Pollen morphology of
the genus Hydnocarpus (Flacourtiaceae)
with notes on related genera
J. Schaeffer
Rijksherbarium, Leiden
Contents
Summary 65
I Introduction 65
II General morphology 66
III Systematic descriptions 66
IV Pollen types 76 Pollen and V morphology taxonomy 77
References 79
Summary
Pollen of ofthe described. Two grains 34 species genus Hydnocarpus(Flacourtiaceae) are pollen types, one of which is subdivided in two subtypes, are distinguished. Within Flacourtiaceae the pollen of Hydnocarpus
In is more or less isolated, but the related genus Chlorocarpa has rather similar pollen. sculpture there exists some resemblance to Paropsia (Passifloraceae).
I. Introduction
The present study forms part of a general pollenmorphological survey of the family
Flacourtiaceae, initiated at the Rijksherbarium in collaboration with Dr. H. Sleumer. The
selected for the first detailed because genus Hydnocarpus was study it proved to have rather characteristic pollen types and is concentrated in SE. Asia and the Malesian region.
Moreover, an adequate amountof well-determinedmaterial as well as a recent revision ot
In the genus (Sleumer, 1954) were available. total 34 species could be investigated, of the total numberof known. representing 85 % species In addition, the pollen of a large other number of Flacourtiaceae was cursorily examined, in order to asses the pollen
of within the morphological relationships Hydnocarpus family. The pollen of the genera and which resemble that of described Neoptychocarpus Chlorocarpa, Hydnocarpus most, is in more detail. Erdtman described Previously, only (1952) has, very briefly, the pollen of Hydnocarpus rather the elmeri which, however, proved to be atypical of genus.
The pollen material was acetolysed for two minutes, mounted in glycerin jelly and photographed with a Leitz apochromatic OI objective (90/1.40). Acetolysed grains, covered with a thin layer of gold, were photographed under a Cambridge All Scanning
Electron Microscope at the Geological Institute, University of Leiden. Unless indicated otherwise, all material examined originated from the Rijksherbarium (L).
65 No. 66 BLUMEA VOL. XX, I, 1972
Mr. I sincere This study was performed under supervision of J. Muller. express my thanks to Dr. H. Sleumer for providing the material on which this study is based, for discussions and taxonomic corrections, and to Mr. Muller for critically examining the
Finally, take this to gratitude to Professor C. G. G. manuscript. I opportunity convey my who has been teacher and under whose J. van Steenis, my in plant taxonomy guidance of in the I have had the privilege working Rijksherbarium for two and a half years.
II. GENERAL MORPHOLOGY
The size pollen grains in Hydnocarpus are single, isopolar and tricolporate. Average
between and Shape, defined by the ratio of length polar axis (P) to varies 17 43 [i. as length equatorial axis (E), is generally spherical but may vary between suboblate, sub- prolate and, rarely, prolate. The equatorial outline is always circular. whenever from well Size measurements were taken, possible, ten developed grains and minimum, average, and maximumsizes are given separately for P and E. Under the endexine and while the latter lightmicroscope an an ektexine can be recognized, is further differentiated into a layer of columellae and a reticulate tectum. The inner surface of the endexine be smooth. The columellae two different appears to occur in types, less in in a) supporting columellae which are more or distinctly arranged single rows a reticulate and which the of the reticulum with their somewhat pattern support muri swollen heads, and b) free columellae, which are shorter, with roundedtops, more variable
in lumina of The in diameter and occurring scattered the some species. presence or absence of these free columellae forms an important criteriumfor the definition ofpollen for types in Hydnocarpus. Size and distribution pattern of the lumina are also important the others this purpose. Some species are characterized by a finer reticulum on poles, by a more even size distribution of the lumina or by finer lumina near the ektoapertures.
There is also a rather marked differencebetween grains with a finely reticulate and those
transitions muri biconvex in with a coarsely reticulate tectum, although occur. The are cross-section, exceptionally conical. The pollen grains are invariably tricolporate. The apertures in the ektexine (ektoapertures) are meridionally elongated slits (colpi); their relative length is indicated by the polar index (PI = width polar field/E) which varies between and with short the width 0.23 (long colpi) 0.75 (short colpi). In grains colpi, of difficult in the is the polar field may be to measure and such cases length of the colpi also indicated. The short colpi in coarsely reticulate grains may be rather inconspicuous. The endexinous wide apertures are generally oval-subrectangular, as as, or slightly sides. wider than colpi and sometimes indistinctly costate on polar
III. SYSTEMATIC DESCRIPTIONS
A. HYDNOCARPUS GAERTN.
Section Taraktogenos (Hassk.) Warb.
annamensis — I: Hydnocarpus (Gagnep.) Lescot & Sleum. Pl. 1—5.
Material studied. Indo-China: Balansa 2400.
Pollen grains suboblate to subprolate, P/E 0.83 (0.96) 1.20, equatorial outline circular,
Size: P E Ektoapertures 1 wide at tricolporate. 27 (28.7) 30//, 24 (29.8) 35 fx. fx equator, sides parallel, tips pointed, PI 0.25 (0.33) 0.42. Endoapertures distinct, oval, 7 p. long, Total wall thickness Endexine 5 [x wide, equatorially elongated. 3 fx throughout. 1 fx thick. columellae rather Supporting reticulately arranged, 0.5 fx in diameter, variable, J. Schaeffer: Pollen morphology of Hydnocarpus 67
circular to oval in free columellae size cross-section, I fx high; o—3 per lumen, variable,
to high. Tectum coarsely to finely reticulate, lumina isodiametric to slightly up 0.5 fx finer elongated, 2—3 fx in diameter, on poles, muri straight to curved, 0.5 fx wide, I /x high.
Hydnocarpus calophylla (Ridl.) Sleum. — Pl. I: 6—9; Pl. VI: I—2.
Material studied. Borneo: Jacobs 5181.
Pollen grains spherical, P/E 0.89 (0.93) 0.97, equatorial outline circular, tricolporate.
wide at sides Size: P 33 (35.9) 39 fx, E 35 (38.6) 41 fx. Ektoapertures 3.5—5 fx equator,
parallel at equator, tapering to pointed tips, PI 0.42 (0.47) 0.51. Endoapertures distinct, direction. Total wall rectangular-oval, 18 fx long, 7 fx wide, elongated in equatorial thickness Endexine thick. columellae 4.5 /X throughout. 2—4 /x Supporting reticulately
arranged, —2 in diameter, rather variable, subangular-circular to -oval in 1.5 fx cross-
free columellae to section, 1.5 [X high; 2—8 per lumen, size variable, up 0.5 fx high.
Tectum coarsely reticulate, lumina irregularly shaped, isodiametric to slightly elongated,
2 in diameter, uniformly distributed, muri straight to curved, I wide, 1 high. —5 fi fx fx
Hydnocarpus hainanensis (Merr.) Sleum.
Material studied. Hainan: F. C. How 70706.
Pollen grains spherical, P/E 0.89 (0.94) 1.04, equatorial outline circular, tricolporate.
Size: P E 1 wide at sides 23 (25) 27 /x, 25 (26.5) 28/x. Ektoapertures fi equator, parallel,
pointed, PI (0.28) Endoapertures indistinct, oval to rectangular, 6 tips 0.19 0.31. ± fx direction. Total wall thickness long, ± 1.5 fx wide, elongated in equatorial 2.5—3 fx
throughout. Endexine 1 thick. columellae reticulately arranged, fx Supporting 0.5 fi in
diameter, rather variable, subangular-circular to -oval in cross-section, —I high; 0.5 fx
free columellae o—2 size to rather per lumen, variable, up 0.5 fx high. Tectum finely reticulate, lumina irregularly shaped, isodiametric to slightly elongated, I in diam- —3 fi
muri to wide. eter, uniformly distributed, straight curved, I fx high, ± 0.5 fx
Bl. — Pl. Pl. Hydnocarpus heterophylla I: 10—13; VI: 3—4.
PNH Material studied. BS — Unesco Philippines: 41600, 23029 (ssp. philippinensis Sleum.). Java: 43, Kostermans c.s. 400 (ssp. heterophy lla).
Pollen suboblate grains to subprolate, P/E 0.78 (0.97) 1.23, equatorial outline circular,
Size: 22 1 wide at tricolporate. P 21 (28.5) 38 fx, E (29.6) 38 fx. Ektoapertures —1.5 fx sides indistinct equator, parallel, tips pointed, PI 0.20 (0.28) 0.40. Endoapertures to rather wide. Total wall thickness distinct, oval, ± <5—10 fx long, ± 3—7 fx 2.5—3 fx throughout.
Endexine I thick. columellae in diam- ± fx Supporting reticulately arranged, 0.5—1.5 /x rather -oval eter, variable, subangular-circular to in cross-section, 1—1.5 fi high; free columellae 0 variable, to to —5 per lumen, size up 0.5 fi high. Tectum finely coarsely reticulate, lumina irregularly shaped, isodiametric to slightly elongated, in diam- 1—4 fx eter, uniformly distributed, muri straight to curved, 0.5—I fx wide, 0.5—1 [X high.
ilicifolia — Hydnocarpus King. Pl. II: 1—8.
Material studied. Laos: Kerr 21293.
Pollen outline grains spherical, P/E 0.93 (1.00) 1.10, equatorial circular, tricolporate. BLUMEA 68 VOL. XX, No. i, 1972
Size: P E 1—2 wide at sides 27 (30.1) 33 fi, 29 (30) 31 fi. Ektoapertures fi equator, parallel,
pointed, PI 0.21 (0.27) Endoapertures indistinct, oval, long, wide. tips 0.33. ±7 fi ± 3 fi Total wall thickness Endexine thick. columellae 3.5 fi throughout. 1.5 fi Supporting reticulately in diameter, rather variable, subangular-circular to -oval in arranged, 0.5 fi
free columellae to cross-section, 1 fi high; 0—2 per lumen, size variable, up 0.5 fi high.
Tectum coarsely reticulate, luminairregularly shaped, isodiametric to slightly elongated, variable in shape, in finer on poles, muri straight to curved, 1 1.5—2.5 /i diameter, fi wide, 1 fi high.
Hydnocarpus kunstleri (King) Warb.
Material studied. Borneo: — Sumatra: Forbes Jacobs 5246. 3033.
Pollen grains spherical, P/E 0.85 (0.99) 1.08, equatorial outline circular, tricolporate.
Size: P 22 (26.1) E 22 (25.9) Ektoapertures wide at sides 30fi, 30 fi. 0.5—1.5 fi equator, elon- parallel, tips pointed, PI 0.23 (0.35) 0.41. Endoapertures oval, 9 fi long, 4 fi wide, gated in equatorial direction. Total wall thickness Endexine 3 fi throughout. 1—1.5 fi thick. columellae reticulately arranged, in diameter, rather variable, Supporting 0.5 fi
to -oval free columellae subangular-circular in cross-section, 0.5 —1 fi high; o—2 per
to Tectum coarsely reticulate, lumina irregularly shaped, isodia- lumen, up 0.5 fi high. metric to slightly elongated, 2—2.5 fi in diameter, uniformly distributed, muri straight to curved, 1 fi wide, 1 fi high.
Hydnocarpus kurzii (King) Warb.
Material studied. Burma: Gillet — Thailand: Kerr (cult. Hawaii) 1943. 15038.
Pollen grains spherical to subprolate, P/E 0.87 (1.02) 1.21, equatorial outline circular, wide tricolporate. Size: P 25 (26.9) 29 fi, E 24 (26.7) 28 fi. Ektoapertures 1.5—2.5 fi at sides equator, parallel, tips pointed, PI 0.22 (0.30) 0.37. Endoapertures oval, 10 fi long,
— in direction. Total wall thickness 3 4 fx wide, elongated equatorial 3 fx throughout.
Endexine I thick. columellae reticulately arranged, —I in diameter, fx Supporting 0.5 fi rather to -oval free columellae variable, subangular-circular in cross-section, 1 fi high; o—i to Tectum reticulate, lumina per lumen, up 0.5 fx high. coarsely irregularly shaped to slightly elongated, in diameter, finer poles, muri straight to curved, I 2—3 fx on fx wide, i fx high.
Hydnocarpus tenuipetala Sleum.
Material studied. Borneo: S 25371.
Pollen outline grains spherical, P/E 0.92 (0.94) 1.00, equatorial circular, tricolporate.
Size: 22 (23.1) (24.5) Ektoapertures 1 wide at sides P 24 fi, E 24 25 fi. —1.5 fi equator, indistinct. Total wall thickness parallel, tips pointed, PI 0.32 (0.36) 0.39. Endoapertures Endexine thick. columellae 3 fi throughout. 1 fi Supporting reticulately arranged, 0.5 —1 fi -oval free in diameter, rather variable, subangular-circular to in cross-section, 1 fi high; columellae lumina o—1 per lumen, up to 0.5 fi high. Tectum finely reticulate, irregularly
isodiametric to finer muri shaped, slightly elongated, 1—2 fi in diameter, on poles, straight to curved, 0.5—x fi wide, I fi high. J. Schaeffer: Pollen morphology of Hydnocarpus 69
Hydnocarpus wrayi King.
Material studied. Perak: Wray 2608.
Pollen grains spherical, P/E 0.89 (0.98) 1.08, equatorial outline circular, tricolporate. sides Size: P (24.3) x E (25.3) 26 Ektoapertures 1 wide at parallel, 23 27 / , 25 /x. fx equator, rather tips pointed, PI 0.19 (0.24) 0.30. Endoapertures indistinct, oval, 7 fx long, 2 fx wide, Total wall thickness Endexine thick. equatorially elongated. 3 fx throughout. 1 /x Sup- columellae reticulately arranged, in diameter, rather variable, subangular- porting 1 [X
circular in I high; free columellae 2 lumen, size variable, to cross-section, fx —5 per up
lumina isodiametric to slightly 1 fx high. Tectum coarsely reticulate, irregularly shaped,
in uniformly distributed, muri straight to curved, I elongated, 1.5—3.5 fx diameter, fx
wide, 1 fx high.
Section Hydnocarpus
Subsection OLIGANTHERA Warb.
Pl. Pl. Hydnocarpus alcalae C. DC. — II: 9 —11; VI: 5 —6,
Material studied. Philippines: PNH 5870.
outline Pollen grains spherical, P/E 0.92 (0.96) 1.04, equatorial circular, tricolporate.
8 Size: P 27 (30.5) 33 fx, E 30 (31.4) 33 fx. Ektoapertures (10) 12 fx long, PI ± 0.62,1—2.5 fx
wide at equator, sides parallel, tips pointed. Endoapertures indistinct, circular, 3 —4 fx diameter. Total wall thickness Endexine thick. in 3.5 /x throughout. 1—1.5 fx Supporting columellae diameter, rather subangular- reticulately arranged, 0.5—1 fx in variable,
circular in cross-section, 1 free columellae —20 size fx high; 5 per lumen, variable, up to
lumina isodiametric to 0.5 fx high. Tectum coarsely reticulate, irregularly shaped, slightly in muri elongated, 2 —5 fx diameter, uniformly distributed, straight to curved, I [X wide,
1 high. fx
Hydnocarpus alpina Wight.
Material studied. Madras: Beddome 1873.
Pollen grains spherical, P/E 1.03 (1.11) 1.24, equatorial outline circular, tricolporate.
wide at sides Size: P 29 (31.8) 36 fx, E 27 (28.6) 31 [X. Ektoapertures 1 fx equator, parallel,
PI 0.21 Endoapertures in equatorial tips pointed, (0.31) 0.39. oval, 9 by 5 //, elongated direction. Total wall thickness Endexine thick. columellae indis- 3 fx. 1 fx Supporting
—I in rather tinctly reticulately arranged, 0.5 [X diameter, variable, subangular-circular free columellae absent. lumina in cross-section, 1—1.5 fx high; Tectum finely reticulate,
isodiametric to irregularly shaped, elongated, I—1.5 fx in diameter, uniformly distributed,
muri to curved, I —1 straight fx wide, 0.5 fx high.
Hydnocarpus anthelmintica Pierre.
Material studied. Philippines (cult.): PNH 36211.
Pollen grains spherical, P/E 0.86 (0.94) 1.04, equatorial outline circular, tricolporate. wide Size: P 27 (29.5) 31 fx, E 30 (31.4) 33 fx. Ektoapertures 12 (14.2) 16 fx long, 1—2 fx
at sides PI equator, parallel, tips pointed, ± 0.65. Endoapertures distinct, oval, 4 fx long, direction. Total wall thickness 2 fx wide, elongated in equatorial 3.5 fx throughout. BLUMEA VOL. XX, No. I, 1972 70
Endexine thick. columellae i.j—2 fx Supporting indistinctly reticulately arranged, 0.5—I
in diameter, rather variable, subangular-circular in cross-section; free columellae fx absent. Tectum finely reticulate, lumina irregularly shaped, isodiametric to elongated,
in diameter, distributed, muri straight to curved, 2 wide, 1 high. ± 0.5 fx uniformly 0.5 / fx
Hydnocarpus beccariana Sleum.
Material studied. Borneo: S 25381.
outline Pollen grains spherical, P/E 0.92 (1.03) 1.X5, equatorial circular, tricolporate.
P wide sides Size: 40 (42) 45 fx, E 38 (40.6) 44 fx. Ektoapertures 1—2 fx at equator, parallel, tips pointed, PI 0.27 (0.34) 0.47. Endoapertures irregularly shaped, isodiametric to oval, direction. Total wall thickness — in elongated in equatorial 4 8 fx diameter, 2.5—3 fx Endexine thick. columellae throughout. 1 fx Supporting indistinctly reticulately arranged,
rather to -oval I 1 [X in diameter, variable, subangular-circular in cross-section, —1.5 fx high; free columellae absent. Tectum finely reticulate, lumina irregularly shaped, iso- diametric to to elongated, 0.5—I fx in diameter, uniformly distributed, muri straight curved, 1 fx wide, 1 fx, high.
Hydnocarpus borneensis Sleum.
Material studied. Borneo: SAN 36211.
Pollen outline grains spherical, P/E 0.97 (1.07) 1.27, equatorial circular, tricolporate.
Size: E (36.4) wide at sides parallel, P 35 (38.9) 45 fx, 34 39 fx. Ektoapertures 1 fx equator, rather tips pointed, PI 0.31 (0.34) 0.39. Endoapertures distinct, oval, 8—11 fx long,
— 6 direction. Total wall thickness 5 fx wide, elongated in equatorial 3—3.5 fx throughout.
Endexine thick. columellae 1 in 1—1.5 fx Supporting reticulately arranged, fx diameter, rather -oval free columellae variable, subangular-circular to in cross-section, 1 fx high; absent. Tectum finely reticulate, lumina irregularly shaped, isodiametric to elongated,
1—2 in distributed, muri to 1 wide, 1 high. fx diameter, uniformly straight curved, fx fx
Hydnocarpus castanea Hook. f. & Thorns. — Pl. II: 12; Pl. III: 1—4; Pl. VII: 1—2.
Material studied. Thailand: Kerr 18342.
Pollen grains spherical, P/E 0.93 (0.98) 1.07, equatorial outline circular, tricolporate.
Size: P (32.8) E (33.4) Ektoapertures I wide at sides 31 34 fx, 32 34 fx. fx equator, parallel, tips pointed, PI0.28 (0.36) 0.48. Endoapertures distinct, oval, 7 fx long, 5 fx wide, elongated in direction. Total wall thickness Endexine thick. equatorial 3 fx throughout. 1 fx Support-
columellae reticulately —1 in diameter, rather variable, subangular- ing arranged, 0.5 fx circular to -oval free columellae absent. in cross-section, 1 fx high; Tectum finely retic-
lumina isodiametric to 1—2 ulate, irregularly shaped, elongated, uniformly distributed, fx in diameter, muri straight to curved, 1 wide, 1 high. fx fx
Hydnocarpus cucurbitina King.
Material studied. Perak: Scortechini 2056.
Pollen grains spherical to subprolate, P/E i.oo (1.14) 1.26, equatorial outline circular, tricolporate. Size: P 22 (26.3) 22 (22.5) Ektoapertures 1 wide at 29 fx, E 23 fx. fx equator, sides parallel, tips pointed, PI 0.27 (0.29) 0.30. Endoapertures oval, 6 by 1.5 fx, elongated in direction. Total wall thickness Endexine thick. equatorial 2 /x throughout. 1 fx Support- Schaeffer: Pollen J. morphology of Hydnocarpus 71
columellae in reticulate 0.5 in diameter, rather ing very indistinctly arranged a pattern, ft
to circular cross-section, high; free columellae absent. variable, subangular in 0.5 ft Tectum finely reticulate, lumina irregularly shaped, isodiametric to elongated, 1 fi in
to curved, wide, 1 high. diameter, uniformly distributed, muri straight 0.5 fi fi
Hydnocarpus curtisii King.
Material studied. Penang: Curtis 1535.
outline circular, Pollen grains spherical, P/E 0.94 (1.06) 1.17, equatorial tricolporate.
wide at sides parallel, Size: P 30 (31.8) 35 ft, E 28 (30.1) 32 ft. Ektoapertures 0.5 fi equator, rather indistinct, oval, long, tips pointed, PI 0.21 (0.26) 0.30. Endoapertures 6 fi 2.5 /i Endexine wide, elongated in equatorial direction. Total wall thickness 3 fi throughout. rather thick. columellae reticulately arranged, —1 in diameter, x n Supporting 0.5 ft circular cross-section, free columellae absent. variable, subangular to in 1 p. high;
Tectum finely reticulate, lumina irregularly shaped, isodiametric to elongated, 1—1.5 fi
—I wide, I high. in diameter, uniformly distributed, muri straight to curved, 0.5 fi fi
Pl. Hydnocarpus elmeri Merr. — Pl III: 5 —8; VII: 3—4.
Material studied. Borneo: Elmer 20736.
outline circular, tricolporate. Pollen grains spherical, P/E 1.00 (1.12) 1.21, equatorial wide sides Size: P 17 (17.7) 19 fi, E 14 (16.9) 17 ft- Ektoapertures I fi at equator, parallel,
indistinct. Total wall thickness 2 tips pointed, PI 0.27 (0.32) 0.38. Endoapertures very fi thick. columellae reticulately arranged, in throughout. Endexine 1 fi Supporting 0.5 fi
rather variable, subangular to oval in cross-section; free columellae diameter, 0.5 fi high,
absent. Tectum finely reticulate, lumina irregularly shaped, isodiametric to elongated,
finer muri straight to curved, wide, 0.5 fi in diameter, slighdy near ektoapertures, 0.5 fi surface conical in cross-section. 0.5 fi high, upper
Hydnocarpus glaucescens Bl.
Material studied. Sumatra, Padang: Beccari 948.
oudine Pollen grains spherical, P/E 0.96 (1.05) 1.14, equatorial circular, tricolporate. sides 22 Ektoapertures 1 wide at parallel, tips pointed, PI Size: P (24) 25 fi. fi equator, indistinct. Total wall thickness Endexine 0.32 (0.36) 0.40. Endoapertures very 3 [i . 1.5 fi
I in rather variable, thick. Supporting columellae reticulately arranged, fi diameter, in oval free columellae 1 lumen, subangular to in cross-section, 1 fi high; —4 per 0.5 fi
Tectum finely to coarsely reticulate, lumina 1.5 in diameter, diameter, 0.5 [i high. —3 fi uniformly distributed, irregularly shaped, isodiametric to elongated, muri straight to
curved, 1 ft wide, 1 fi high.
Hydnocarpus nana King.
Material studied. Perak: King*s collector 3828.
outline circular, Pollen grains spherical, P/E 0.81 (0.88) 1.00, equatorial tricolporate. wide sides (17.9) E (20.3) 21 Ektoapertures 0.5 at equator, parallel, Size: P 17 19 /i, 19 fx. [X Endoapertures rather indistinct, oval, ± by 1 tips pointed, PI 0.45 (0.48) 0.50. 4 [X,
direction. Total wall thickness 2 throughout. Endexine —1 elongated in equatorial fx 0.5 fx BLUMEA VOL. No. 72 XX, i, 1972
thick. columellae rather Supporting reticulately arranged, 0.5 fi in diameter, variable,
circular to oval in free columellae absent. cross-section, 0.5 fi high; Tectum finely retic-
lumina irregularly shaped, isodiametric to elongated, 1 in ulate, —1.5 fi diameter, uniformly distributed, muri straight to curved, wide, high. 0.5 [i 0.5 fi
Sloot. — Pl. — Hydnocarpus pentagyna III: 9 13
Material studied. Borneo: SAN 15170, S 22116.
Pollen outline grains spherical to prolate, P/E 0.90 (1.10) 1.36, equatorial circular,
wide at tricolporate. Size: P28 (33.9) 41 /t, £28(30.7) 35 fi.Ektoapertures 0.5—1 fi equator,
sides PI 10 parallel, tips pointed, ± 0.23. Endoapertures distinct, oval, ± )jl long, ± 5 wide, elongated in equatorial direction. Total wall thickness throughout. Endexine 3 fi thick. columellae I fi Supporting indistinctly reticulately arranged, O.J fi in diameter,
subangular-circular to -oval in cross-section, 1 high; free columellae absent. —1.5 fi
lumina isodiametric to Tectum finely reticulate, irregularly shaped, elongated, 0.5—1 (jl muri in diameter, uniformly distributed, straight to curved, 0.5—1 fi wide, 1 fjL high.
Hydnocarpus pentandra (Buch.-Ham.) Oken; H. laurifolia (Dennst.) Sleum.
Material studied. India: Hook. f. & Thorns, s.n.
Pollen outline grains spherical, P/E 0.91 (1.07) 1.23, equatorial circular, tricolporate.
Size: 1 wide at sides P 21 (24) 27 fi, E 20 (22.3) 24 fi. Ektoapertures —2 fi equator, parallel,
pointed, PI (0.33) Endoapertures oval, 10 long, wide, elongated tips 0.27 0.39. fi 3—4 fi direction. Total wall thickness Endexine thick. in equatorial 3 fi throughout. 1 fi Sup- columellae indistinctly reticulately arranged, in diameter, rather variable, porting 0.5 fi
to -oval in cross-section, I high; free columellae absent. Tectum subangular-circular fi lumina isodiametric finely reticulate, irregularly shaped, to elongated, 1—2 fi in diameter, muri uniformly distributed, straight to curved, 1 fi wide, 1 fi high.
Hydnocarpus subfalcata Merr.
Material studied. Borneo: S 18087.
Pollen grains spherical, P/E 0.93 (0.98) 1.07, equatorial outline circular, tricolporate. sides Size: P 25 (28.5) 32fi, E 26 (29.2) 32 fi. Ektoapertures 1—2 fi wide at equator, parallel, indistinct tips pointed, PI 0.46 (0.50) 0.56. Endoapertures or distinct, 3 fi wide, length variable, equatorially elongated and sometimes nearly merging with each other. Total thickened wall thickness 4—4.5 fi. Endexine 1.5—2 fi thick, occasionally somewhat up to around the columellae 2.5 fi endoapertures. Supporting reticulately arranged, 0.5 —1 fi in
diameter, rather variable, subangular-circular to -oval in cross-section; free columellae lumina 1—6 per lumen, size variable, up to 0.5 fi high. Tectum coarsely reticulate, irregu- larly shaped, isodiametric, 2—4 fi in diameter, uniformly distributed, muri straight to curved, 1 fi wide, 1 fi high.
Hydnocarpus sumatrana (Miq.) Koord.
Material studied. Philippines: FB 3916.
Pollen grains spherical, P/E 1.11 (1.15) 1.21, equatorial outline circular, tricolporate.
Size: wide at sides P 35 (35.3) 37 fx, E 29 (31.7) 35 fx. Ektoapertures 1 fx equator, parallel, indistinct. Total wall thickness tips pointed. PI 0.35 (0.37) 0.39. Endoapertures 3 fx. J. Schaeffer: Pollen morphology of Hydnocarpus 73
Endexine thick. columellae reticulately i in diameter, rather i [X Supporting arranged, fx free columellae absent. Tectum variable, subangular-circular in cross-section, 1.5 [X high; diam- finely reticulate, lumina irregularly shaped, isodiametric to elongated, 0.5 —I fi in
muri straight to curved, I wide, high. eter, uniformly distributed, fx 1.5 fx
Hydnocarpus venenata Gaertn.
Material studied. Ceylon: Thwaites 1630.
outline tricolporate. Pollen grains spherical, P/E 0.97 (1.00) 1.04, equatorial circular, wide sides Size: P 27 (28.1) 29 fx, E 27 (28.2) 29 fx. Ektoapertures 0.5—x fx at equator,
8 parallel, tips pointed, PI 0.21 (0.26) 0.32. Endoapertures distinct, oval, fx long, 3 fx wide, direction. Total wall thickness throughout. Endexine elongated in equatorial 4fx 1.5 /x rather thick. Supporting columellae reticulately arranged, 0.5—1 [X in diameter, variable,
free columellae o—2 lumen, subangular-circular to -oval in cross-section, 1 fx high; per lumina irregularly isodiametric to up to 0.5 fx high. Tectum coarsely reticulate, shaped,
in finer on muri straight to curved, 1 wide, slightly elongated, 2—4 fx diameter, poles, fx
1 fx high.
Hydnocarpus yatesii Merr.
Material studied. Sumatra: R. Si Toroes 4644.
outline Pollen grains spherical, P/E 0.88 (0.96) 1.05, equatorial circular, tricolporate. sides Ektoapertures —1 wide at Size: P 21 (21.6) 22 fx, E 21 (22.5) 24 fx. 0.5 fx equator, long, wide, parallel, tips pointed, PI 0.18 (0.23) 0.27. Endoapertures oval, 7 fx 2—2.5 Total wall thickness Endexine elongated in equatorial direction. 5 fx throughout. I—1.5 fx columellae in rather variable, thick. Supporting reticulately arranged, 0.5—1 fx diameter,
to -oval in I free columellae I lumen, subangular-circular cross-section, fx high; —4 per Tectum coarsely reticulate, lumina irregularly shaped, size variable, up to 0.5 fx high. isodiametric uniformly distributed, muri to slightly elongated, 2—3 fi in diameter,
to —x high. straight curved, 0.5 fx wide, 0.5 fx
Subsection PLEIANTHERA Warb.
Hydnocarpus crassifolia Sleurn.
Material studied. Borneo: Beccari 3972.
Pollen grains spherical, P/E 0.91 (0.99) 1.10, equatorial outline circular, tricolporate.
Size: P (35.2) 38 E (35.4) 38 Ektoapertures 8—10 long, PI ± —1 32 [JL, 32 fx. fx 0.75, 0.5 [x indistinct. Total wall thickness wide at equator, sides parallel, tips pointed. Endoapertures
Endexine thick. columellae reticulately arranged, diameter 5 —6 fx, 2.5—3 fx Supporting
rather variable, subangular-circular or -oval in cross-section, 1 high; i.j—2.5fx, —1.5 fx
to reticulate, free columellae 4—14 per lumen, size variable, up I fx high. Tectum coarsely lumina irregularly shaped, isodiametric to slightly elongated, 2.5—3.5 fx in diameter, uniformly distributed, muri straight to curved, 1.5 [X wide, 1.5 fx high.
Hydnocarpus octandra Thwait.
Material studied. Ceylon: Thwaites 2640.
outline Pollen grains spherical, P/E 0.93 (0.99) 1.14, equatorial circular, tricolporate.
28 wide sides Size: P 28 (29.4) 32 fx, E (29.7) 32 fx. Ektoapertures 1—2 fx at equator, BLUMEA VOL. XX, No. i, 1972 74
oval.Total wall thickness parallel, tips pointed, PI 0.35—0.40. Endoapertures indistinct, 4fx. Endexine thick. columellaereticulately arranged, in diameter, 1.5 fx Supporting x—1.5 fi free columellae rather variable, subangular-circular to -oval in cross-section, 1 fx high;
size variable, to high. Tectum coarsely reticulate, lumina 3—12 per lumen, up 0.5 fx irregularly shaped, isodiametric to slightly elongated, 3 —5 fx in diameter, uniformly distributed, muri straight to curved, I fx wide, 1—1.5 fx high.
Hydnocarpus woodii Merr. — Pl. IV: 1—6,
Material studied. Borneo: SAN 20839.
outline Pollen grains spherical, P/E 0.86 (1.03) 1.14, equatorial circular, tricolporate. sides 28 (28.4) Ektoapertures —1 wide at Size: P (29.2) 31 /x, E 24 31 [X. 0.5 fx equator, parallel, tips pointed, PI 0.28 (0.34) 0.40. Endoapertures rather indistinct, 6 fx long, 3.5 /z Total wall thickness wide, elongated in equatorial direction. 3.5—4./u throughout.
Endexine thick. columellae arranged, ± 1 in diameter, 1.5 /x Supporting reticulately fx free columellae rather variable, subangular-circular or -oval in cross-section, 1 fx high;
to high. Tectum coarsely reticulate, lumina irregularly shaped, i—7 per lumen, up 0.5 fx isodiametric in distributed, muri to slightly elongated, 2—4 fx diameter, uniformly straight to curved, 1—I.J fx wide, 1 fx high.
Section Asteriastigma (Bedd.) Warb.
Pl. anomala Sleum. — Pl. IV: VII: —6. Hydnocarpus (Merr.) 7 —13; 5
Material studied. Borneo: SAN 35247.
outline Pollen grains suboblate to spherical, P/E 0.83 (1.00) 1.02, equatorial circular,
28 E Ektoapertures 11 (13.7) is long, tricolporate. Size: P (32) 37 fx, 30 (32.1) 37fi. fx
wide at PI 0.60—0.65, Endoapertures distinct, oval, 2 —4 fx equator, ± tips pointed. direction. Total wall thickness — 10 —6 wide, in equatorial y fx long, 5 fx elongated 3—5 fx Endexine thick. columellae reticulately arranged, throughout. 1—1.5 fx Supporting
in to -oval in cross-section; free columellae 1—13 i—1.5fi diameter, subangular-circular
to Tectum lumina irregularly per lumen, size variable, up 0.5 fx high. coarsely reticulate, distributed, muri shaped, isodiametric to elongated, 2—6 fx in diameter, uniformly
to curved, I high. straight [X
Hydnocarpus gracilis (Sloot.) Sleum.
Material studied. Sumatra: Krukojf 4085.
outline tricolporate. Pollen grains spherical, P/E 0.94 (1.04) 1.07, equatorial circular, sides Size: E (29.7) Ektoapertures 1 wide at parallel, tips P 29 (31) 32 fx, 28 31 fx. fx equator, pointed, PI 0.40 (0.48) 0.53. Endoapertures distinct, oval, 10 fx long, 5 fx wide, elongated Total wall thickness Endexine thick in equatorial direction. 3.5—5 fx. 1.5 fx thick, 2.5 fi around the endoapertures. Supporting columellae reticulately arranged, 1—2 /x in -oval free diameter, rather variable, subangular-circular to in cross-section, 1 fi high;
size Tectum reticulate, columellae 2 —7 per lumen, variable, up to 0.5 fx high. coarsely lumina irregularly shaped, isodiametric to slightly elongated, 2 —7 fx in diameter, uni-
to curved, x high, wide. formly distributed, muri straight fx 1.5 fx J. Schaeffer: Pollen morphology of Hydnocarpus 75
Hydnocarpus macrocarpa (Bedd.) Warb. ssp. burmanica Sleum.
Material studied. Burma: Parkinson 402.
Pollen outline grains spherical, P/E 0.93 (1.01) 1.12, equatorial circular, tricolporate.
Size: P 27 (28.1) 30 fx, E 26 (27.9) 30 fx. Ektoapertures 12 (13) 14 /x long, PI 0.61—0.67,
wide at xo 2.5—3 fi equator, tips pointed. Endoapertures distinct, oval, by 4—5 fx, elongated in equatorial direction. Total wall thickness throughout. Endexine 4 fx 1.5 fi thick. columellae rather Supporting reticulately arranged, 0.5—I fx in diameter, variable, subangular-circular in high; free columellae size cross-section, 1 fx 2 —7 per lumen,
to lumina isodia- variable, up 0.5 fx high. Tectum coarsely reticulate, irregularly shaped,
metric to elongated, 2 —4 fx in diameter, uniformly distributed, muri straight to curved,
—I wide, high. 0.5 (x 1.5 fx
Hydnocarpus polypetala (Sloot.) Sleum.
Material studied. Borneo: Jacobs 5313, Kostermans 21392.
Pollen grains spherical to subprolate, P/E 0.88 (1.05) 1.26, equatorial outline circular,
Size: wide at tricolporate. P 28 (34.5) 41 /x, E 26 (32.9) 41 fx. Ektoapertures 1—3 fx equator,
sides parallel, pointed, PI (0.38) Endoapertures distinct, oval, 10 tips 0.27 0.47. —14 fx direction.Totalwall thickness long, 5 —7fx wide, elongated in equatorial 4—6fx through-
out. Endexine thick. columellae reticulately arranged, in I—2 fx Supporting I—1.5 fx rather -oval diameter, variable, subangular-circular to in cross-section, 1—2 fx high;
free columellae size to Tectum o—5 per lumen, variable, up 1 fx high. coarsely reticulate,
lumina irregularly shaped, isodiametric to slightly elongated, 2 in diameter, uni- —5 fx
to wide. formly distributed, muri straight curved, 1—1.5 fx
Section Kerrandrias Sleum.
Craib. — Pl. Hydnocarpus calvipetala V: 1—3.
Material studied. Thailand: Kerr 16518.
Pollen outline grains spherical, P/E 0.92 (1.03) 1.17, equatorial circular, tricolporate. Size: E wide sides P 27 (31.2) 33 fx, 27 (30.1) 32 fx. Ektoapertures 1.5—3 fx at equator,
parallel, tips pointed, PI 0.20 (0.27) 0.37. Endoapertures distinct, oval, 8 fx long, 5 fx wide, in direction. Total wall thickness Endexine elongated equatorial 3.5 fx throughout. 1—1.5
thick. columellae reticulately arranged, —1 in diameter, rather fx Supporting 0.5 fx
to -oval in cross-section, free columellae 0 variable, subangular-circular 1 fx high; —5 per lumen, scattered, size variable, to high. Tectum coarsely reticulate, lumina up 0.5 fx
isodiametric to irregularly shaped, slightly elongated, 3 —5 fx in diameter, on poles 2 —3 fx in muri diameter, straight to curved, I fx wide, I fx high.
B. CHLOROCARPA ALSTON
Alston. — Pl. Chlorocarpa pentaschista V: 4—9.
Material studied. Ceylon: Thwaites 2918 (BM).
Pollen outline grains subprolate, P/E 1.11 (1.18) 1.24, equatorial circular, tricolporate.
Size: wide at sides P 38 (39.6) 41 fx, E 32 (33.6) 36 /x. Ektoapertures 1 fx equator, parallel, 76 BLUMEA VOL. XX, No. I, 1972
indistinct. Total wall thickness Ende- tips pointed, PI 0.22 (0.27) 0.31. Endoapertures 2 fx.
xine —1 thick. columellae indistinctly reticulately arranged, —1 0.5 fx Supporting 0.5 /x
in rather roimd to oval in 1 free columellae diameter, variable, cross-section, fx high;
absent. Tectum finely reticulate, lumina irregularly shaped, isodiametric to elongated,
in 0.5 —1 [X diameter, uniformly distributed, muri straight to curved, 0.5 fx wide, 1 fx high.
C. NEOPTYCHOCARPUS BUCHHEIM
Neoptychocarpus apodanthus (Kuhlmann) Buchheim.
Material studied. Brasil, Amazonas: Silva 57815.
Pollen outline grains spherical, P/E 0.88 (i.oo) 1.07, equatorial circular, tricolporate.
28 wide sides Size: P 22 (26.4) 30 fx, E 25 (26.4) fx. Ektoapertures 0.5 fx at equator, parallel, PI tips pointed, 0.36—0.52. Endoapertures distinct, irregularly isodiametric to oval,
long, wide. Total wall thickness 2 around the 2.5—7fx 3—5 fx fx, endoapertures 5 fx. Endexine thickened sides of the width of these ± 0.5 fx, mainly on polar endoapertures,
to columellae in thickenings up 2.5 fx. Supporting single rows, reticulately arranged, 0.5 fx rather circular oval free columellae in diameter, variable, to in cross-section, ± 1 fx high;
to lumina o—i per lumen, size variable, up 0.5 fx high. Tectum coarsely reticulate, irreg-
isodiametric to in smaller the ularly shaped, slightly elongated, 1.5—3 fx diameter, near
muri straight to angular, —1 wide, high. apertures, 0.5 fx 0.5 fx
IV. POLLEN TYPES
From the preceding systematic descriptions it will be clear that the pollen of the Hydno-
is of the same general and that all other characters carpus species type vary continuously between the species.
Based on a combination of characters, the following pollen types can nevertheless be recognized:
free columellae columellae often Type I: Lumina5S 2 fx in diameter, absent, supporting irregularly arranged, ektoapertures medium long.
free columellae columellae Type II: Lumina 2 [X, present or absent, supporting in a of variable. reticulate pattern, length ektoapertures
Subtype Ha: 0—6 free columellae/lumen, lumina often finer on poles, ekto-
apertures long.
Subtype lib: 0—20 free columellae/lumen, lumen size uniform, ektoapertures short, often indistinct.
If assumed that the reticulate which a it is small, finely pollen type occurs in majority of Flacourtiaceous has been the ancestral of genera, type, a phylogenetic sequence Hydno- in carpus pollen types as shown fig. 1 can be envisaged. J. Schaeffer: Pollen morphology of Hydnocarpus 77
lib, PI 0.49
increase free columellae
Ha, PI 0.32
development free columellae further enlargement lumina
I, PI 0.37
enlargement lumina
basic of reticulate Flacourtiaceae type finely pollen
in Fig. 1. Pollenmorphological trends Hydnocarpus.
This would indicate that the coarsely reticulate grains with short ektoapertures and abundant free columellae (pollen type lib) form a derived pollen type. Hydnocarpus the closest resemblance the basic Flacourtiaceous elmeri (pollen type I) shows to pollen
type.
V. POLLEN MORPHOLOGY AND TAXONOMY
The distribution of the the and is investigated species over types subtypes presented
in TableI. From this table it would appear that subsection Oliganthera of section Hydno-
variation carpus has preserved the most complete range of with both primitive and advanced types present. Subsection Pleianthera has only advanced pollen types. Section but Taraktogenos lacks the primitive types has developed a high percentage of Ila-type Sections and Kerrandrias show each different derived grains. Asteriastigma a type. Hut-
chinson (1967) has recently distinguished Taraktogenus Hassk. as a separate genus, while
Sleumer earlier it of (1954), following authors, only accepts as a subgenus Hydnocarpus.
Pollenmorphologically it clearly belongs to Hydnocarpus, which supports Sleumer's opinion.
Since itis also ofinterest to provide evidence for the taxonomic position of Hydnocarpus in within Flacourtiaceae, a general survey of the pollen types present this family was made.
This survey was restricted to one or two representatives of each genus, as far as material
was available, and is of a preliminary nature.
In general, pollen grains in Flacourtiaceae are fairly small, spherical to prolate and
tricolporate. Ektoapertures are distinct, slitlike, endoapertures are variously developed and often elongated in equatorial direction. In some cases the endoapertures are much
reducedand only developed as equatorial bridges in the colpate ektoapertures. Columellae
are indistinct or distinct and often reticulately arranged. BLUMEA VOL. XX, No. I, 1972 78
Kerrandrias calvipetala Hydnocarpus Asteriastigma macrocarpa polypetala anomala gracilis in subdivision Pleianthera crassifolia octandra woodii
! | ' 1 1 1 . 1 1 1 1 1 11 | j 1 1 | | i 1 taxonomic
Hydnocarpus and anthelmintica Oliganthera alcalae yatesii glaucescenssubfalcata venenata alpina beccariana borneensis castanea cucurbitina curtisii elmeri nana pentagyna pentandra sumatrana types Pollen
I. Taraktogenos calophylla annamensis hainanensis heterophylla ilicifoliakunstleri kurzii tenuipetala wrayi Table
Section b a II II Subsection I J. Schaeffeh: Pollen morphology of Hydnocarpus 79
Based the of the the be mainly on development tectum, following groups can recog- nized:
smooth or columellae A. Tectum finely reticulate, < 0.5 fi in diameter, grains generally small.
reticulate B. Tectum to coarsely reticulate, columellae > 0.5 fi in diameter, grains fairly large (Hydnocarpus type).
C. Tectum gemmate (Gynocardia type). D. Tectum areolate-perforate ( Camptostylus type).
E. Apertures porate (Lozania type).
table the distribution of these the indicated. will be In II pollen types over generais It clear that the The of the Hydnocarpus pollen type (B) is relatively infrequent. pollen genus and this the Chlorocarpa belongs to type B-I is also genus which is taxonomically nearest
similar is this to Hydnocarpus. Another genus with pollen Neoptychocarpus, but differs, as be from the this the and thinner can seen description in paper, in costate endoapertures
wall. Besides, it belongs to quite a different tribe (Casearieae). The resemblance with
pollen of Trichostephanus (Trichostephaneae) is not so striking, although it comes close to
Hydnocarpus type I, from which it differs in the broader muri. Outside distinct Flacourtiaceae there is a resemblance in sculpture with the pollen of
Paropsia, a genus of Passifloraceae which has sometimesbeen referred to Flacourtiaceae. The
aperture development in Paropsia, however, shows considerable difference and is typical for Passifloraceae (Pacque, personal communication).
In conclusion it can be stated that, in contrast to the relative pollen morphological of uniformity Flacourtiaceae, within Hydnocarpus a derived pollen type has developed,
characterized by a coarsely reticulate sculpture, free columellae,and a tendency to reduction
is certain of the ektoapertures. Within the genus this tendency to a degree correlated with
the taxonomic subdivision.
REFERENCES
G. Pollen and ERDTMAN, 1952. morphology plant taxonomy, Angiosperms. II. HUTCHINSON, J. 1967. The genera of flowering plants
SLEUMER, H. 1938. Monographic der Gattung Hydnocarpus Gaertner. Bot. Jahrb. 69: 1 —94.
H. —106. SLEUMER, 1954. Flacourtiaceae. Flora Malesiana I, 5: 1 BLUMEA VOL. XX, No. I, 1972 80
TABLE II. Distribution of pollen types (A—E) in relation to tribal subdivision (I—XI) in
Flacourtiaceae.
I Oncobeae VI Homalieae
A Rawsonia A ByrsanthusByrsanthus
A DasylepisDasykpis A Gerrardina
A Scottellia A Homalium
A Berberidopsis A CalanticaCalantica
A ErythrospermumErythrospermum A Trimeria
D Camptostylus A Dissomeria A Poggea
A Peterodendron VIIVu Phyllobotryeae
A Grandidiera no material available
A Oncoba
A Xylotheca VmVul Flacourtieae
A Caloncoba Flacourtiinae
A Lindackeria A Eichlerodendron
A Mayna A Xylosma A Carpotroche A Azara
A Buchnerodendron A Ludia
A Prockiopsis A Tisonia A Aphloia
II Pangieae A Flacourtia
Hydnocarpinae A DouyalisDovyalis B HydnocarpusHydnocarpus A Neopringlea B Chlorocarpa C Scaphocalyx Idesiinae
C Trichadenia A Bennettiodendron
Cc Gynocardia A Cathayeia ((== Idesia) A Pangium A Poliothyrsis A Ryparosa A Itoa
C Eleutherandra
C Baileoxylon IXIX CasearieaeCascaricae A Lunania
Kiggelariinae D Samyda A Kiggelaria A Tetrathylacium
A Laetia inIll AbatieaeAnatieae A Bartholomaea
A AbatiaAbalia A Euceraea
A AphaeremaAphaerema A Ophiobotrys
A Osmelia
IV TrichostephaneaeTrichostephaneae A Ryania B TrichostephanusTrichostephanus A HecatostemonHecatostemon
A Zuelania
V Scolopieae D Casearia Scolopiinae B Neoptychocarpus A Scolopia A Hemiscolopia X Bembicieae A StreptothamnusSlreptothamnus A Bembicia
A Pseudoscolopia
XI Lacistemeae
Prockiinae A Lacistema
A Prockia E Lozania
A Banara
A Pineda
A Pleuranthodendron J. Schaepfer: Pollen morphology of Hydiiocarpus 81\I
I. annamensis H. Plate Fig. 1—5: Hydnocarpus (Balansa 36211).— Fig. 6 —9: calophylla (Jacobs 5181). —
Fig. 10—13: H. heterophylla ssp. heterophylla (Kostermans 400). All x 1000.
6 82\II BLUMEA VOL. XX, No. i, 1972
II. Plate 1—8: H. — Fig. Hydnocarpus ilicifolia(Kerr 21293). Fig. 9—11: alcalae (PNH 58780). Fig. 12: H. All castanea (Kerr 18324). x 1000. J. Schaeffer: Pollen morphology of Hydnocarpus 83\III
— Plate III. — H. elmeri Fig. 1—4: Hydnocarpus castanea (Kerr 18324). Fig. 5—8: (Elmer 20736). Fig.
H. — H. All 9 —10: pentagyna (S. 22116). Fig. 11—13: pentagyna (SAN 15170). x 1000. 13LUMEA VOL. No. 84\IV XX, I, 1972
Plate IV. Fig. 1—6: Hydnocarpus woodii (SAN 20859). — Fig. 7—13: H. anomala (SAN 35247). All
X 1000. J. Schaeffer: Pollen morphology of Hydnocctrpus 85\V
V. 4 —9: (Thwaites Plate Fig. 1—3: Hydnocarpus calvipetala (Kerr 16518). Fig. Chlorocarpa pentaschista
2918). All x 1000. 86\VI BLUMEA VOL. XX, No. I, 1972
(Jacobs Plate VI. Stereoscan photomicrographs. Fig. 1, x 1000. Fig. 2, x 2000: Hydnocarpus calophylla
5181). Fig. 3, x 1000, Fig. 4, x 5000: H. heterophylla ssp. heterophylla (Kostermans 400). Fig. 5, X 1000,
H. Fig. 6, X 2500: alcalde (PNH 58780). J. Schaeffer: Pollen morphology of Hydiwcarpus 87\VII
VII. Stereoscan castanea Plate photomicrographs. Fig. 1 x 2000, Fig. 2, x 5000: Hydnocarpus (Kerr H. H. 18324). Fig. 3, x 2500, Fig. 4, x 5000: elmeri (Elmer 20736). Fig. 5, x 1000, Fig. 6, x 2500: anomala (SAN 35247). BLUMEA VOL. No. 88\I XX, I, 1972
Plate I. Scypkostegia borneensis Stapf — A. young plant of about two years old. — B. late anaphase I, = meiotic chromosomes in — C — F. mitotic chromosome showing a pollen mother cell, n 9. complements, D 2n = 18; C —D. chromosome complement of a vegetative cell of a flower-bud, with illustrating the inked inked chromosomes of C; E—F. chromosome complement of a roottip cell, with F illustrating the chromosomes of E. — (B & D —F, X 3000; C, x 1500. A & E, Nooteboom & Aban 1600; B & C, Nooteboom & Aban 1539).