BLUMEA 21 (1973) 153—178
Comparative leaf anatomy of
Kokoona and Lophopetalum
W.T. Jansen and P. Baas
Rijksherbarium, Leiden
Contents
Summary 153
Introduction 153
Materials and methods 154
Results 156
Generic and specific descriptions 156
in Development of the stomatal complex Kokoona ovatolanceolata! 173
Discussions 173
Stomata 173
Vascular of and midrib system petiole 175
Variability of anatomical characters in certain species 175
Diagnostic and taxonomic value ofsome leaf anatomical characters 177
General conclusions 177
Acknowledgements 178
References 178
Summary
is The leaf anatomy of Kokoona and Lophopetalum described in detail. Separation of the two genera as effectuated differences vascular of the distal end of the by Hou (1963) is supported by in anatomy petiole, is in Other which invariably more complex Lophopetalum than in Kokoona. differential characters (p. ooo)
ofthe of As based on leaf itis define only apply to part species Lophopetalum. anatomy impossible to groups the of closely related species within the genera because of a lack of mutual correlation between different characters. in shows leaf anatomical Lophopetalum particular some striking anatomical features such as a very
variable and complex arrangement of vascular tissue in petiole and midrib, complex types of cyclocytic
and and Most of these characters in of anisocytic stomata, crystarque cells, epidermal papillae. occur part the The genus only. variabilitybelow the species level ofcharacters such as epidermal wall thickness, number of cells dimension of cell absence of in subsidiary per stoma, guard pairs, presence or stomata upper epider- Because mis etc. is reported for some species. of the great range of anatomical variation it is impossible to identify species, using leaf anatomical characters only.
Introduction
In his revision of the family Celastraceae for the Flora Malesiana Hou considered the
Kokoona and Arn. the genera Thwaites Lophopetalum Wight ex as distinct, on grounds of differences in floral and seed characters (1963: 264). Herewith he refutedKurz's suggestion
(1870: 73) to reduce Kokoona to Lophopetalum. A natural classification of the wood of
This carried the first author under *) paper reports on the results ofa post graduate study out by the super-
vision of the second author.
153 BLUMEA VOL. No. 154 XXI, I, 1973
Kokoona and Lophopetalum coincides with Hou's botanical classification (Balan Menon
1964: 20). A study of pollen morphology by Hou (1969) resulted in the discovery of for both Kokoona another character supporting the generic status and Lophopetalum.
This of leaf undertakenin order to have a more study anatomy was complete survey of of whether the characters these genera and to find out there is leaf anatomical support of for their separation. Furthermore, the results such a study might be helpful in the identification is for these of sterile material which very difficult genera (Hou 1963: 260
and A further contribution the ofthe leaf in 264). to knowledge anatomy Celastraceae was
regarded as another justification of this detailed study.
and Some data on the leaf anatomy of some species of Kokoona Lophopetalum can be found Stenzel in publications by (1893: 50, 70), Metz (1903: 27—28), Loesener (1892: Solereder and 192—193), (1908: 87—96), Metcalfe and Chalk (1950: 389—391). In the
present paper descriptions and tables of characters ofall species of Kokoona andLophopeta- known and will be lum, at present (Hou, 1962, Byrnes, 1971), given.
In order to gain some impression of leaf anatomical variation within a species (be it
either ecologically or genetically controlled) 11 specimens of Kokoona littoralis and selected for These known show Lophopetalum javanicum were study. species are to much Of studied. Ofall leaf morphological variation (Hou, 1963). K. sessilis 6 specimens were
remaining species at least i, mostly 2 specimens were studied.
MATERIALS AND METHODS
All material taken from dried in the was specimens present Rijksherbarium, Leiden,
for a of Kokoona which was except specimen coriacea, kindly provided by Mr. J. P. M.
Brenan, Keeper ofthe Herbarium, Royal Botanic Gardens, Kew. Because of the possibility of misidentification of material the leaves taken from non-flowering were always speci- with flowers. taken from mens They were a branchlet just below the basis of a flowering shoot in order to be certain that fully developed leaves were studied.
mounted Permanent preparations, in Euparal, were made of transverse microtome sections of the central part of the laminaand both distal and basal end of the petiole and of free hand sections of the leaf surface. Transverse sections were stained with a safranin- haematoxylin mixture. Hand sections were stained with safranin only. This procedure of was followed for one specimen each species. Semi-permanent shdes, mounted in made glycerine jelly, were ofall other specimens of which transverse sections were made. made Cuticular preparations of all specimens were using equal volumes of hydrogen- peroxyde 30% and glacial acetic acid. The cuticles were stained with Sudan IV and mounted in glycerine jelly. the width of the cell Measurements were carried out as follows: guard pairs at their from measured with widest part and the length pole to pole were a linear eye piece micrometer in all cuticular preparations. Lamina and cuticle thickness were measured in
distinction between glycerine jelly preparations. In most specimens no sharp was present uncutinised cell wall, cuticular layer, and cuticle (cf. Stace, 1965). Thereforethe total outer
wall thickness of the in the anticlinal periclinal epidermal cells halfway between walls was the cuticle. indices calculated measured instead of the thickness of Stomatal were according to the formula:
number of stomata
S.I. — x 100 number of epidermal cells + stomata + subsidiary cells
This formula deviates from the one given by Salisbury (1927) in the presence of the number of subsidiary cells in the fraction's denominator. I have not found whether the W. T. JANSEN & P. BAAS: Leaf anatomy of Kokoona and Lophopetalum 155
of of vascular tissue in midrib and in Kokoona and Fig. 1. Types arrangement petiole Lophopetalum. and 6—8 Types 1 and 3 are simple, the other types are complex. Type 1 and 3—5 are ‘open’, types 2 are ‘closed’.
cells should be counted or taken with the stomata in subsidiary apart together any publi- cation in which the stomatal index is used. It seems logical to include the number of
cells total subsidiary in the sum of cell types used in the fraction's denominator.Probably
Salisbury worked with plants with anomocytic stomata which would account for the omission of subsidiary cells.
Average values of stomatal sizes are based on 25 measurements. Stomatal indices are based in the of the lamina between midrib and leaf on 10 counts part margin at a 156 BLUMEA VOL. XXI, No. I, 1973
of least from either of them. These carried distance at 3 mm counts were out in cuticu- of lar preparations using a grid eye piece micrometer at a magnification X 1000.
For the description of stomatal types the terminology of Van Cotthem (1970) was The and for followed. terms complex anisocytic complex cyclocytic were used in addition
of where subdivisions of sub- types occurring in some species Lophopetalum one or more cells evident. As in between aniso- sidiary are shown fig. 24, transitional types cyclocytic,
cytic, and the complex types occur (arrows). In many specimens two or more of the types and in the leaf. of fig. 24 intermediates occur same of the of different vascular the of Because occurrence complex systems in petioles
Lophopetalum, the types were classified as in fig. I. The classification is artificial and was made for convenience only.
Collection numbers and abbreviations, used according to Hou (1969: 103) are recorded
before each specific description. Numbers marked with* were used for both transverse cuticular sections and cuticular preparations. All unmarked numbers were used for
preparations only.
Abbreviations used in the text:
1 = long S.V. = in surface view
w = wide T.S. = in transverse section
RESULTS
The leaf anatomical data have been recorded using both descriptions and tables. The
the generic descriptions are as comprehensive as possible, but in specific descriptions only
those characters not suitable for listing in a table are mentioned. The specific descriptions and with the are therefore unbalanced and should be used together with table I or II
generic description. The complex vascular systems in Lophopetalum are not elaborately
described, but only reference to figures is made.
GENERIC AND SPECIFIC DESCRIPTIONS
Kokoona & Thwaites (Table I; Plate I, 2 3, III, 2; Fig. 24)
S.V.: Cuticle smooth to granular. Unspecialized epidermal cells polygonal, with straight
curved anticlinal walls which from thin thick. Thin areas of to slightly range to very adaxial cuticle or pit-like structures absent. Minor veins mostly not prominent in epi-
dermis, not to fairly prominent in abaxial epidermis. Stomataconfined to abaxialepidermis,
for one of Kokoona or except specimen ovatolanceolata (see specific description), cyclocytic with anisocytic, 2—9 subsidiary cells, 13—38 /im 1., 10—34 fim w., length-width ratios
1. Outlines of cells difficultto observe in cuticular 0.9—1.5, mostly ca. subsidiary prep- because of the cuticle the with thick arations being very thin here except in specimens
anticlinal epidermal cell walls where only those adjacent to the guard cells have in-
conspicuous cuticular flanges. Outer and inner stomatal ledges not conspicuous. Polar Granular T-pieces absent. material sometimes present in the stomatal pore. Subsidiary
index 6.2 cells partly or almost completely underlying the guard cells. Stomatal —12.4.
Cork warts present in varying amounts.
dorsiventral or to isobilateral, thick. Unspecialized T.S.: Lamina tending 140—470 fxm epidermal cells rectangular, tall to flattened, those of abaxial epidermis usually lower than
those of adaxial epidermis. Anticlinalwalls often tapering towards the mesophyll, in some the middle walls bulging in (Plate I, i). Outer periclinal —12 thick. Hypoder- species 4 /im of adaxial abaxial of mis consisting I, rarely 2 or layer(s) square to flattened, sometimes cells pentagonal, translucent which are mostly somewhat bigger than adjacent epidermal W. T. P. BAAS: Kokoona and JANSEN & Leaf anatomy of Lophopetalum 157
adaxial of cells cells. Mesophyll composed of I—4 layer(s) palisade parenchyma (up to 5
as and of and thickness. of times as long wide) spongy tissue varying compactness Part mesophyll cells sometimes withthickened, lignified, pitted walls. Midribadaxially shallow- ly grooved, flattened, or raised, abaxially always prominently raised, supplied with a
closed not simple open or vascular system (type I or 2, fig. i), whether or accompanied by single or grouped sclerenchyma fibres. Cells with lignified pitted walls often present in ground tissue. Vascular bundles of minor veins embedded in mesophyll, supplied with
few often or sheathed sclerenchyma caps or sclerenchyma fibres, partly completely by a Petiole similar layer of translucent parenchyma cells. supplied with a vascular system to that of midrib and with grouped or solitary sclerenchyma fibres, rarely without scleren- chyma. Cells with lignified pitted walls often present in medullary tissue. Brachysclereids
or absent of and midrib clusters present in ground tissue petiole in and/or solitary. Crystals solitary or both solitary and clustered, scarce to abundant, most frequent in petiole,
minor in cells absent. midrib, and near veins, absent to scanty mesophyll. Crystarque
Kokoona I. coriacea King
Material: Malaya, King's collector 4226*.
S.V.: Stomata anisocytic-cyclocytic, with 3(4) subsidiary cells. T.S.: Anticlinal walls of adaxial epidermal cells tapering to narrow towards the mesophyll. Hypodermal cells some- what bigger than adjacent epidermal cells. Brachysclereids solitary and in conspicuous clusters.
2. Kokoona filiformis (Laws.) Fischer
Material: Thailand,Kerr 12450*.
S.V.: Stomata with —6 anisocytic and/or cyclocytic, 3, rarely 2 or 4 subsidiary cells.
T.S.: Lamina tending to isobilateral. Hypodermal cells somewhat bigger than adaxial epidermal cells. Brachysclereids absent.
3. Kokoona littoralis Laws. (Plate I, 3)
Material: Malaya, KEP. 94398*, S.F. 30177*; Sarawak, S. 17893*,S. 23654*, S. 23693, S. 9537, Smythies
15242*', Sumatra, Endert 349*, Endert 34E.1P.639, Soepadmo 168, Buwalda 7113.
S.V,: Stomata anisocytic-cyclocytic, with (2)3 —5(6) subsidiary cells. Granular material stomatal Anticlinal walls of sometimes present in pore. T.S.: unspecialized epidermal cells from thin thick. Anticlinal walls towards ranging to very tapering to narrow mesophyll thick-walled in specimens. Hypodermis consisting of I or 2 layer(s) of cells that are slightly bigger than adjacent epidermal cells in thick-walled specimens only. Spongy mesophyll cells in some specimens with thickened, lignified, densely pitted walls. Brachysclereids solitary or in conspicuous clusters.
Note: It is remarkable that the strikingly thick-walled epidermis of K. littoralis
(similar to situationin K. sessilis illustrated in Plate to be a variablecharacter. as 1,2) appears The whole from thin-walled thick-walled range fairly to extremely is present in the material Plate The that investigated. I, 3 represents an intermediate form. possibility we deal with different in slow leaf stages extremely development was ruled out by the finding of extremely thick-walled epidermal cells in some rather young leaves. BLUMEA VOL. XXI, No. I, 1973 158
+ + ± ± + + + -f- + ± ± crystals clustered ± ± ± ± —— + + + + + ± ± + + +
crystals solitary — ± — — ± — ± + + ± ± ±± ± — — +
brachysclereids + — + + + + + ± ± ± — ± — ± + +4- +
raised 1,2 1,21,2 system vase, 22 I1 1 1 1 1,2 2 petiole I I I I I1 I1 I1 I I1 I1 I1 I1 = R 11
systemsystem vase, 2 I1 I1 I1 I1 1 1 1 1 1 1 1 1 2 1 raised; midrib 1 1 1 1 1 I 2 1 1
1
fig. ad. shape SR ad. midrib of R F G G G G F R R slightly F G G G G G G F F F R R R to absent = =
SR flattened T.S. T.S.
in cells epidermal sq-fl sq-fl sq-fl sq-fl er-sq sq-fl er sq-fl sq-fl sq-fl sq-fler-fl sq-fl er-fl er-sq er-sq er-sq according = er shapes scarce; grooved; fl adaxial of types = 6 6 6 11 11 6 11 6 square; 6 6 6 II 11 6 II 12 12 1313 ± = walls cell periclinalpericlinal G = 6—8 4— 6—8 4— 4— 7— 9— 3—5 4—5 4— 7—9 9— 5-6 3—5 ab.ab. of thickness 10— 10— II— +systems: = — sq abundant; flattened; erect; 66 II11 10 1313 8 66 1212 1313 II walls cell periclinal = = 4— 9— 6—8 6—8 8— 3—5 5—7 7—9 5-8 KOKOONA vascular ad.ad. of thickness 7—9 9— 11— 3—5 4—5 6—8 4— 9— 8— F er II—
1:
18 Table 2020 18 2020 1818 2828 2121 1818 22 1818 2626 21 2222 2222 1919 1919 31 2929 2020 2020 2020 19 2020 1818 33 2929 38 16— 19— 16— stomataof length 18—15— 16—14— 18—15— 16—14— 22—24— 16—14— 16—13— 14—17—2214—17— 16—13— 22—24— 17—19— 17—19— 16—18— 16—14— 16—17— 20—25—31 21—24— 16—18— 16—18— 16—18— 16—14— 16—18— 16—14— 22—29—33 24—27— 30—32—38
20 16 1919 1717 2525 1919 18 19 1717 23 2020 2020 2020 19 34 2525 1414 1616 1515 1515 1515 1313 2828 2929
stomata 16—16— 15— 18—20 stomata of width 7—6——+8 9 ——+ 15—18—2015—18— 14—12—14—16 n—8—24—27—32n 3^—30— 18—15— 12—14— 16—17— 13—16— 16—17— 15—17— 15—17— 22—25—34 10—12— extremes 21—23— 16—14— 13—15—18 13— 13— 19—21—23 16— 14—17—19 22—19— 13—14— 12—13— 12—14— 12—13— 10—12— 26—23— 24—27— 24—27—32
5 I I1 I index 8.98-9 7.67-6 7.8 9.191 6.26.2 8.0 6.2 between index stomatal 12.0 12.0 IO.410.4 II.511. 12.4 II.11. 10.3 II. 10.5 µm
% 140140 ±20 lamina of thickness 280 6—330 8 6—2804——±8 6—3004—24——21—23—22—±7—8 4008—16—15—17—18—4709—13—14—16——16—250 14—16—10 15—13—16—13—11— 13— 2502504— 532074—5— 7—6—28020—22—25—14—87—15—17—16—93209—22—21—24—16—14—17—9 2305—— 2004—3—519— 16—10—12—18— 14—16—13—18— 250—23—26—9—22—29—+ 220—24—27—+ 370 10—12 11—10—12 in averages
indices thickness sizes: \N. 639 634 µm 635 168 iP. iP. 1158 in 24—21—22—13—16—13—15— 19—iP.16—17—18—20 16—17—19— 16—15—17—18— 16—12—13—18— 14—12—14— 16—12—13—18— iP. 14—16—10—12— character 13 stomatal stomatal 7113 47614 9159 12045 3P. 736 sizes lamina 94893 34E. 71 9159 7910 34E. 92 5375 30177 15242 349 ovatolanceolata 32394 34E. 36296 zeylanica coriacea littoralis 17893 23654 23693 9537 ochraceaochracea reflexa E. E. sessilis K.K.filifomxisfiliformis Soepadmo Buw. SAN. Kost. And. End. Sutr. Kost. FRI.FRI. K. K. KEP S.F. S. S. Sm. End. S. S.S- End. K.K. K. And. bb. K. End. bb.bb. bb. bb. K. S.F. K. Legend: N.n. specimenspecimen S. S. S. I.1. 2. 3-3. 4. 5. 6. 7.7. 8. Kokoona and W. T. JANSEN & P. BAAS: Leaf anatomy of Lophopetalum 159
4- Kokoona ochracea (Elm.) Merr.
Material: Indonesian Borneo, Kostermans 9159*; Sabah, SAN. 47614*.
the im- S.V.: Stomata anisocytic-cyclocytic with 3 or 4 subsidiary cells, often giving cells side of pression ofparacytic, but with one big lateral cell and 2 smaller on the other the guard cell pair. T.S.: Hypodermal cells bigger than adjacent epidermal cells. Brachy- sclereids solitary.
5. Kokoona ovatolanceolataRidley
Material: Indonesian Borneo, bb. 32394*; Brunei, Brun. 0822*, Brun. 5324*; Sarawak, Anderson 7910*, old Leiden*. S. 9224; 2-years seedling from the greenhouse of the Hortus Botanicus,
S.V.: Cuticle granular. Stomata cyclocytic, frequent in adaxial epidermis of bb. 32394, with 3—6 subsidiary cells. Granular material sometimes present in the stomatal pore. of flattened which be T.S.: Hypodermis absent in cultivated seedling, composed cells may slightly bigger or smaller than epidermal cells in other specimens. Clustered crystals abundant in hypodermis of Anderson 7910 and in adaxial epidermis of bb. 32394. Abundance of the adaxial of bb. and absence in Note: stomata in epidermis 32394 led the this No other material Anderson 7910 to checking of more specimens for feature. than the former was found to have stomata in the adaxial epidermis.
6. Kokoona reflexa (Laws.) Ding Hou
Material: Indonesian Borneo, Sutrisno 92; Sumatra, bb. E. 736, bb. E. 3P. 1158, bb. 34E. 1P. 633*, Endert
P. Kostermans 34E. 1 634*, 12045.
with S.V.: Stomata anisocytic-cyclocytic 3 or 4 subsidiary cells, resembling paracytic ochracea. differentiated stomata as in K. T.S.: Hypodermis poorly from mesophyll.
7. Kokoona sessilis Ding Hou (Plate I, 2)
Material: Malaya, S.F. 36296*, KEP FRI. 5375*.
walls of cells with thin- S.V.: Anticlinal epidermal very thick. Stomata cyclocytic 4—7 walled cells. Granular material stomatal subsidiary present in pore. T.S.: Lamina tending to isobilateral. Anticlinal walls of unspecialized epidermal cells bulging (cf. Plate I, 1).
Hypodermal cells ca. 1/3 in height ofepidermal cells. Brachysclereids solitary or in conspicu- ous clusters.
Note: The outstanding thickness ofthe anticlinalwalls ofthe unspecialized epidermal cells also found of of was in some specimens K. littoralis and in the adaxial epidermis
Lophopetalum pachyphyllum (Plate I, 2).
8. Kokoona zeylanica Thwaites (Plate III, 2)
Material: Ceylon, Thwaites C.P. 2584*.
S.V.: Abaxial cuticle with reticulate thickenings on inner side. Stomata cyclocytic with
6—9 subsidiary cells. T.S.: Hypodermis consisting of an abaxial layer of cells which are considerably bigger than adjacent epidermal cells, interrupted by stomatal air cavities. BLUMEA VOL. XXI, No. 160 I, 1973
2—8: in Figs. Lophopetalum,vascular system as seen T.S. ofpetioles and midribs, all X 13. — 2. L. beccaria- num (SAN. 52957), midrib. — 3. ibid., basal end of petiole. — 4. L. glabrum (C.F. 34564), distal end of
— —8.L. — — — petiole. 5 javanicum. 5. bb. Cel/V—145, distal end ofpetiole. 6. ibid., basal end ofpetiole. 7.
SAN. 25059, distal end of petiole. — 8. ibid., basal end of petiole. W. T. JANSEN & P. BAAS: Lea anatomy of Kokoona and Lophopetalum 161
cells with thin Part of palisade parenchyma periclinal division walls, part of spongy
mesophyll cells with slightly thickened walls. Vascular system of basal end of petiole
composed of 6 free vascular bundles.
The of the side of the Note: presence of a hypodermis on the lower instead upper lamina is noteworthy in this specimen.
Lophopetalum Wight ex Arn. Plate and and (Table II; I, i, 4, 5, 6; II, 1—6; III, 1 3—6; Figs. 2 —22).
S.V.: Cuticle smooth to coarsely granular and/or slightly striated. Unspecialized epider- Thin mal cells polygonal, with straight to undulated anticlinal walls. areas of cuticle in
loops of undulations and/or cell corners or pit-like structures in cell corners sometimes
present. Anticlinal walls fairly thin except in adaxial epidermis of L. pachyphyllum, cuticular flanges rarely pitted. Minor veins mostly not prominent in adaxial epidermis, in fairly prominent abaxial epidermis of most species. Stomata present on both surfaces
or confined to abaxial epidermis, anisocytic, helicocytic, cyclocytic, or complex aniso- or often different with cyclocytic, present as types in one specimen, 3—5(11) subsidiary
11 ratios Outlines ofsubsidi- cells, —38 fim 1., 7—29 /mi w., length-width 1.0—1.3—1.7. thin here. and ary cells sometimes difficult to observe because of the cuticle being Outer
inner stomatal ledges inconspicuous to fairly conspicuous, small polar T-pieces rarely Granular material absent from stomatal present. pores or very inconspicuous. Subsidiary Stomatal in cells partly underlying guard-cells. index 3.7—11.3%. Cork warts present
varying amounts. T.S.: thick. cells Lamina dorsiventral, 150—450 gm Unspecialized epidermal rectangular,
tall to flattened, those of abaxial epidermis mostly lower than those ofadaxial epidermis.
Anticlinal walls sometimes tapering towards the mesophyll, those of adaxial epidermis
of L. pachyphyllum tapering and bulging like in Kokoona sessilis (cf. Plate I, i). Outer periclinal walls of both epidermides sometimes with thin places or pitlike structures cuticle next to cell corners, corresponding to thin areas of as seen in surface view. Abaxial
epidermis in some species with low or high papillae. Thickness of outer periclinal walls
some of i adaxial I—l 2 [xm. Hypodermis present in species only. Mesophyll composed —s of tall and layers square to very palisade parenchyma cells spongy tissue of varying and compactness thickness. Part of mesophyll cells with thickened, lignified, pitted walls in many species. Midrib adaxially shallowly grooved to prominently raised,abaxially always prominently raised, with a simple open to complex closed vascular system without with whole or i—3 medullary bundles; the system surrounded by an interrupted to closed ring of sclerenchyma fibres. Cells with thickened, lignified, densely pitted walls the Vascular present in medullary tissue except in L. subobovatum and L. sessilifolium. bundles of minor veins embedded in mesophyll except for the species with prominent veins in surface view where dorsiventral bundle sheath extensions to the epidermis sheathed are present, provided with sclerenchyma caps, whether or not by a layer of translucent parenchyma cells. Petiole supplied with a vascular system ranging from closed basal complex open to in end, whether or notwith one or more medullary bundles, distal end with least latero-dorsal bundles in always at 1 medullary bundle, wing and/or fibre strands wing present in distal end. Sclerenchyma fibres mostly present like hi midrib. Cells with thickened, lignified, densely pitted walls often present in medullary tissue. noted in L. L. Brachysclereids only pachyphyllum and one specimen of javanicum. sometimes Crystals mostly solitary, clustered or both clustered and solitary. Crystarque cells in present some species. 162 BLUMEA VOL. XXI, No. i, 1973
Figs. 9—13: Lophopetalum multinervium, vascular systems as seen in T.S. of petioles and midribs, all X 13.
— midrib. — ibid. basal end of — Kostermans basal end of — 9. bb. 8953, 10. petiole. 11. 10375, petiole. 12. bb. distal end of — Kostermans distal end of 8953, petiole. 13. 10375, petiole.
I. Lophopetalum arnhemicum Byrnes
Material: Australia, Byrnes 1180*.
S.V.: Stomata complex cyclocytic. Minor veins prominent in adaxial epidermis. T.S.:
Features as listed in table II.
beccarianum 2. Lophopetalum Pierre (Plate III, 5 & 6; figs. 2 & 3)
Material: Sabah, Ping Sam A. 1896*; Sabah, SAN. 37508, SAN. 52957*; Sarawak, S. 23829.
with thin of S.V.: Outer periclinal walls of epidermal cells areas of cuticle in loops
Polar undulations and/or cell corners. Stomata anisocytic-complex anisocytic. T-pieces faint to fairly conspicuous. T.S.: Mesophyll with numerous thickened, lignified, pitted walls. Midrib: vascular in SAN. Petiole: vascular system very complex 52957 (fig. 2). basal end of system in SAN. 32937 as in fig. 3. Kokoona and 163 W. T. JANSEN & P. BAAS: Leaf anatomy of Lophopetalum
3. Lophopetalum duperreaneum Pierre
Material: South Vietnam (Cochinchina), Pierre 4082*.
S.V.: Stomata anisocytic-complex anisocytic-complex cyclocytic. Incomplete stomata
with one guard cell infrequently present. T.S.: Hypodermis present at adaxial side, clearly
differentiated near veins, faindy differentiated in other areas.
floribundum 4. Lophopetalum Wight
Material: Malaya, KEP. 99907*, FRI. 4800*.
of cells S.V.: Outer periclinal walls epidermal without or with rather inconspicuous thin
areas of cuticle in loops of undulations and/or cell corners. Stomata complex anisocytic in of cells FRI. 4800, complex cyclocytic in KEP. 99907. T.S.: Part spongy mesophyll in
FRI. 4800 with lignified, densely pitted walls.
Note: The two specimens show rather striking differences, e.g. in abaxial epidermal cell stomatal and of cells and pattern, sizes types, presence crystarque lignified mesophyll cells (Table II).
5. Lophopetalum glabrum Ding Hou (Plate III, 1, 2, 4; fig. 4)
Material: Brunei, CF. 34564*-, Sarawak, S. 20487.
of with thin of cuticle of S.V.: Outer periclinal walls epidermal cells areas in loops adaxial Stomata undulations and cell corners, particularly conspicuous in epidermis. sometimes of anisocytic, mostly complex anisocytic or helicocytic. T.S.: Part spongy with walls. Petiole: vascular distal end of mesophyll lignified system as in fig. 4 in CF. 34564, slightly different in basal end and in S. 20987(Table II).
6. Lophopetalum javanicum (Zoll.) Turcz. (Plate I, 5; III, 3; figs. 5 —8)
6602 bb. Material: Indon. Borneo, Endert 5150*, Kostermans & 9580; Sabah, SAN. 23059*; Celebes,
— bb. bb. Cel/V 145*2 New Guinea, 30540; Moluccas, bb. 29957 ; Philippines, Sulit 12360; Sumatra, 6426*, Endert 48E. lP. 538*, Forbes 3151*.
S.V.: Stomata often complex, anisocytic-cyclocytic-helicocytic. Polar T-pieces faintly
T.S.: bb. cutinized present. Hypodermis present in Cel/V —145 and probably in bb. 29957(2 cell layers: epidermis + hypodermis notedin cuticular preparations), at adaxial side. Part of mesophyll cells with thickened, lignified, pitted walls in some specimens. Petiole: vascular & & 8 system very complex in bb. Cel/V—145 (figs. 5 6), as in figs. 7 in SAN.
25059.
7. Lophopetalum ledermannii(Loes.) Ding Hou
Material: New Guinea, bb. 30964*, Aet & Idjan 388*.
S.V.: Outer periclinal walls of epidermal cells with thin areas of cuticle in loops of undulations and cell in Aet & in corners Idjan 388, inconspicuously so bb. 30969. Stomata mostly complex, anisocytic-cyclocytic. Minute polar T-pieces usually present. T.S.:
Hypodermis present in Aet & Idjan 388 only, poorly differentiated. Part of mesophyll cells with thickened, lignified, pitted walls. 164 BLUMEA VOL. XXI, No. i, 1973
vascular in T.S. of and all L. Figs. 14—22:Lophopetalum, systems as seen petioles midribs, x 13. — 14—18.
— S. — FRI. — — pachyphyllum. 14. 16212, midrib. 15. 0632, midrib. 16. ibid., basal end of petiole. 17. S. basal of end 16212, end petiole. — 18. ibid., distal end of petiole. — 19. L. pallidum(K.L. 1566), basal of
— — L. — petiole. 20. ibid., distal end of petiole. 21. torricellense (BW. 5683), distal end of petiole. 22. L. wightianum (Endert 59E. 1P. 567), basal end of petiole. Kokoona W. T. JANSEN & P. BAAS: Leaf anatomy of and Lophopetalum 165
8. Lophopetalum macranthum (Loes.) Ding Hou
Material: New Guinea, Gjellerup 701*, NGF. 37402*.
S.V.: Outer periclinal walls of epidermal cells with rather thin areas of cuticle in cell corners at adaxial side. Stomata frequently complex, anisocytic-cyclocytic. T.S.: Features as listed in table II.
9. Lophopetalum micranthum Loes. (Plate I, 6)
Material: New Guinea, Versteeg 1773*.
S.V.: Abaxial cuticle coarsely granular except over the stomatal complex. Stomata complex cyclocytic. T.S.: Part of mesophyll cells with thickened, pitted, lignified walls. Midrib: vascular of number of system composed a more or less free vascular bundles.
10. Lophopetalum multinervium Ridley (figs. 9—13)
Material: Indon. Borneo, Kostermans 10375*: Sumatra, bb. 8953*.
Small S.V.: Stomata complex cyclocytic. polar T-pieces mostly present. T.S.: Hypo- dermis of one or two adaxial cell layers present, poorly differentiatedin some places. Part of mesophyll with thickened, lignified, pitted walls. Midrib flattened and broad, vascular system as in fig. 9. Petiole: vascular system very complex, particularly in basal end (figs,
in distal end in io & n). Vascular system as figs. 12 & 13.
11. Lophopetalum pachyphyllum King (Plate I, 1; II, 1 & 2; figs. 14—18)
Material: Sarawak, S. 16212*: Malaya, FRI. 0632*.
S.V.: Anticlinal walls of epidermal cells very thick in adaxial epidermis; adaxial periclinal walls with in cell pit-like structures corners; abaxial cells with well-developed papillae.
Stomata cyclocytic, difficult to observe because of overarching papillae ofsubsidiary cells
veins walls of (Plate II, 2). Papillae on arranged in rows (Plate II, 1). T.S.: Anticlinal sessilis adaxial epidermal cells as ik Kokoona (Plate I, i). Lumina of epidermal cells narrow in acute to obtuse tall. Midrib: vascular thick-walled, papillae. Papillae ca. 30—60/an in in vascular system as fig. 14 (S. 16212) or fig. 15 (FRI. 0632). Petiole: system very
basal end distal end as 18. Asterosclereids complex in (figs. 16,17), in in fig. present through- out mesophyll.
Laws. 12. Lophopetalum pallidum (Plate I, 4; figs. 19 and 20)
Material: Indon. Borneo, bb. 18997*; Malaya, KL 1566*.
S.V.: Cuticle coarsely granular in KL. 1566 striated in bb. Anticlinal walls of , 18997. cells epidermal conspicuously pitted. Stomata anisocytic or cyclocytic, rarely complex anisocytic. T.S.: Outer periclinal walls ofabaxial epidermal cells dome-shaped (very lowly lamina papillate). Stomata sunken below surface. Petiole: vascular system of KL 1566 as in and fig. 19 (basal end) fig. 20 (distal end).
This L. Note: species, together with wightianum, shows a very simple arrangement of vascular and midrib. the basal of the of tissue in petiole Particularly part petiole L. the the pallidum recalls situationin Kokoona. Yet, medullary bundles are present in distal end of the petiole in this species. 166 BLUMEA — VOL. XXI, No. 1, 1973
brachysclereids. fl; + + — ± ± — ± + ± ±± + ± \V \ 1\
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388 567 543 1 3219 701 2236 iP. i -2PPS7 Idjan » 12400 811 /eder/fia/mii/eder/fia/wii 374.02 1037510375 16814 coll. 17833 5683 1053 coll. 30969 & 0632 71523 12326 Su2lC macratithummacranthum multinervium 8953 18997 1566 59E. l>£>.bb. micranthum multinervium pachyphyllumpachyphyllum 16212 pallidum rigidum sessilifolium 13737 subobovatumsubobovatum torricellense wallichii wightianum A.Z.. bb. Aet Gjellerup bb.18997 Haviland L. NGF. Kost. bb. SAN. Native SAN. KEP. T.G.H. Garrett Native Gallatly S. FRI. K.L. .L. KEP. Brass End. L.L. L. bb. L. bb. B.W. End. 7.y. L. L.L. L. L. S. L."'' L. L. L. 8. 9"9. 10. ii.11. 12. 13-13. "14. 15. 16. 17-17. 18. 168 BLUMEA VOL. XXI, No. i, 1973
13. Lophopetalum rigidum Ridley
Material: Sabah, SAN. 16814*; Sarawak, Haviland 2236*.
S.V.: Outer periclinal walls of epidermal cells with thin areas of cuticle in loops of
and cell undulations corners. Stomata anisocytic in SAN. 16814, anisocytic to complex
anisocytic in Haviland 2236. T.S.: Part of mesophyll cells with thickened, lignified walls.
sessilifolium 14. Lophopetalum Ridley
Material: Sarawak, S. 13737*, Native collector 543*.
S.V.: Outer periclinal walls of epidermal cells with thin areas of cuticle in loops of
undulations and cell corners. Stomata anisocytic, complex anisocytic, or helicocytic. T.S.:
Features as listed in Table II.
15. Lophopetalum subobovatumKing
Material: Malaya, KEP. 71523*] Sabah, SAN. 37833*.
S.V.: Stomata anisocytic or complex anisocytic-complex cyclocytic. Small polar
at adaxial side. T-pieces usually present. T.S.: Hypodermis poorly differentiated, present Palisade parenchyma cells extremely tall.
16. Lophopetalum torricellense Loes. (fig. 21)
Material: New Guinea, Brass 12326*, BIV. 5683*, Hartley 12400.
Small S.V.: Stomata complex, anisocytic-cyclocytic. polar T-pieces present in BW. Petiole; vascular distal end o( 5683. T.S.: system very complex in BIV. 5683 (fig. 21).
Solitary crystals abundant in epidermis ofBIV. 5683.
17. Lophopetalum wallichii Kurz (Plate II, 3 and 6)
Garrett Native Material: Thailand, 1053*, collector 321 g*.
S.V.: Stomata anisocytic-cyclocytic, occasionally complex. T.S.: Outer periclinal walls of abaxial epidermal cells low dome-shaped (sometimes lowly papillate) (Plate II, 3).
cells. Minute crystals present in epidermal
18. Lophopetalum wightianum Arn. (Plate II, 4 and 5; fig. 22)
Material: Burma (Tenasserim), Gallatly 811*] Sumatra, Endert 5gE. lP. 567*] and nail varnish replicas of
3 more specimens to check for presence of papillae.
S.V.: Stomata mostly anisocytic, sometimes cyclocytic or complex anisocytic. Many irregular cork formations and necrotic cells in lower epidermis of Gallatly 811. T.S.:
Abaxial epidermal cells lowly papillate in Gallatly 811. Guard cells sunken. Petiole: vascular
in in basal end. in system as fig. 22 Solitary crystals abundant epidermis of Gallatly 811; in subepidermal layer of Endert 59E. lP. 367. W. T. JANSEN & P. BAAS: Leaf anatomy of Kokoona and Lophopetalum 169\I
T.S. adaxial showing bulging Plate I. 1. Lophopetalum pachyphyllum (S. 16212), through epidermis, — cuticu- anticlinal walls and pit-like structures in outer cell corners, X 825. 2. Kokoona sessilis (FRI. 5375),
— littoralis cuticular preparation lar preparation of thick-walled epidermis, X 330. 3. Kokoona (S. 17893), wall of abaxial epidermis with intermediate cell thickness, X 530. — 4. Lophopetalum pallidum (ibb. 18997), cuticular preparation of abaxial epidermis with anisocytic stoinata, X 330. — 5. Lophopetalum javanicum
(Sulit 12360), cuticular preparation of abaxial epidermis with complex anisocytic stomata, X 530. — 6. Lophopetalummicranthum (Versteeg 1773), cuticular preparation of abaxial epidermis with complex cyclocytic stomata and granular cuticle, X 530. 170\II I3LUMEA VOL. XXI, No. I, 1973
Plate II. 1—4. Scanning electron photomicrographs of abaxial leaf surfaces. — 1. Lophopetalum pachy-
rows over — of phyllum (S. 16212), papillae, note arrangement in vein, x 800. 2. ibid., group papillae overarching a stoma and covered with waxy particles, X 2000. — 3. Lophopetalum wallichii (Garret 1053), and between low confined in — papillae stomata, waxy particles to areas papillae, x 2000. 4. Lophopetalum 81 wightianum (Gallatly 1), low papillae and waxy material, x 4000, fungalhyphe present on all leaf surfaces.
— with — 6. wallichii coll. 5. ibid., upper epidermis prismatic solitary crystals, x 330. Lophopetalum (Native with minute cubical 3219), upper epidermis crystals, X 530. W. T. JANSEN & P. BAAS: Leaf anatomy of Kokoona and Lophopetalum 171\III
Plate III. Lophopetalum glabrum (S. 20987), cuticular preparation of abaxial epidermis with undulated anticlinal walls and complex anisocytic to helicocytic stomata, X 330. — 2. Kokoona zeylanica (Thwaites C.P. section abaxial with Note submersion of 2584), free hand of epidermis cyclocytic stomata. subsidiary
— section cells, x 530. 3. Lophopetalumjavanicum (Forbes 3151), free hand of abaxial epidermis with com- — plex anisocytic stomata. Submersion of subsidiary cells only partial, x 530. 4. Lophopetalum glabrum
with thin of (S. 20987), cuticular preparation of adaxial epidermis areas cuticle showing up as light spots, A. x 310. — 5. Lophopetalum beccarianum (Ping Sam 1896), free hand section of adaxial epidermis with thin
cuticle in wall — 6. ibid. T.S. of areas of outerpericlinal showing up as light spots, x 530. petiole with cells in crystarque ground tissue, X 330. W. T. JANSEN & P. BAAS: Leaf anatomy of Kokoona and Lophopetalum 173
Camera lucida of leaf of Kokoona ovatolanceolata. stained Fig. 23. drawings young epidermis Densely ceils dotted. Arrows indicate of cell division. Mitotic division products inequal spindles present. Note parallel in in to guard cell pair a and perpendicular to future guard cell pair b.
DEVELOPMENT OF THE STOMATAL COMPLEX IN KOKOONA OVATOLANCEOLATA
Several old ofK. ovatolanceolata in the Hortus Botani- two years seedlings were present cus, Leiden, which enabled us to study the ontogeny of the stomatal complex in this
Free hand sections of aceto-orceine species. very young leaves were stained with 2% or surface aceto-carmine. The stomatal development as seen in view is as follows: Following inequal cell divisions protoderm cells give rise to a small stoma mothercell, distinguishable of and by its intense uptake the stains used, a bigger daughter cell which soon becomes indistinguishable from the other neighbouring cells.
The stoma mother cell divides equally and gives rise to the two guard cells. In mature the cells the cells almost specimens subsidiary underly guard partly or completely. Con- sidering the different stages of development observed, this must be due to enlargement of the guard cells. The final numberof subsidiary cells depends on the number of cell divi- sions in the neighbouring cells during the early stage of leaf development. The develop- ment of stomata in K. ovatolanceolata thus conforms to the mesoperigenous type (see fig. 23).
DISCUSSIONS
Stomata
The is Kokoona presence of different stomatal types within one genus remarkable. has diverse both anisocytic and cyclocytic stomata. Lophopetalum is even more in stomatal addition and in type in showing in complex types a tendency for helicocytic stomata part of the The fact that several the leaf and that often species. types may occur on same the transitionsbetween the types (see fig. 24) have beenrecorded, limits taxonomic use of the type of stomata in both Kokoona and Lophopetalum. Only the restriction of complex
of the of of these as types to part species Lophopetalum is some help in recognizing species the is belonging to Lophopetalum. Moreover, submersion of subsidiary cells more pro- nounced in Kokoona than in Lophopetalum (see Plate III, 2 and 3).
The ontogeny of stomata in K. ovatolanceolata is roughly similar to that reported for other of Celastraceae Pant and Kidwai It differs in genera by (1965). slightly showing
173 VOL. No. 174 BLUMEA XXI, i, 1973
Stomatal in Fig. 24. types Lophopetalum. a: anisocytic; c.a. complex anisocytic; c: cyclocytic; c.c.: complex indicate of between different cyclocytic; h: helicocytic. Arrows possibilities relationships types, presumably throughformation of extra division walls.
less the cell addition to those to divisions more or perpendicular to guard pairs in parallel the latter the guard cells. Pant and Kidwai only reported for the genera they studied.
of the stomatal in would be for An ontogenetic study types Lophopetalum important a of The reliable interpretation of the diversity found in the stomatal complex that genus.
of cells some much the of arrangement subsidiary in species gives so impression having Plate that originated from a single cell (cf. III, i) one may tentatively anticipate meso-
in those The and genous stomata species. complex anisocytic cyclocytic types are presumably elaborations arising through subdivision of the subsidiary cells. Subdivision of subsidiary cells has also been recorded for other genera of Celastraceae by Rehfous Metcalfe and Chalk (1914). Unfortunately, his paper was misinterpreted by (1950: 390) and Pant and Kidwai (1966: 288) who suggested that Rehfous regarded the guard cells to be subdivided, though in fact he referred to subdivisions of the subsidiary cells.
The helicocytic stomata in some species ofLophopetalum (Plate III, 1) are never a very The cells of dominant feature. spiral of subsidiary declining size as described for this type
is by Payne (1970) never elaborate and subdivisions parallel to the outer guard cell walls obscure the helicocytic pattern. W. T. Jansbn & P. Baas: Leaf anatomy of Kokoona and Lophopetalum 175
Several authors (e.g. Gupta, 1961) have regarded the stomatal index as a reliable diag- nostic feature at the species level. The variation recorded in TableI and II for K. littoralis andL. javanicum clearly show that this is not valid for those species. The conclusion that stomatal character also the index is a very variable in general is supported by current studies on Icacinaceae and Winteraceaein the Rijksherbarium.
Vascular system of the petiole and midrib
Kokoona the vascular of the of the In system petiole is always simple type (fig. i, in in types 1 and 2). The two types are represented one petiole some specimens (K. sessilis and K. ovatolanceolata, see TableI).
the and much In Lophopetalum interspecific intraspecific differences are bigger. In many
of from the basal end distal end of the specimens a great change type to the petiole was found This makes the of vascular (e.g. L. javanicum, L. multinervium). variation type system of the petiole an unreliable character on the species level, although series of transverse sections might give more information.
Comparison of transverse sections of the petioles of Kokoona and Lophopetalum in the distal in of the As stated end results an easy separation two genera. before, we found no of Kokoona with vascular the distal of the and specimen a complex system in part petiole, in of no specimen ofLophopetalum with a simple vascular system the distal end the petiole was found either.
The vascular system of the midrib usually varies within one specimen. This is due to the often looks splitting offfrom the midrib of the lateral veins. It like the vascular system in the distal end of the petiole, but at the places of insertion of lateral veins it may be more complex.
Variability ofanatomical characters in certain species
Kokoona littoralis
A study of 11 specimens of Kokoona littoralis, a species with leaves that show much variability in shape, texture, and size (Hou, 1963:261), has shown that there is anoverlap anatomical in characters with those of the other species and that many leaf anatomical characters are variable in this species. Thickness of anticlinal walls of unspecialized epidermal cells: Extremely thick walls found like Kokoona sessilis thin walls were in S. 9537, just in (Plate I, 2), very in most of specimens. Presence specimens with intermediate wall thickness (S. 17893 and to a continuous variation of the character lesser extent Smythies 15242) suggests a (Plate I, 3). Number of cells: The number from the thick- subsidiary highest ranges 3—5(6) in walled the lowest from in of the thin-walled specimens, (2)3 or 4 most specimens.
Diameter of the guard cell pairs: Thin-walled specimens mostly have small stomata
than thick-walled much stomata (in (less 20 jum wide), specimens bigger any case over
20 The specimen Smythies is an with thick walls and guard fim wide). 15242 exception, cell of wide. pairs 13—15 —18 /tm
Kokoona reflexa
encountered In the six specimens used for cuticular preparations much less variation was than in K. littoralis. The two specimens used for transverse sections only showed some differences in the shape ofthe adaxial epidermal cells and the frequency of brachysclereids. 176 BLUMEA VOL. XXI, No. x, 1973
Kokoona ovatolanceolata
The of stomata in the of occurrence numerous upper epidermis one specimen (bb. is also particularly because it is the 32394) very remarkable (see p. 159), only specimen of the out the whole genus with this feature. Dr. Hou kindly checked and confirmed identity of the specimen.
Lophopetalum javanicum and other Lophopetalum species variationin Lophopetalum javanicum, a species witha high degree ot macro-morphological leaf characters, was chosen for a study of n specimens for cuticular preparations, 6 of will which were also used for transverse sections. Variability of several characters be discussed belowand compared with that in some other species. which Lamina thickness: This is a highly varying character, is not very surprising, because of the fact that lamina thickness of sun and shade leaves from the same tree can be
different. Tables I and II show that most species Kokoona as lamina very (cf. well) vary in thickness.
of in Presence stomata upper epidermis: The whole range from virtually absent to
lamina in for which abundant over the whole occurs Table II, herbarium material from
nail varnish in total. In different regions was checked, using replicas, 32 specimens ca. 70% the adaxial The of the specimens stomata were found in epidermis. number of specimens whether of in appeared to be insufficient to see the presence stomata the adaxial epidermis exhibits The absence of is also variable a geographical pattern. presence or stomata in some other species, e.g. L. ledermannii, L. torricellense. variable Sizes of stomata are rather in L. beccarianum, L. floribundum, L. ledermannii, L. and pallidum, L. rigidum, L. subobovatum, L. wightianum. In L. javanicum they are rather stable, but Endert 48E. lP. 338 differs widely from the other specimens (Table II). Undulations of anticlinal walls of cells rather for epidermal are constant L. javanicum and also for the other species.
Thickness of outer walls of cells: all of periclinal epidermal 1—2 jum in specimens
L. javanicum, but other species show great variation(J(L. floribundum, L. subobovatum). ofadaxial ofmidrib: but Shape part Not constant inL. javanicum (S. 2y05g), constant in most other species.
Vascular systems of petioles and midribs, see p. 175 of Crystarque cells were found in some specimens L. javanicum'■ and in one specimen of other L. ledermannii only. In species the presence or absence of crystarque cells seems to be constant. Particularly if they are infrequent there is a fair chance of missing them in a transverse section, which may have slightly influenced our results as listed in Table II.
Crystarque cells have previously not been recorded for the Celastraceae. These unusual
from The cells are now known 7 families (see Baas, 1972). fact that they are not a constant of indicates of their feature in some species Lophopetalum a restriction diagnostic value in this genus.
The in presence or absence of crystals, known to be a very variable feature many plant be for the material of studied. This also groups, appears to quite constant Lophopetalum is true for Kokoona.
Diagnostic and taxonomic value of some leafanatomical characters
the of From variation the characters discussed above it becomes apparent that these are of doubtful restricted and value Kokoona and very or diagnostic taxonomic in Lophope- talum. Very outspoken and unusual anatomical characters like extremely thick-walled W. T. JANSEN & P. BAAS: Leaf anatomy of Kokoona and Lophopetalum 177
cells and cells be of value the epidermal crystarque appear to no diagnostic on species of level for some species. Other characters like stomatal size, numbers subsidiary cells, of of the vascular in the presence or absence of hypodermis, degree complexity system also characters of petiole etc. are exhibiting a great range variation below the species level.
and much less so and Papillae conspicuously present inL. pachyphyllum in L. wightianum
L. wallichii (Plate II) have been recorded to be inconstant in species of Cratoxylum (Gutti- but ferae) by Baas (1970). In both specimens of L. pachyphyllum they were present, only of the showed low Both one five L. wightianum specimens studied papillae. L. wightianum
and L. pachyphyllum belong to pollen group C ofHou (1969). Apparently this correlation of epidermal papillae and the presence of conspicuous subglobose verrucae on the pollen
of these because the of is not for species is insignificant, presence papillae constant L. L. different wightianum, and wallichii with very low papillae belongs to a pollen group.
Thin areas of cuticle and/or pit-like structures in the corners of epidermal cells are
constant features for L. beccarianum, L. floribundum, L. glabrum, L. ledermannii, L. macran-
thum, L. pachyphyllum, L. rigidum, and L. sessilifolium. They are absent from all other
ofthe species studied. Most above mentioned species belong to pollen group B as classified and by Hou. However, L. pachyphyllum L. sessilifolium belong to his group C and D
of that respectively. Moreover, some species of group B do not show thin areas cuticle so correlation ofpollen characters with this leafanatomical character also seems insignificant. other found of the distribution showed No characters were which over the species a clear correlation with pollen characters.
In of variation recorded spite the tremendous here, an attempt was made to compose a synoptical key to the species based on leaf anatomical characters. Tests of small the key, however, revealed that only a very proportion of the species could be identified here. satisfactorily and we therefore refrain from publishing it
GENERAL CONCLUSIONS
view Leaf anatomy lends some support to the that Lophopetalum and Kokoona are distinct fromeach other. the vascular ofthe distal end of the genera Only anatomy petiole the provides an absolute character by which two genera can be recognized. The other differential listed hold for of be characters, below, only part the species and may inter- preted as trends of generic differentiation.
Kokoona Lophopetalum
Vascular of distal of Vascular of distal of 1. system part petiole simple. system part petiole complex.
2. Stomata never complex. Stomata often complex.
absent. in 3. Hypodermis very rarely Hypodermis present 3 species only.
4. Anticlinal epidermal cell walls straight to slightly Anticlinal epidermal cell walls straight or sinuous from curved. or ranging straight to sinuous.
in 5. Thin areas ofcuticle and/or pit-like structures Thin areas and/or pit-like structures present in 8
cell corners absent. species.
6. Crystarque cells absent. Crystarque cells present in 12 species.
It is related of within either impossible to recognize closely groups species (sections) the level be Lophopetalum or Kokoona. Each character, as far as it is constant at species may used to divide the genera into two groups, but these never coincide for the different characters there correlation with based employed, nor is a groupings on pollen morpholo- Hou also refrained from the because of similar lack of gy. (1963) subdividing genera a correlation of macromorphological characters. BLUMEA VOL. XXI, No. I, 1973 178
ACKNOWLEDGEMENTS
due Dr. Hou in Os Thanks are to Ding for his suggestions and interest this study, to Mr. J. van for his
B. help in drawing the figures, and to Mr. N. Kieft and Mr. C. L. Marks for their share in producing
Plates. Dr. C. the Mr. J. Bongers kindly tested the synopticalkey. Prof. Kalkman critically read the manus- cript.
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