniform, ±
Notes. The genus Lophopetalum, which has been redelimited by myself (Hou, 1963),
of rather far consists seventeen species; it is homogeneous as as the gross morphology
into is corcerned. It can hardly be subdivided natural groups by any character or com- be bination of characters without overlapping. However, the pollen grains can grouped into at least four distinct types.
Diagrammatic sketches of pollen tetrads of two Lophopetalum species were published
his Fl. Cochinch. vol. sub t. & as the sketches by Pierre in For. 4, 307 A B, early as 1894; of in t. 307B were reproduced by Loesener (1942) under the name L. wightianum.
Group A.
Pollen grains in tetrahedral tetrads, tricolporate, outline of single grains in polar view wide circular. Colpi meeting two and two, long, rather narrow. Ora subcircular, as as in diam. Sexine baculae muri thin, lumina colpi, 4.2 fx very finely reticulate, indistinct, uniform and diam. round, in size shape, 0.5 fx in
1. Lophopetalum wallichii Kurz (syn.: L. celastroides Laws.) — Pl. 5: A—D.
Pollen tetrads diam. Individual 60.0 fx (63.8 fx) 67.5 fx in grains 30.0 fx (38.3 fx) 42.5 fx in diam.
Larsen & THAILAND: Kerr 1650; Sarensen, Hansen 1007.
Notes. In this extra-Malesian resembles L. gross morphology, species closely pallidum from from the Malay Peninsula. However, their pollen grains are distinct each other, here. belonging to different groups
In the each of the tetrad This the present species, grain is tricolporate. is only species in this genus to have such pollen.
those the and Cam- The tetrads are similar to of African Bequaertia, Tristemonanthus, pylostemon (except C. laurentii de Wild., which has tri-, rarely tetraporategrains, cf. Van
Campo & Halle, 1959). The tetrad is of the tetrahedral such when viewed from type. Sometimes, a tetrad, a different angle, resembles a '+' (cf. pi. 5 C —D; Faegri & Iversen, 1964, pi. VIII, fig. 23).
the The single grains are similar in general appearance to those ofKokoona, but aperture construction appears shghtly less complicated.
Group B.
Pollen grains in tetrahedral, rarely rhomboidal, tetrads. Colpi absent. Ora circular,
each but shifted in L. from generally not opposite other, (e.g. subobovatum, ±45° away the of the cf. 6: & diam. tricolporate position preceding group, pi. I J), 2.9—5.7 fx in Sexine baculae indistinct muri very finely to coarsely reticulate, or distinct, 0.5 fx wide, uniform and diam. |umina in size shape, 0.5—1 fx in Ding Hou: Pollen Sarawakodendron and related of (Celastraceae) some genera 109
The size and the cxine of the Note. This is a large group, including thirteen species. subdivide this grains in sonic species are rather characteristic. It may be possible to group
into smaller units if detailed studies, based on extensive collections, will be made by a
palynologist. The tetrads in this group are similar to those of the African Campylostemon laurentii
(cf. Van Campo & Halle, 1959).
2. Lophopetalum beccarianum Pierre
Pollen tetrads (41.3 in diam. Individual (26.8 37.5 fx /x) 42.5 fx grains 25.0 fx fx) 30.0 fx in diam.
Sabah: Puasa Patrick Pin Sam BORNEO. 4542; A-1896.
3. Lophopetalum glabrum Ding Hou Pollen tetrads in diam. Individual (24.6 35.0 fx (38.3 fx) 40.0 fx grains 22.5 fi /x) 27.5 fx in diam.
BORNEO. Brunei: CF 34564. — Sarawak: S 23809 (Jugah anak Kudi).
4. Lophopetalum rigidum Ridl.
Pollen tetrads in diam. Individual ix (29.5 47.5 fi (50.0 fx) 52.5 ft grains 27.5 /x) 30.0 fx in diam.
BORNEO. Sarawak: Anderson 3361 /6.
floribundum 5. Lophopetalum Wight Pollen tetrads (42.8 diam. Individual (27.6 a) 40.0 fx fx) 45.0 /tin grains 25.0 fx 30.0 fx in diam.
MALAY PENINSULA: KEP 0659.
6. Lophopetalum macranthum (Loes.) Ding Hou Pollen tetrads diam. Individual 57.5 fx (60.8 fx) 62.3 fx in grains 37.5 fx (39.5 fx) 40.0 fx
in diam.
NEW GUINEA: Gjellerup 701.
7. Lophopetalum micranthum Loes. Pollen tetrads diam. Individual 47.5 fx (52.5 fx) 55.0 fx in grains 32.5 fx (34.7 fx) 35.0 fx
in diam.
NEW GUINEA: Versteeg 1773.
8. Lophopetalum pallidum Loes. — Pl. 6: F—H. Pollen tetrads diam. Individual 52.5 fx (55.6 fx) 57.5 fx in grains 30.0 fx (34.2 fx) 37.5 fx in diam.
MALAY PENINSULA: Phytochem. Surv. Malaya, Kuala Lumpur, 1566.
9. Lophopetalum javanicum (Zoll.) Turcz. — Pl. 6: A—C. Pollen tetrads diam. Individual diam. 50.0 μ (51.6 μ) 55.0 μ in grains 32.5 μ in
Thailand: Kerr 11970. BORNEO. Sabah: SAN 21376 (J. Singh).
10. Lophopetalum multinervium Ridl. — Pl. 6: D—E.
Pollen tetrads diam. Individual diam. 50.0 fx (54.4 fx) 57.0 fx in grains 35.0 fx in
BORNEO. Kostermans & — Indonesian: 7720 10375. Sarawak: S 1264 (Tready). 110 BLUMEA VOL. XVII, No. I, 1969.
11. Lophopetalum ledermannii (Loes.) Ding Hou Pollen tetrads in diam. Individual 55.0 fx (59.7 fx) 62.5 fx grains 35.0 fx (37.3 fx) 37.5 ft in diam.
NEW GUINEA: Docters v. Leeuwen 9622.
torricellense — Pl. 12. Lophopetalum Loes. 5: E—F.
Pollen tetrads diam. Individual 62.5 fx (68.3 fx) 75.0 fx in grains 40.0 fx (43.8 fx) 47.5 fi in diam.
NEW GUINEA: Brass Brass & 12326; Versteegh 11903.
13. Lophopetalum subobovatum King — Pl. 6: I—K. Pollen tetrads in diam. Individual 50.0 fx (54.8 [x )57.5 fx grains 30.0 fx (33.9 fx) 40.0 fx in diam.
MALAY PENINSULA: KEP 71323. BORNEO. Sarawak: Beccari P.B. 2639.
14. Lophopetalum duperreanum Pierre, Fl. For. Cochinch. 4 (1894) t. 307A.
Pollen tetrads in diam. Individual 55.0 [X (61.3 fx) 65.0 fx grains 37.5 fx (38.4 fx) 42.5 fx in diam.
COCHINCHINA: Pierre 4082.
Group C.
Pollen grains in tetrahedral, rarely rhomboidal, tetrads. Colpi absent. Ora circular,
three at of position variable, very rarely meeting together junctures grains, 4.2—6.4 fx diam. in Sexine very finely reticulate, bearing conspicuous subglobose verrucae, regularly surface of and rooted baculate spaced, c. 10 fx apart on sexine in layer.
15. Lophopetalum wightianum Arn. — Pl. 7: B—G. Pollen tetrads tetrahedral, (72.3 in diam. Individual 65.5 fx /x) 82.5 fx grains 42.5 fx diam. diam. (45.8 fi) 50.0 fx in Verrucae 2.5 fi (4.3 fx) 7.5 fx in
INDIA. Canara: R. F. Terr. Hohenacker 324. Pierre COCHINCHINA: 421.
SUMATRA: bb. 49E.1P.339 (Endert).
INDONESIAN BORNEO: Kostermans H. Winkler 4972; 2623.
16. Lophopetalum pachyphyllum King — Pl. 7: A. Pollen tetrads in diam. Individual (37.2 55.0 fx (63.7 fx) 67.5 fx grains 32.5 fx fx) 45.0 fx diam. diam. in Verrucae c. 2.5 fx in
MALAY PENINSULA: Dr King's collector 7323.
Notes. In kinds of in a the present two species there are two pollen arrangements tetrad: tetrahedral and rhomboidal and Each (cf. Wodehouse, 1935, Erdtman, 1945). has three sometimes individual grain distinct apertures. In a tetrahedral tetrad, the three
each from situated the of the apertures, a respective grain, are at junction grains (pi. 7, D), comparable with the situation in Helia brevifolia (Gentianac.) (Erdtman, 1952).
The most conspicuous pollen character of these two species is presented by the
subobovoid or subglobose verrucae of the exine. They resemble the enlargement ofa
Their and structure shouldbe studied ultra-thin sections. pilum (pi. 7, G). origin by using Ding Hou: Pollen of Sarawakodendron (Celastraceae) and some related genera 111
The tetrads of these two species are quite similar to those of the polyads in the African
Hippocratea vignei Hoyle (Van Campo & Halle, 1959). and related and their Lophopetalum wightianum L. pachyphyllum are closely species similar other. pollen grains are to each
Group D.
Pollen grains in polyads, consisting of 8, rarely 16, grains joined together, the
arrangement not regular, sometimes simple tetrahedral tetrads also present. Colpi absent. Ora rather and very indistinct, 2—3 per grain, position irregular. Sexine regularly very finely reticulate, muri in width, lumina in diam., isodiametric. 0.5 fx 0.5 fx
17. Lophopetalum sessilifolium Ridl. — Pl. 8: A—F. 2-tetrad Polyads of 57.5 fx (65.1 fx) 75.0 fx X 42.5 fx (48.6 fx) 55.0 fx. Polyads of 4-tetrad
80.0 x Individual 70.0 fx (72.0 fx) fx 55.0 (60.8 fx) 70.0 fx. grains 27.5 fx (30.0 fx) 32.5 fx in diam. in polar view.
BORNEO. Sarawak: J. A. R. Anderson 63;i; Native collector 414 & 543.
Notes. In this be and gross morphology, species can easily recognized distinguished
from the other species of Lophopetalum, why it has been placed at the beginning of the is L. key (Hou, 1963, p. 264). It closely allied to L. beccarianum, floribundum, L. glabrum, and the these four L. rigidum. However, pollen grains of species belong to group B.
It is surprising to find polyads consisting of two, rarely four, tetrads in Lophopetalum. tetrads Polyads consisting of4 were already known from two African and one American
species of Hippocratea (sensu Hallé, 1962) belonging to the former Hippocrateaceae (Van
Campo & Halle, 1959, andWaanders et al., 1968). However, contrary to the two African
of where the of in the species Hippocratea polyads are composed 4 tetrads, present species
the 16 grains are not regularly arranged.
8 of Polyads of 2 tetrads or grains have not been recorded for any other species Celastraceae (sens. lat.). At first I thought that they originated by breakdown from 4-tetrad
in anthers well in made from or 16-grain polyads. However, open as as preparations
a young anther, 2-, rarely 4-, tetrad polyads are always, and some simple tetrads are
16 anther sometimes present. The polyads of grains do not occur in every or flower. The individual obscure grains of the polyads possess only pores. The pores are usually and be sometimes two or three of them can observed on a pollen grain. I am neither certain of the of the of the number and arrangement grains in a polyad nor exact of the The ofthe further detailed disposition pores. pollen present species requires study.
ACKNOWLEDGEMENTS
I am gratefultoProf. Dr C. G. G. J. van Steenis, Director ofthe Rijksherbarium, Leiden,for his kindness
to read the manuscript. I am indebted to those ofmy colleagues and friends who have helped me in the
course of this study, particularly to Dr J. van Brummelen for discussions on techniques, to Mr J. Muller for his help in matters of pollen morphology, also by enlarging the pollen descriptions, and for his interest in this and Leenhouts for information editorial paper, to Dr P. W. on matters.
I wish to take this opportunity to thank Prof. G. Erdtman and his staff, PalynologicalLaboratory, Solna, for the and extended there. Sweden, courtesy, facilities, assistance to me during my visit 112 BLUMEA VOL. XVII, No. I, 1969.
REFERENCES
ANDERSON, S. T., 1965. Mounting media and mounting techniques, in: B. Kummel & D. Raup, Hand- book of paleontological techniques: 587—598. BALAN MENON, P. K., 1964. Perupuk wood, revision of nomenclature and classification. Mai. For. 27:
18—21, 2 figs.
CAMPO, M. VAN, & N. HALLE, 1959. Les grains de pollen des Hippocrateacees d'Afrique de L'Ouest. Bull.
Inst. Fr. Afr. Noire 21, ser. A: 807—899, pi. 49—88, fig. 1—4. Reprinted in Pollen et Spores vol. I,
no. 2, in 1959. introduction ERDTMAN, G., 1943. An to pollen analysis. 239 pp., 28 pis. & 15 figs. On Svensk 1945- the occurrence of tetrads and dyads. Bot. Tidskr. 39: 286—297, 2 figs. and Off-set edition 1952. Pollen morphology plant taxonomy, Angiosperms. 539 pp., 261 figs. (with in addenda pp. 543 —553, fig. 262) 1966.
i960. The acetolysis method. A revised description. Svensk Bot. Tidskr. 54: 561—564.
Textbook of 2nd ed. FAEGRI, K., & J. IVERSEN, 1964. pollen analysis, rev. 237 pp., 8 pis. & 23 figs. HALLE, N., 1962. Monographic des Hippocratiiacees d'Afrique occidentale. M6m. Inst. Fr. Afr. Noire 64:
1—245, 76 figs-
Hou, D., 1963. Celastraceae I, in: Flora Malesiana I, 6: 227—291, fig. 1 —22.
Celastraceae ditto: 1964. II. 389—421, fig. 23—37.
1967. Sarawakodendron, a new genus of Celastraceae. Blumea 15: 139—143, fig. 1.
LOESENER, TH., 1942. Celastraceae, in: E. & P. Nat. Pfl. Fam. ed. 2, 20b: 87—197, fig. 21—61.
PALMER, A. R., 1965. Preparation of plates for paleontologic publication, in: B. Kummel & D. Raup,
Handbook of paleontological techniques: 456—459.
Bot. PRAGLOWSKI, J. R., 1959. On the swelling of pollen grains in glycerine jelly. Not. 112: 175. of the with references Wentia PUNT, W., 1962. Pollen morphology Euphorbiaceae special to taxonomy. 7:
I—116, pi. 1 —23, fig. I —2.
ROBSON, N., 1965. Taxonomic and nomenclatural notes on Celastraceae. Bolet. Soc. Broteriana 39: 5 —55.
D. SAMUELSSON, K. E., 1965. Photomicrography ofrecent and fossilized pollen, in: B. Kummel & Raup, Handbook of paleontological techniques: 623 —636, 4 figs. New SMITH, A. C., & I. W. BAILEY, 1941. Brassiantha, a new genus of Hippocrateaceae from Guinea.
J. Arn. Arb. 22: 389—394, 1 pi.
TRAVERSE, A., 1965. Preparation of modern pollen and spores for palynological reference collection, in: B. Kummel & D. Raup, Handbook of paleontological techniques: 598—613, 5 figs.
WAANDERS, L., J. J. SKVARLA, & C. C. PYLB, 1968. Fine structure of Hippocrateaceaepollen walls. Pollen
et Spores 10: 189—196, s pis.
R. & WODEHOUSB, P., 1935. Pollen grains. 574 pp., 14 pis. 123 figs. Reprinted in 1959- and related Ding Hou: Pollen of Sarawakodindren (Celaslraceae) some genera 113\1
Plate 1. Sarawakodendron filamentosum Ding Hou. A—B. Two successive foci of sexine; C. polar view; view of F. section in view of short axis D. optical cross section; E. equatorial aperture; optical equatorial a
— Kalkman. grain; G. optical section in equatorial view of a long axis grain. Xylonymus versteeghii H—J.
F. M. L. Polar view, three successive foci; K. equatorial view. — Hedraianthera porphyropetalum v. Polar
M. N. view. B & D & view; optical cross section; equatorial (A, E, x 2000; C, F—N, x 1000. A—G,
Hou Brass Ding 333! H-K, BW 4686; L—N, 20018).
113 BLUMEA VOL. 114\2 XVII, No. i, 1969
Plate Brassiantha C. Smith. A—B. successive foci of muri curved and 2. pentamera A. Two sexine, winding; C. D. sunken E—G. polar view; optical cross section, showing especially the colpi; equatorial view, three and successive foci, F. showing the thickened marginsofthe ora, and G. showing the slit-like colpus narrow-
Merr. H. I. cross lalongate os. — Kokoona ochracea (Elm.) Polar view; optical section; J. equatorial view, showing the distinct annulus. — K. coriacea King. K. Polar view; L. optical cross section; M. equatorial
view. Brass Kostermans K Dr (A & B, X 2000; C—M, x 1000. A—G, 8954, H—J, 5143; —M, King’s coll. 4226). Ding Hou: Pollen of Sarawakodendron (Celaslraceae) and some related genera 115\3
Laws. A—B. Two foci of muri in A Plate 3. Kokoona littoralis successive reticulum, showing (bright) E—F. and bacules (black dots) in B; C. polar view; D. optical cross section; equatorial view, two successive Polar foci, E. showing the distinct annulus, and F. optical section. — K. sessilis Ding Hou. G. view of
H. I. — K. Hou. Polar a rather young grain. reflexa (Laws.) Ding view; optical section; J. equatorial view; K—M. three successive foci of axis & C — Hou a long grain. (A B, x 2000; M, x 1000. A—F, Ding 550;
G, SF 36296; H —M, Achmad 945). 116\4 BLUMEA VOL. XVII, No. I, 1969
A. view — K. Plate 4. Kokoona zeylanica Thwaites. Polar of a rather young grain. ovatolanceolata Ridl.
— E. B. Optical cross section; C. equatorial view, showing a distinct annulus. K. filiformis (Laws.) C. C.
Fischer. D. Optical cross section of a tetracolporate grain; E. polar view of a tetracolporate grain; F. optical
of G—I. view of three successive foci of the cross section a tricolporate grain; equatorial aperture, same G. distinct H. grain: showing a annulus at high focus; slightly lower focus than G, showing the os much
end of wider than the width of colpus at the equatorial part; I. one an optical section, showing the two exine. B distinct layers of (A—F, x 1000; G—I, x 2000. A, Thwaites C.P. 2584; & C, Anderson 7910; D—I, Kerr 12750). Ding Hou: Pollen of Sarawakodendron (Celastraceae) and some related genera 117\5
wallichii tetrahedral tetrad with on Plate 5. Lophopetalum Kurz. A. One one tricolporate grain top in B. —D. in viewed optical cross section; optical cross section; C two tetrahedral tetrads as A, at different
out in cross-like with the lower of focus each tetrad. — angle, appearing in a arrangement two grains L. Loes. E—F. Two in torricellense successive foci of a tetrahedral tetrad polar view. (All x 700. A—D,
Sorensen et al. 1007; E—F, Brass 12326). VOL. No. 118\6 BLUMEA XVII, i, 1969
A—C. Three foci ofa tetrahedral tetrad, Plate 6.Lophopetalum javanicum (Zoll.) Turcz. successive showing D. E. rhomboidal — multinervium Ridl. tetrahedral each ofthe three grains with three apertures. L. tetrad;
a the sexine in various tetrad. —L.pallidumLoes. F—H. Three successive foci of tetrahedral tetrad, showing
Three successive foci of a tetrahedral tetrad. X foci in F. — L. subobovatum King. I—K. (All 700. A—C,
F I— KEP — Malay 71523). SAN 21376; D—E, Kostermans 7720; H, Phytochem. Survey 1566; K, Pollen Sarawakodendroti and some related Ding Hou: of (Celastraceae) genera 119\7
A. Plate 7. Lophopetalumpachyphyllum King. Polar view of a tetrahedral tetrad, showing one aperture
— L. Arn. ofa tetrahedral tetrad, showing especially on each respective grain. wightianum B. Optical section
view of rhomboidal D. view of tetrahedral the verrucae on the sexine; C. polar a tetrad; polar a tetrad, foci three each from situated at the E—F. two successive showing apertures, a respective grain, junction; in G. an section ofthe grain wall, showing three verrucae relation of the sexine; a small portion of optical
Dr B— Kostermans to the exine. (A—D, X 700; E—G, X 2000. A, King’s coll. 7525; G, 4972). VOL. No. 120\8 BLUMEA XVII, I, 1969
Plate 8. Lophopetalum sessilifolium Ridl. A—B. Two successive foci of a two-tetrad polyad, showing the
rhomboidal tetrad in A and the lower in C. two-tetrad in another kind of upper one B; a polyad arrange- with three three foci of ment (2 + 4 + 2), some grains showing two or apertures; D—F. successive a four-tetrad the four rhomboidal in the middle polyad, showing upper grains (a tetrad?) D, eight grains in E and the lower four grains in F (a rhomboidal tetrad?). (All x 700. Native collector 414).