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ISSN 0374-7859
THE GARDENS’ BULLETIN
SINGAPORE
Volume 37
(1984)
A periodical reflecting the interests and
activities of the Botanic Gardens
Singapore
Published by the Botanic Gardens
Parks and Recreation Department
Ministry of National Development
Singapore 1025
Printed by Jin Jin Printing Industry Pte Ltd
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CONTENTS
Volume 37
PART 1 — Ist June 1984
KENG, HSUAN:
Florae Malesianae Precursores LVIII, Part Two. The Genus
PPE PTT CACCAC) TINY HeStansese tenes eae soon a eo cew nonce isaccoech oe scuiecoeewt ous dewedetens
MABBERLEY, D. J.:
A Monograph of Melia in Asia and the Pacific: The history
A TECURSCO ATE AT Ga LCrstaniilt AG ehacseee- ae areca sas a acic enrosncsic onticninss doanidys sasbiewtaedeaeeccdsence
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FOONG, THAI WU, AND NORMAN WELLS:
Physical Effects of Soil Compaction and Initial Growth of
PCUCiMpyCnaritia (eepuminosae) inia’ Clay-lOam SOM .....0..<0c0ss-cscee0eccwcsceecsousenecesscsees
LIM, AH LAN:
The Embryology of Garcinia mangostana L. (Clusiaceae) ...............0cceceeeceeeeceeneeeeeeeenes
PUTZ, FRANCIS E., AND N. MICHELE HOLBROOK:
Multiple Branching at Nodes of S)mingtonia populnea (Hamamelidaceae) ...................--
VIANE ROL. L.:
Trichomanes proliferum Bl\.: A new record for the fern flora of Singapore .....................-
PART 2 — Ist December 1984
WILDE, W. J. J. O. DE:
A New Account of the Genus Horsfieldia (Myristicaceae), Pt 1 ...............2.cececeeeececeeeeees
LIM, AH LAN:
The Reproductive Biology of Rambutan (Nephelium lappaceum Linn., Sapindaceae) .......
WONG, KHOON MENG:
A Revision of Rennellia (Rubiaceae) in the Malay Peninsula .....................0cececeeeeeeeenees
GHANI, FARAH D.:
Keys to the Cultivars of Keladi (Colocasia esculenta — Araceae) in Peninsular
EAL AVS APPR eee ser TC eee nce soci ae <i sick oca'sccll ssasieva ya teenls «cies Seiscacdscweb Sesbuatsens
RIE W. K.:
Notes on the Systematy of Malayan Species of Chionanthus (Oleaceae) ..................0.020+
KORTHOF, H.M. & J. F. VELDKAMP:
A Revision of Aniselytron with Some New Combinations in Deyeuxia in SE. Asia
COAG) odadoh6 e5as baaeeencs soo ete ee ORR REE TEE CER R EE ere eee eee
KIEW, R.:
Preliminary Pollen|Study of the Oleaceae'in Malesia .............<.sc0.c..2s0.0.s+cecceeneseeseseeeee
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ISSN 0374-7859
THE GARDENS’ BULLETIN
1) SINGAPORE
Oy VOL. 37 (Part 1)
(, CONTENTS
‘) KENG, Hsuan:
Florae Malesianae Precursores LVIII, Part Two. The Genus
Ist June 1984
PAGES
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@ The Embryology of Garcinia mangostana L. (Clusiaceae) .............22222222005- 93-103
% PUTZ, FRANCIS E., AND N. MICHELE HOLBROOK:
Multiple Branching at Nodes of Symingtonia populnea (Hamamelidaceae) ........... 105-109
ee VIANE, R. L. L..:
Trichomanes proliferum Bl\.: A new record for the fern flora of Singapore .......... 111-114
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THE GARDENS’ BULLETIN
SINGAPORE
VOL- 37 (Part 1) Ist June 1984
CONTENTS
KENG, HSUAN:
Florae Malesianae Precursores LVIII, Part Two. The Genus
SeueIaeNFIRAEIVITCACEAC MITE IVI ACSIA petre rererc sities cee ove erage © seer Siolavnga So aceid oe usu dele eters
MABBERLEY, D. J.:
A Monograph of Melia in Asia and the Pacific: The history
qo “igiee (Cacare Pat he 20 1 Cl ES] Ome ee
LEE, S. K., AND A. N. RAO:
Pam RE GSHOM SOME sEOpiCal IRTEES) . 25. feo atei= ba lod RSS siele nsec ac Sole Saterselete ane slow
FOONG, THAI WU, AND NORMAN WELLS:
Physical Effects of Soil Compaction and Initial Growth of
Acacia pycnantha (Leguminosae) in a Clay-loam Soil ............--.....0---00+-5-
LIM, AH LAN:
The Embryology of Garcinia mangostana L. (Clusiaceae) ............--..--..-+----
PUTZ, FRANCIS E., AND N. MICHELE HOLBROOK:
Multiple Branching at Nodes of Symingtonia populnea (Hamamelidaceae) ...........
RAaNE, Ro L. L::
Trichomanes proliferum Bl.: A new record for the fern flora of Singapore ..........
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Florae Malesianae Precursores - LVIII, Part Two*
The Genus Gordonia (Theaceae) in Malesia
HsuAn KENG
Department of Botany, National University of Singapore, Singapore
EFFECTIVE-PUBLICATION DATE: 5 Sept. 1984
Abstract
This is a taxonomic treatment of 21 species of Gordonia (of the family Theaceae) found in the Malesian
region. Among these, there are two new (G. borneensis, G. sarawakensis) and two newly combined
(G. integerrima, G. vulcanica) species. Besides, two excluded (G. brevifolia, G. lobbii) and two doubtful
(lanceifolia, G. sarasini) ones are briefly mentioned. A complete list of scientific names and their
synonyms is presented in an index.
I. INTRODUCTION
The genus Gordonia was established by John Ellis in 1771 in honour of James
Gordon (1728-1791), an English nurseryman. The type species Gordonia lasianthus
(L.) Ellis (basinym: Hypericum lasianthus L.), is a tree originally from the coastal
plain areas along Loblolly Bay in eastern North America.
Gordonia Ellis is a conserved generic name against the earlier name, Lasianthus
Adanson (1763). In addition, a number of synonyms were proposed by different
authors. Some of them perpetuate to this day. The most controversial one is
Laplacea HBK (1822), which is based on a plant (Laplacea speciosa Kunth ex HBK)
from Ecuador in South America. The original Gordonia, namely G. /asianthus Ellis,
differs considerably from the type species of Laplacea, namely L. speciosa HBK.
They differ in various aspects of pedicels, bracteoles, sepals, petals, androecia and
gynoecia. These differences gradually break down when many more species were
subsequently discovered and described. Even in recent literature, several authors
(e.g. Melchior, Kobuski) maintain Laplacea as a separate genus mainly on the
ground of the usually 5 short, free styles; while others (e.g. Burkill, Merrill, Sealy)
advocate that these two genera, Gordonia and Laplacea, should be merged. A
formal fusion was made by the present writer (Keng 1980).
For the Asiatic Gordonia species, the earliest synonym is Polyspora Sweet (1826).
It was named after a plant originally described under the name Camellia axillaris
Roxb. ex Ker. The flowers of this plant are almost sessile, the bracteoles and bracts
pass gradually into sepals which protect the rest of the flower parts in bud, the style
is stout and 5-fid at the apex. Other notable synonyms are Antheeischima and
Closaschima of Korthals (1842), who tended to think that plants from America and
Asia could not be possibly accommodated in the same genus. Korthals’ two new
genera were based on two Malesian plants, Gordonia excelsa Bl. and G. ovalis
Korth., respectively.
* Part One: The Genus Pyrenaria (Theaceae) in Malesia, Gard. Bull. Sing. 33 (1980) 264-289.
2 Gard. Bull. Sing. 37(1) (1984)
Like Korthals, Pitard, (1902) also emphasized the geographic separation between
Asia and America, and established the genus Nabiasodendron for the Asiatic
Gordonia species. Besides some external morphological characters, Pitard mentioned
some dubious internal anatomical features to stress that the Asiatic species should
be classified in a separate genus. For critical discussions of these and other
synonyms of Gordonia, see Burkill (1917) and Sealy (1958).
On the other hand, the first botanist who referred the Asiatic plants to Gordonia
is Blume (1825). The Javanese plant was at first described as Schima excelsa Bl. in
1823; two years later, it was transferred by himself to Gordonia, namely, G. excelsa
(BI.) BI.
II. A GENERAL ACCOUNT OF THE TAXONOMIC CHARACTERS
The Malesian Gordonia species are mostly small or medium-sized trees. Some
species, however, are lofty trees which can reach 60 m or more in height, while
others are shrubs which are usually found in high altitudes (e.g. G. imbricata and
G. vulcanica).
The leaves are alternate, well-spaced, mostly spirally arranged on branchlets. The
leaf-bases are acute, attenuate or rounded, sometimes decurrent along the petiole
and forming two narrow wings (e.g. G. multinervis, G. singaporeana and G. oblong-
ifolia), which are especially prominent in young trees and saplings. The nerves are
usually visible and often intermingled with smaller, parallel veins and complicately
looped and merged near the margin into submarginal veins or reticulations. The
secondary veins are invisible in a few species (e.g. G. imbricata and G. ovalis). The
tertiary veins and reticulations, in general, cannot be clearly seen, except after
treatment with dilute sodium hydroxide (as seen in some of the illustrations accom-
panying this paper). The venation is found to be useful in separating some related
species and in identifying sterile material of Gordonia.
The flowers are borne in leaf-axils, often in the upper axils, usually solitary, rarely
2-3 in a cluster. Each flower is usually subtended by one bract and two bracteoles
which are generally caducous. In G. maingayi, however, the bracteoles, bracts and
sepals, not being clearly differentiated, together form perules subtending the corolla
at anthesis. While in most of the other species, the bracteoles, bracts and sepals are
+ differentiated, the bract and bracteoles are caducous, only the sepals being
present at anthesis. The peduncles vary in length, the flowers are sessile or subsessile
in some species.
The differentiation between calyx and the corolla in most species is clear: sepals
are 5-6 in number, thick and hairy; petals are also 5-6, generally thick (at least in
their lower middle part) and briefly joined at the base into a short corolla-tube and
shed together after anthesis. In G. sarawakensis, the petals are 8-10 in number, but
they are relatively narrow, arranged in one series; whereas in G. borneensis and
G. polisana, the petals are also 8-10 in number, but are very broad, arranged in two
distinct series. In the two latter species, the petals of the outer series are smaller and
somewhat intermediate between sepals and petals.
Gordonia in Malesia 3
The androecium consists of numerous stamens which are in 3-4 whorls, usually
briefly fused at the base, sometimes in 5 (or 4) groups. The filaments are glabrous
or hairy, or more often the upper part glabrous and the lower, hairy. They are
always adnate to the base of the corolla and shed together after anthesis.
The gynoecium consists of a usually hairy, ovoid or subglobose ovary, one
(solitary or branched) or several (5, rarely 3 or 4 more) free styles, of which the tip
is enlarged into a stigma. Exceptions are found in G. ovalis and G. sarawakensis
in which species the styles are absent and the stigmas lie on top of the ovary. In each
ovary-locule, 3-5 ovules are found.
The fruits are capsular, angulate, cylindric-oblong to ellipsoid, rarely broadly
ovoid (e.g. G. borneensis) or subglobose (e.g. G. sarawakensis), dehiscing
loculicidally from the top downwards. As they are mostly 5-locular, they thus often
dehisce into 5 valves. In those species with ovary-locules other than 5, the number
of valves is generally in accordance with that of their ovary-locules; thus 3 (-4) in
G. scortechini, and 7-8 in G. sarawakensis. In the last named species, septicidal
lines can be clearly seen in the lower part of the fruit. In fully matured fruits of all
Gordonia species, the valves break away for most of their length from the stout
columella, but remain attached to it at the base for some time and eventually
disintegrate.
The seeds are usually ovoid or ellipsoid, always flattened, 2-3 in each locule, with
a large obliquely attached apical wing. Superficially they resemble the seeds of some
conifers (e.g. Pinus, Abies or Keteleeria). The embryo is large, slightly bent,
surrounded by a thin layer of endosperm.
Ill. TAXONOMIC TREATMENT
Gordonia Ellis
Gordonia Ellis, Phil. Trans. London 60 (1771) 518, ¢. 7] [Type species: Gordonia
lasianthus (Linn.) Ellis]; Benth. in B. & H. Gen. PI. 1 (1862) 186 (incl. Laplacea);
Szyszyl. in E. & P. Pflanzenfam. 3, 6 (1893) 185 (incl. Haemocharis); Melchior
in E. & P. Pflanzenfam. ed. 2, 21 (1925) 136, f. 63 (incl. Laplacea); Burkill,
J. Str. Br. Roy. As. Soc. 76 (1917) 133 (incl. Haemocharis); Merrill, J. Str. Br.
Roy. As. Soc. 86 (1922) 332; Sealy, Rev. Gen. Camellia (1958) 8; H. Keng, Gard.
Bull. Sing. 33 (1980) 308. Nom. cons.
Lasianthus Adans., Fam. 2 (1763) 398 [non Jack (1823), nec Zucc. ex DC (1836)].
Laplacea HBK, Nov. Gen. Sp. 5 (1822) 207, ¢. 46/.
Lindleya Nees. Flora 4 (1821) 299.
Haemocharis Salisb. ex Martius et Zucc., Nov. Gen. Sp. 1 (1824) 106.
Polyspora Sweet, Hort. Brit. ed. 1 (1826) 61.
Antheeischima Korth., Kruidk. (1842) 137, ¢. 27.
Closaschima Korth. op. cit. 139.
4 Gard. Bull. Sing. 37(1) (1984)
Carria Gardn. Calc. J. Nat. Hist. 7 (1847) 6.
Dipterospermum Griff., Notul. 4 (1854) 564.
Nabiasodendron Pitard, Act. Soc. Linn. Bordeaux 57 (1902) Cpt. Rend. Sc. 54.
Small to medium-sized trees, rarely shrubs. Leaves simple, coriaceous, alternate,
spirally or distichously arranged, entire or serrate. Flowers bisexual, axillary,
solitary or 2-3 (rarely more) congested in a cluster, shortly pedunculate or
subsessile; bracteoles and bracts 2-3 or more; sepals 5-6, unequal; petals 5-6, rarely
9-10, unequal or subequal, usually briefly fused at the base; stamens numerous, in
3 or 4 whorls, shortly connate at the base and often briefly adnate to the corolla,
sometimes in 4-5 less distinct fascicles; anthers versatile, on a short or long filament;
ovary mostly 5-locular, rarely 8-10 or 3-4 locular; ovules 2-8 (usually 3-5) in each
locule, on axile placentation in two vertical rows; styles mostly 5, sometimes more
or only 3, fused to varying extent proximally, or sometimes totally free. Fruit a
woody capsule, ovoid-cylindric, bluntly angulate, dehiscing loculicidally from apex
to base along a persistent central columella. Seeds usually 2-5 in each locule,
compressed, with a thin membranous, oblique, unilaterally attached wing; testa soft
woody; embryo oblong, straight or slightly oblique; endosperm of a thin layer,
enveloping the embryo.
A genus with about seventy species; occurs in SE. Asia and America. Forty or so
Asiatic species are found from India, Sri Lanka, Burma, Thailand, Indochina,
S. China to Taiwan and southwards to Malesia (the Malay Peninsula, Sumatra,
Borneo, Java, the Philippines, Celebes to New Guinea). Thirty or so American
species are concentrated in Central America and the West Indies (most of them
formerly arranged under the generic names Haemocharis or Laplacea), with a few
species in the northern parts of South America, and only one species (which is the
type species) in the southeast of the United States of America.
About 21 species occur in Malesia.
KEY TO THE MALESIAN GORDONIA SPECIES
1. Average leaves very large, generally over 15 cm long
2. Peduncles of flowers generally less than 0.5 cm long
3. Leaf-apex usually acuminate; nerves 10-11 pairs; flowers 5-6 cm across (the Malay Peninsula)
20. G. taipingensis
3. Leaf-apex usually rounded and mucronate; nerves up to 18 pairs; flowers 3-4 cm across (the
Malay Peminsula) ccs ct crsece castes tae caer deka lec uae poe eicn ch a onc VeReccauren ae 12. G. multinervis
2. Peduncles of flowers usually over 1.5 cm long
4. Petioles conspicuously winged; sepals persistent in fruit
Gordonia in Malesia 5
4. Petioles not winged; sepals caducous in fruit; capsules broadly ovoid, about 5 cm long (Borneo:
SADE): «ISS Sod on 6 Ge SEO ODE ne Sere eS Sooo re 4. G. grandiflora
1. Average leaves smaller, generally less than 15 cm long
6. Average leaves relatively small, usually less than 9.5 cm long
7. Leaf-apex generally obtuse or rounded, often emarginate; plants often found in montane or
lower montane forest
8. Petals mostly 5, rarely 6, in one whorl; leaves thick coriaceous
9. Leaves elliptic, 4.5-9.5 cm long (the Malay Peninsula & Borneo) ...................
ee eee eh AEE eal save Stele cael s, Atinls te ete e Se eRe hh ee wes 7. G. imbricata
9. Leaves ovate or broadly oblong 2-5 cm long; capsules 2.5-3 cm long (Sumatra)......
et Bde $e4 Son Nee oo Se Oe rd ne oe ee 21. G. vulcanica
8. Petals 8-10, in two whorls; leaves thin-coriaceous (the Philippines) ...... 16. G. polisana
7. Leaf-apex mostly acute or acuminate, pointed or blunt and rarely rounded; plants generally
found in lowland or at medium altitudes
10. Ovary 3- (rarely 4-) locular, the 3 (rarely 4) styles free; flowers very small (1.5-1.8 cm
ERA SSNilee Wi tLAVELCHIFISUNA) Nu ec hve s «0 cles Wage aerbe-w «age Pe eos + mae 18. G. scortechinti
10. Ovary 5- (rarely 4-) locular; styles fused together into a column, sometimes branched above;
flowers various
11. Number of bracteoles, bracts and sepals rather large (around or over 10) and less clearly
differentiated, together forming overlapping perules surrounding the flowers at
AUIMeSISH EN eg Vic AVEEEMINSULA) phe cree cts cust ahaa sieve vs 2 os aoe 10. G. maingayi
11. Number of bracteoles, bracts and sepals generally less than 8, and usually differentiated;
normally bracteoles and bract caducous and only the sepals present at anthesis
12. Leaf-apex generally acuminate and obliquely caudate; capsule cylindric, pointed,
3.5-5 cm long (the Malay Peninsula) ..................... 15. G. penangensis
12. Leaf-apex acute or sometimes acuminate, but rarely caudate; capsules usually less
than 3 cm long
13. Leaves 3-6 cm wide, often glaucescent beneath; flowers 3-3.5 cm across;
styles solitary, with a club-shaped tip (Borneo: Sarawak & Sabah) ........
|S Sch ahha s «Sac D5 ers eee eee ee Be NESS a 5. G. havilandi
13. Leaves generally less than 3 cm wide, not glaucescent below; flowers less than
2.5 cm across; styles either branched or absent
14. Nerves visible; style 1.5-2.5 mm long, 5-branched above (Borneo)
ere es ee ei on a2 St aha «dag alte Bee eet s 11. G. marginata
14. Nerves nearly invisible; style absent, only 5 tiny protrusions (less than
1 mm long) on top of the ovary representing the stigmas (Sumatra)
HABE ORE ES SE ee SA Se Lae cr Renernern chor Pear ee 14. G. ovalis
6. Average leaves generally between 10 to 14 cm long
15. Petals 7-8, in one series; ovary 8-10 locular, style absent or very short; fruit broadly ovoid or
subglobose (Borneo: Sarawak & Sabah)..............---+----+---+- 17. G. sarawakensis
15. Petals 5-6; ovary 5- (rarely 4-) locular; style present; fruit generally ovoid or cylindric
6 Gard. Bull. Sing. 37(1) (1984)
16. Leaf-margin entire, subentire, very rarely serrulate (Java, Bali, Celebes) .............
Bera tea Sec Aaaa amore os von Comm ano Sema mstoe Ss Cumort 8. G. integerrima
16. Leaf-margin serrate or serrulate
17. Peduncles of flowers usually less than 0.5 cm long
18. Leaves subsessile or sessile, the blade usually tapering towards the base and
winged; capsules 3 or 5 cm long
19. Leaves thin-coriaceous; nerves 10-12 pairs; flowers 4—5 cm across (the Malay
Peninsula) svncc cy oe ones an a eee 19. G. singaporeana
19. Leaves membranous; nerves 12-18 pairs; flowers 2-3.5 cm across (the Malay
Peninswla)) ss Stas nese ie oe aioe te oe Oe 12. G. multinervis
18. Leaves petiolate, petiole 5-8 mm long; leaf-base not winged; capsules 2 or
25cm longa(the Malayabeninsula) msec ieee 6. G. hirtella
17. Peduncles of flowers generally 0.5-1 cm long, sometimes longer
20. Style columnar, the tip enlarged, discoid, shallowly 5- (rarely 4-) lobed
21. Flowers 2.5-3 cm across; capsule 3.5-4.5 cm long (Sumatra and Java)
ee ere eee ent S On dio c.G0 boy Ca ccnudo nd omatrec 3. G. excelsa
21. Flowers 7-8 cm across; capsules 3-4 cm long (the Philippines)
sw Sigiaieca audtaaee Sra" p ah aie TENE ota See a Pere SLE eee Oe ee eee 9. G. luzonica
20. Style distinctly 5-branched above (Celebes, Lesser Sunda Isls., Moluccas and
NewiGuiniea) ly « 2eytas teil Ss hoe eon Shree src ace Lomas 1. G. amboinensis
1. Gordonia amboinensis (Miq.) Merr., J. Str. Br. Roy. As. Soc. 86 (1922) 332.
Fig. 1
Laplacea amboinensis Miq., Ann. Mus. Bot. Lugd. Bat. 4 (1968) 114 (based on
Lignum mucosum vel Caju lapia Rumph., Herb. Amb. III (1743) 203,
tab. 130).
Haemocharis amboinensis (Miq.) Burk., J. Str. Br. Roy. As. Soc. 76 (1917) 141,
158.
Gordonia rumphii Merr., Interpret. Herb. Amb. (1917) 368.
Gordonia brassii Kobuski, J. Arn. Arb. 21 (1940) 135. Syn. nov.
Gordonia papuana Kobuski, op. cit., 136 (incl. var. acuminata & var. montana);
W. R. Barker, Brunonia 3 (1980) 8 f./. Syn. nov.
Gordonia sp. Kobuski, op. cit., 139 (citing Brass & Versteegh 13169 from
W. Irian).
Tree, to 20 (-25) m tall. Bark grey to brown, smooth or shallowly fissured; inner
bark beefy red. Young twigs and buds covered with short silky hairs or glabrous.
Leaf-blades thin coriaceous or membranous, lanceolate, narrowly ovate to ovate-
elliptic, 7-14 (-20) cm long, 3-5.5 (-8) cm wide, apex bluntly acuminate, sometimes
obtuse or retuse, base acute or attenuate; margin shallowly crenate to undulate, or
subentire; nerves 9-12 pairs, intermingled with less conspicuous veins and merged
Gordonia in Malesia 7
and looped into submarginal reticulation; glabrous above, glabrescent and sometimes
papillate beneath; petiole 0.5-1 (-1.5) cm long, thickened. Flowers in the upper
axils, solitary; peduncle 0.5-1 (-2) cm long, slender. Bracteoles and bracts 2-3,
caducous. Sepals 4-5, unequal, deltoid, lunate or broadly ovoid, 4-6 mm long,
coriaceous, densely covered with silky hairs externally. Corolla 3-4 cm across,
creamy yellow or white; petals 5-6, broadly ovate to subrounded, concave, 1.5-2 cm
long, the outer surface sericeous in the central portion and below, glabrous around
the margin. Androecium 7-8 mm long, the filaments glabrous, except the base
which is covered with short hairs. Gynoecium 5-8 mm long, shortly silky; ovary
globose or ovoid, 3-4 mm in diameter; style (4-) S-branched above. Capsule
Fig. 1. Gordonia amboinensis (Miq.) Merr.
Ambon, Teysmann H. B. 5587 (Herb. Lugd. Bat. No. 908, 249-227) (lectotype); Halmahera,
Neth. Ind. For. Serv. 31360 (fr.).
8 Gard. Bull. Sing. 37(1) (1984)
oblong cylindric, 2.5-3 (-3.5) cm long, puberulous, dehiscing into 4 or 5 valves.
Seeds about 2 cm long including the wing.
DISTRIBUTION. Malesia (Celebes, Moluccas, Lesser Sunda Isls. and New Guinea)
and Bismarck Archipelago (New Britain and New Ireland). (All specimens cited
below from L).
Celebes. Minahassa, Koorders 18910, 18912, 18913, 18914; Molili, Neth. Ind. For. Serv. 481.
Moluccas. G. Sembilan, Halmahera, Pleyte 341; Ternate, Neth. Ind. For. Serv. 24534. P. Buru, Neth.
Ind. For. Serv. 31360, Ceram, Eyma 2214, Kuswata & Soepadmo 240. Ambon, Robinson 276 (isotype
of Gordonia rumphii Merr.); De Vriese & Teijsmann Herb. Lugd. Bat. No. 908, 251-414; Teysmann
H. B. 1970 (Herb. Lugd. Bat. No. 908, 251-224, & -225), H. B. 5587 (Herb. Lugd. Bat. No. 908,
Fig. 2. Gordonia borneensis H. Keng sp. nov.
Kalimantan, Kostermans 35063 (fl.); Kostermans 13627 (fr.).
Gordonia in Malesia 9
249-227, lectotype of Laplacea amboinensis Miq.)
Lesser Sunda Isls. Bali, Mt. Batukau complex, Kostermans et al. KK-SS 137, 174. Flores, near Wae
Mao, Kostermans 838; Monde Hill, Schmutz 3552, 3711.
Irian Jaya (representative specimens only). Sidoarsi Mts., Hollandia D. 3t, Jwanggin BW 9037; Sibil
Valley, Star Mts., Ka/kman 4285; near Andai, SW. of Manokwari, Koster BW 11798; near Soendei, Isle
of Biak, Moll BW 9721; Cycloop Mts., Hollandia Dist., van Royen 3635; Mt. Tohkiri, Vogelkop
Peninsula, van Royen & Sleumer 7253; Tobie, Kebar Valley, Schram BW 7970; Aisao, Japen Isl., Schram
BW 10584.
Papua New Guinea (representative specimens only). Arau, Eastern Highlands Dist. Brass 32063;
Mt. Dayman, Maneau Range, Brass 23243; Torricelli Mts., W. Sepik Dist., Darbyshire 358; near Lake
Birip, Wabag, W. Highlands Dist., Flenley ANU 2738; Above Akuna, E. Highlands Dist., Hartley
11986; upper Oriomo, Western Dist., Havel 17246; Kwa Mountain, Rossel Isl., Henty NGF 27071; near
Frieda River, W. Sepik Dist., Henty & Foreman NGF 42579; Mt. Rawlinson, Morobe Dist., Hoogland
9262; Mt. Hunstein, Sepik Dist., Hoogland & Craven 10951; near Hagen Station, W. Highlands Dist.,
Hoogland & Pullen 5965; Lake Erobo, S. Highlands Dist., Powell 2423; Ingembit, Western Dist.,
Ridsdale, Henty & Galore NGF 31943; Marapuna, Eastern Highlands Dist., van Royen NGF 15053;
Mt. Kumme, Central Dist., van Royen NGF 20346, 20416; Above Kiburu, Southern Highlands Dist.,
Schodde 1371; Mt. Simpson, Milne Bay Dist., Schodde 5522; Mt. Pigini; Central Dist., Stevens LAE
50449; near Kapiaggo, Western Highlands Dist., Vandenberg, Womersley & Galore NGF 39995; near
Nondugl, Eastern Highlands Dist., Womersley 4860.
ECOLOGY. From lowland rain or swamp-forest to lower montane or montane
moss forests, in Eucalyptus-Gironniera, Castanopsis-Quercus, or Nothofagus
forests; on sandy or clay soil, also on limestone ridge and in disturbed areas and
grassland. Altitudes 50-2000 m. Fl.: Apr.-Nov., fr.: Jun.-Jan. (few collections
only).
VERNACULAR NAMES: adikelp (Japen), alimp (Tagoba), bado (Motus), baif
(Gab-gab), benelemonde (Hattam), bwabwa (Wafu), dapiri (Mid. Waria), dimi
(Kiwai), iniaili (Je), kawal-gugn (Mini), kerkebo (Flores), kimkaroo (Tehid),
la (Enga), kilimdan (Sepit), koka kaber (Knambiadi), kuku (Kutub), La (Enga),
naningning (New Britain), oytungo (Aseki), reik (W. Biak), sjioe (Andjai), tawan
(Wipi), timor (Mendi), toani (Ormoe), tokoi (Manikiong), totona Rombo
(Garumaia), tugera (Waskuk).
NotE. Kobuski (1940) examined 8 specimens of Gordonia from New Guinea then
available to him and recognised 4 entities — one uncertain species and two new ones
of which one consisted of two varieties. Barker (1980) in a recent revision correctly
pointed out that there is only one homogeneous species in New Guinea, showing
little or no evidence of polymorphism.
When the New Guinea material was compared with those from Moluccas, Celebes
and the Lesser Sunda Islands, they were found to agree in leaf, flower and fruit
characters. I therefore adopt Gordonia amboinensis (Miq.) Merr. as the name for
this widely distributed species.
Regarding the field notes of numerous collectors, the following two are noteworthy:
on Brass 23243 was stated that the flowers are 6-6.5 cm in diameter; this appears
much larger than the usual size (3-4 cm in diameter). Schram BW 10584 indicated
10 Gard. Bull. Sing. 37(1) (1984)
the flowers are pink; all the others are recorded as white, creamy white or creamy.
Some specimens from Flores (e.g. Schmutz 3711) have much smaller capsules (less
than 2 cm long) which are probably immature or from trees growing on extremely
poor soil. They also may represent a new entity.
Gordonia amboinensis is closely allied to G. excelsa. The two are quite similar in
shape and venation of their leaves. However, they can be differentiated by their
shape and size of the flowers and geographically they are more or less demarcated.
Fig. 3. Gordonia excelsa (B\.) BI.
Java, Cult. in Hort. Bog. No. 16/7 (Anno 1901); Koorders 14735 (fr.).
Gordonia in Malesia 11
2. Gordonia borneensis H. Keng, sp. nov.
Figs 2
Arbor 20-30 m alta. Folia angusto-elliptica, angusto-obovata vel oblanceolata, 18-25 (-30) cm longa,
apice acuta vel cuspidata, basi attenuata vel cuneata, coriacea tenui, nervis lateralibus 20-22, petiolo 3-5
mm longo. Flores flavi, axillares, solitarii, 6-7.5 cm diametro; bracteae 2, caducuae; sepala 5, subrotun-
data, 1-1.2 cm longa; petala circa 10, 1.5-3.5 cm longa. Gynoecium 2.5-3.5 cm longum; styli ad apicem
discoideum. Capsula obovoidea, 2-3 cm longa, valvis 5.- Typus: E. Kalimantan, Kostermans 7084, in L.
A tree, 20-30 m tall. Bark greyish brown, smooth, or minutely fissured; young
branches stout, glabrescent. Leaf-blades thin coriaceous, narrowly elliptic, narrowly
obovate or oblanceolate, apex acute or obliquely cuspidate, base attenuate or
cuneate and narrowly winged, 18-25 (-30) cm long, 6.5-7.5 (-10) cm wide; margin
finely crenulate-serrulate for the most part, subentire or entire near the base; side
veins 20-22 pairs, further intermingled with less distinct and smaller ones in between
and intricately interlooped near the margin; glabrous on both surfaces, thinly
puberulous on the midrib beneath; petiole 2-5 mm long, thickened. Flowers
solitary, in upper leaf-axils, sometimes several together on the branch tips; peduncle
1.5-2.5 cm long, often elongate and much thickened after anthesis. Bracts 2, a short
distance below the calyx, caducous. Sepals 5, suborbicular, 1-1.2 cm long cor-
iaceous, silky-hairy externally. Corolla 6-7.5 cm across, yellow to bright yellow (fide
Kostermans); petals about 10, in 2 series, sericeous externally, petals of the outer
series thicker (thin coriaceous) and smaller (1.5-2 cm long), broadly oblong or
sub-rounded, somewhat intermediate between sepals and the inner petals; petals of
the inner series thinner (membranous) and larger (2.5-3.5 cm long), broadly ovate
or broadly oblong, apex often notched, base sometimes abruptly narrowed, 2.5-3.5
cm long. Androecium 1.2-1.5 cm long; stamens numerous, in 3-4 whorls, connate
below, the filaments glabrous. Gynoecium 2.5-3.5 cm long; style 1, 1.5-2 cm long,
the tip discoid, shallowly 5-lobed into stigmas; ovary ovoid, 1.-1.5 cm long, densely
covered with yellowish brown hair. Capsule obovoid, S-angulate, 2-3 cm long; calyx
persistent. Seeds about 2 cm long including the wing.
DISTRIBUTION. Malesia (Brunei, Sabah and Kalimantan).
Brunei. Kuala Sebatu, Ashton BRUN 354 (BO, L, SING).
Sabah. Monsapol F. R., Sipitang, Aban Gibot SAN 65950 (L); Kalabakan, Tawau, Bakar SAN 24974
(BO, L, SING); Benawood, Tawau, Felilis & Sumbing SAN 88140, 88418 (SAN); Sandakan, J. Singh
SAN 53427 (L, SING); Ulu Dusun, Sandakan, Tarodop SAN 87724 (SAN); Beaufort, Wood & Kapis
SAN 16984 (BO, L, SAN, SING).
Kalimantan. Tg. Bangko, near mouth of Sg. Mahakam, Kostermans 7084 (L, holotype; BO, isotype);
Sangkulirang, Karangan River, Kostermans 13627 (BO, L); Sg. Tiram Balikpapan, Kostermans 35063;
Landak, W. Borneo, Ngabang 6410 (L).
EcoLocy. In primary forests, on low ridges in sandy soils or on seasonally
inundated riverine alluvium. Alt. 20-100 m. Fl. May-Aug.; fr.: Aug—Oct.
VERNACULAR NAMES: obah (Sabah), rawali batu (Kalimantan).
NOTE. This species differs from G. grandifolia Merr. and G. oblongifolia Steenis
in that the flowers have about 10 petals in two series (instead of 5 petals in one series)
and in the much smaller fruits and seeds.
Fig. 4. Gordonia grandiflora Merr.
Sabah, Wood SAN 16185 (fr.); Meijer SAN 94245 (fl.).
3. Gordonia excelsa (Bl.) Bl., Bijdr. (1825) 130; Miq., Fl. Ind. Bat. 2 (1857) 489;
Koord. & Val., Med., ’s Lands Pl. Tuin (1896) 289 (incl. var. macrocarpa
K. & V.); Koord., Exk. Pl. Jav. 2 (1912) 608; Koord. -Schum. Syst. Verz. fam.
180 (1913) 35; Burk., J. Str. Br. Roy. As. Soc. 76 (1917) 155; Merr. Contr. Arn.
Arb. 8 (1934) 106; Back. & Bakh. f. Fl. Jav. 1 (1963) 320. Fig. 3
Schima excelsa Bl., Cat. (1823) 80.
Antheeischima excelsa (Bl.) Korth., Kruidk. (1842) 138, ¢. 27.
Gordonia acuminata Choisy in Zoll. Syst. Verz. 2 (1854) 144.
Small or large tree, to 40 m tall. Bark smooth, greyish brown. Young twigs
slender, covered with soft yellow hairs or puberulous; old branches glabrescent.
12
Gordonia in Malesia 13
Leaf-blades thin coriaceous or membraneous, elliptic or narrowly elliptic or oblong-
lanceolate, apex obtuse or acuminate, base attenuate, (7-) 10-14 (-20) cm long, (2.5-)
3.5-4.5 (-6) cm wide, equal or sometimes slightly bent and unequal-sided; margin
remotely serrulate; midrib impressed above and elevated below; nerves 12-15 pairs,
connected and merged into submarginal reticulation; glabrous above, appressed
pubescent beneath; petiole 0.5-1 cm long, puberulous. Flowers in axils of the upper
leaves or subterminal, usually solitary; peduncle 0.5-1 (to 2 cm in fruit) cm long,
stout, velutinous. Bract and bracteoles about 3 caducous. Sepals 5-6, unequal,
reniform to suborbicular, 7-8 mm long, sericeous. Corolla 2.5-3.5 cm across, white,
fragrant; petals 5-6, deltoid-reniform, often concave and notched, 1-1.7 cm long,
1.2-2 cm wide, thin coriaceous, sericeous on both sides except the edge of the inner
surface, which is glabrous. Androecium about | cm long; filaments nearly glabrous
and briefly united below. Gynoecium about | cm long; style 1, stout, 5-6 mm long,
glabrescent, the tip disk-shaped, shallowly undulate or shortly lobed into 5 stigmas;
ovary globose, 4-5 mm across, densely sericeous. Capsule cylindric, 3.5-4.5 (-5) cm
long, bluntly angulate, pubescent, dehiscing into 5 valves. Seeds 2-2.5 cm long
including the wing.
DISTRIBUTION. Malesia (Sumatra & Java).
Sumatra. Without precise locality, Forbes 2053 (L). West Coast, Neth, Ind. For. Serv. 2860, 2878 (L).
Java. G. Loechoer, Bakhuizen v/d Brink 7146 (L); G. Salak, Blume Herb. Lugd. Bat. 908, 251-415,
-42] (lectotype), -422, -43] (L); without locality, Houtsoorten van Gedeh 638 (L), Junghuhn 73, 83;
Besuki, Koorders 14735 (holotype of G. excelsa Bl. var. macrocarpa K. & V.), 20381, 28675, 32342 (L),
Sukari FRI 7525; Java, Teysmann s.n. in 1860 (L); Preanger, Winckel 486 (L) Wind s.n. in 1918 (L).
Eco_Locy. In primary forests, altitude 600-1700 m . Fl.: Apr.-Nov. fr.: Jun.-
Feb.
NoTE. This is the first Asiatic species described under Gordonia, a genus
previously known only from N. America.
4. Gordonia grandiflora Merr., J. Str. Br. Roy. As. Soc. 86 (1922) 331;
Masamune, Enum. Phan. Born. (1942) 472. Fig. 4
Small or large tree, 10-28 m tall. Bark greyish brown fissured or scaly. Branchlets
stout, glabrescent. Leaf-blades coriaceous, elliptic, narrowly elliptic or obovate,
sometimes oblanceolate, 15-25 (-35) cm long, 6.5-8 (-10) cm wide, apex obtuse or
abruptly acute, base attenuate, decurrent and narrowly winged or sometimes
rounded; margin crenulate; nerves 13-16 pairs; glabrous above, scattered with short
hairs beneath; Petiole 3-5 mm long, thickened. Flowers solitary, in upper leaf-axils;
peduncles 2-3 cm long, stout. Bracts and sepals not seen. Corolla 8-9 cm across,
white; petals 5, membranous, pubescent externally, broadly oblong or obovate,
4-4.5 cm long, narrowed beneath, sometimes notched or bilobed above. An-
droecium 1.5-2 cm long, the filaments pubescent, connate only at the base and ad-
nate to the corolla. Gynoecium not seen. Fruit (nearly mature) broadly ovoid, about
5 cm long, dehiscing into 5 valves. Seeds about 4 cm long including the wing.
14 Gard. Bull. Sing. 37(1) (1984)
DISTRIBUTION. Malesia (Sabah).
Sabah. Rosab, near Kudat. P. Castro 972 (A, isotype [photo]); Ranau, Kinabalu National Park, Aban
Gibot SAN 76577, 76605, 76608 (SAN), Meijer SAN 94245 (SAN); Kota Belud, Kandasan, Wood & Kapis
SAN 16185 (B, SING).
EcCOLoGy. In primary forest, on hill side or on dry slopes; alt. 50-1,300 m.
Fl.: Sept. & Nov. (2 collections); fr.: Feb. (1 collection).
NOTE. This species, as mentioned in some of the field notes is common in the
Kinabalu National Park. However, among the material available, only Meijer 94245
Fig. 5. Gordonia havilandii Burk.
Sarawak, Haviland 1995 (syntype, fl.); Rosali S 15184 (fr., immature). Sabah, Amin G. et al.
SAN 93856 (fr., mature).
Gordonia in Malesia 15
bears a detached corolla, and Wood & Kapis 16185 (2 specimens) bears two nearly
mature fruits, the others being sterile. A photo of the type (Castro 972) was made
available through the courtesy of the Director of the Arnold Arboretum, Harvard
University.
In Merrill’s original description, the sepals are ‘‘coriaceous, broadly ovate to
orbicular-ovate, rounded, glabrous or slightly pubescent, 1.5-2 cm long and
somewhat accrescent in anthesis’’, the ovary is ‘‘pubescent’’ and the style is
“*slender, glabrous, up to 2.5 cm long’’. He also mentioned that the flower is ‘‘10-11
cm in diameter’’. This is larger than in Meijer 94245, which is 8-9 cm across.
This species is related to G. oblongifolia of Sumatra. Because of the scarcity and
imperfection of fertile materials in both species, close comparisons cannot be made
at the present.
5. Gordonia havilandii Burk., J. Str. Br. Roy. As. Soc. 76 (1917) 157; Merr.,
op. cit. spec. no. 20 (1921) 389. Fig. 5
Small to medium-sized tree about 25 m tall and 30 cm in diameter. Bark smooth,
hoop-marked, mauve brown and grey. Young twigs slender, covered with yellow or
black hispid hairs. Older branches greyish brown, glabrescent. Leaf-blades char-
taceous or thin coriaceous, elliptic, rarely narrowly or broadly elliptic, obtuse or
acuminate, occasionally cuspidate, base cuneate or attenuate, 7-10 (-12) cm long,
3-6 cm wide, symmetrical or sometimes slightly unequal; margin entire or nearly so;
midrib impressed above, tomentose and glaucescent beneath; nerves 9-10 pairs, less
conspicuous above; petiole slender, 1-1.5 cm long, puberulent. Flowers subterminal
and in upper axils, solitary or 2-3 together; peduncles 2-3 mm long, hispid.
Bracteoles and bracts 2-3 silvery puberulous, orbiculate-deltoid, 2-3 mm long.
Sepals 5-6 suborbicular, 7-8 mm long leathery, sericeous externally. Corolla 3-3.5
cm across, yellow or pale yellow; petals 5-6, suborbicular to reniform, 0.8-1.2 cm
long, thin leathery, concave, sericeous externally. Androecium 6-7 mm long, the
filaments thinly sericeous in the lower part, and united at the base. Gynoecium
about 1 cm long; style 1, 6-7 mm long, glabrescent; stigmas 5, very short; ovary
globose, 3-4 mm across, densely sericeous. Capsule cylindric, 2.5-4.5 cm long,
1.2-2 cm across, pointed, bluntly 5 angled, thinly strigose; calyx persistent. Seeds
about 2-4 cm long including the narrow wing.
DISTRIBUTION. Malesia (Borneo, known only from Sarawak and Sabah).
Sarawak. Mt. Serapi, alt. 900-1000 m, Haviland 67 (SING, holotype), 1995 (L, SING, syntype);
Burgoh Range, near Summit, Anderson et al. S 29323 (L); Ulu Kenyana, Mukoh, Ashton S 195492
(SING); Mt. Meroyang, Ulu Lundu, Ashton S 1886] (L); Semengoh F. R., Banyeng & Benang S. 25490
(L, SING), Banyeng & Jugah S 26882 (L), Rosli S 15184 (L); Mt. Santubong, Chew 1386 (L, SING);
Bt. Lambir, Miri, Dan bin Hj. Bakar 3090 (L); Sg. Belaban, Lawas, Ilias S 26319 (L).
Sabah. Telupid, Sandakan, Aban Gibot & Joseph SAN 94036; Kiabau, Beluran, Amin G. et al. SAN
39856; Sg. Korang, Lamag, Madani SAN 76315; Tongod, Sundaling SAN 90348 (all in SAN).
EcoLocy. In heath or fagaceous forests, from lowland to the summit of sand-
stone hills; alt. 15-1850 m. Fl.: Aug.-Nov., fr.: Dec, Mar. & Aug.
16 Gard. Bull. Sing. 37(1) (1984)
VERNACULAR NAME: /inggai.
NOTE. This species can be recognized by the elliptic leaves with a tomentose and
glaucescent undersurface.
6. Gordonia hirtella Ridl., J. Str. Br. Roy. As. Soc. 73 (1916) 142; Burk., op. cit.,
149, f. 7; Ridl., Fl. Mal. Pen. 1 (1922) 204; H. Keng in Ng, Tr. Fl. Mal. 3
(1978) 285. Fig. 6
Tree, 20 m high; bole coarsely and shortly fluted. Bark grey, smooth-rugulose.
Young branches slender, silky-pubescent. Leaf-blades thin coriaceous, elliptic,
elliptic-lanceolate or narrowly ovate; apex acute or acuminate, base cuneate or
=
Fig. 6. Gordonia hirtella Ridl.
Malaya, Ridley 7350 (isotype, fl.); Whitmore FRI 20049 (fr.).
Gordonia in Malesia 17
attenuate, 6-13 (-15) cm long, 2.5-4 (-5) cm wide; margin crenulate-serrate or sub-
entire; nerves and reticulations barely visible above, indistinct below; shining dark
green and glabrescent on the upper surface, light green, strigose on and around the
midrib especially near the base on the lower surface; petiole 3-6 mm long, densely
hispid. Flowers subterminal or in leaf-axils, usually solitary, densely covered with
greyish yellow hairs externally; peduncles subsessile or very short (2-3 mm long),
appressed silky. Bracteoles and bracts about 3, 1-2 mm long. Sepals about 5, ovate
to broadly orbicular, 3-5 mm long. Corolla 2-2.5 cm across, cream in colour; petals
5, oblong to broadly obovate, 1-1.2 cm long, briefly joined at the base. Androecium
4-5 mm long, the filaments connate below and adnate to the corolla tube.
Gynoecium 5-6 mm long, the ovary densely covered with greyish yellow hairs, the
style and its branches glabrescent. Capsule 2-2.5 cm long, 5-valved, puberulous;
sepals persistent. Seeds 1.5-2 cm long including the wing.
DISTRIBUTION. Malesia (the Malay Peninsula)
The Malay Peninsula, Perak, Batu Puteh, Wray 1116 (syntype, SING). Selangor, Bt. Kutu, Ridley
7350 (isotype, SING); Bt. Etam, Kelsall 1848 (SING). Pahang, Sg. Telom Ridge, Whitmore FRI 20049
(L).
EcCOLoGy. In forests and on ridges; alt. 400-1000 m. Fl.: June, fr.: May.
Note. Burkill (l.c.) stated that this species ‘‘occurs on the central chain of
mountains of the Malay Peninsula from Gunong Batu Puteh in Perak to the west
of Tapah ... through Bukit Kutu in Selangor ... to Bukit Etam ... on the Selangor
Negri Sembilan boundary’’. Apart from the specimens cited by Ridley and Burkill,
which were all collected near the end of the last century, the only single recent
collection referable to this species was the collection made by Whitmore from
Sg. Telom ridge in NW. Pahang.
7. Gordonia imbricata King, J. As. Soc. Bengal 59, 2 (1890) 204, Ann. Roy. Gard.
Calc. 5 (1896) 148, p/. 179; Burk., op. cit., 157, f. 75; Ridl. Fl. Mal. Pen. 1
(1922) 203; H. Keng in Ng, op. cit., 285. Fig. 7
Low shrub or stout tree, to 15 m tall. Bark brown, finely cracked. Young twigs
stout, glabrescent or thinly covered with scattered pilose hairs; older branches often
finely fissured. Leaf-blades coriaceous or thick coriaceous, narrowly elliptic to
oblanceolate or elliptic; apex obtuse or rounded, sometimes shallowly retuse, base
cuneate, 4.5-9.5 (-12) cm long, 2-4 (-5) cm wide; margin remotely undulate or
entire, sometimes revolute; midrib sulcate above; nerves about 10-12 pairs, barely
visible above, and obscure beneath; drying green; glabrous above, brownish,
verruculous and subglaucous beneath; petiole stout, 0.5-1.5 (-2) cm _ long,
puberulous. Flowers axillary or subterminal, solitary or 2-4 and congested near the
tip; peduncle absent or very short (1-2 mm long, stout). Bracteoles, bracts and
sepals forming an involucre about 1-1.5 cm high, densely woolly externally, and
increasing in size from the lower bracteoles (deltoid, 2-3 mm long) to the upper
sepals (broadly ovoid to suborbicular, 8-10 mm long). Corolla 2.5-3.5 cm across,
white to butter yellow (fide Whitmore); petals 5, suborbicular to reinform, 1.2-1.7
cm long, for the most chartaceous and glabrous except the lower portion which
18 Gard. Bull. Sing. 37(1) (1984
is coriaceous and woolly and similar to the sepals. Androecium 8-10 mm long, the
filaments velutinous and connate below, Gynoecium 8-9 mm long; style thick,
columnar, 2-3 mm long, briefly 5-branched near the tip, glabrous except the lower
portion; ovary globose, 5-6 mm long and wide, densely covered with golden
velutinous hairs. Capsule cylindric, 4-5 cm long, strigose, dehiscing into 5 valves;
sepals persistent until the dehiscence of the capsule. Seed 3-3.5 cm long including
the wing.
DISTRIBUTION. Malesia (the Malay Peninsula and Borneo (Sabah)).
The Malay Peninsula. Perak, G. Kerbau, Robinson s.n. in June 1913, Scortechini 402 b (SING,
duplicate of type). Kelantan, G. Stong, Symington 37695 (SING); Gua Ninik, Henderson SFN 19745
(SING); G. Rabong, Soepadmo & Mahmud 1063 (L). Pahang, G. Tahan, Kloss 12108, 12296 (SING),
\& cm 2
Y
Fig. 7. Gordonia imbricata King
Malaya, Ng KFN 98043 (fr.); Whitmore KFN 8727 (f1.).
Gordonia in Malesia 19
Ridley 16021, 16263 (SING), Md. Haniff & Md. Nur 7885 (SING), Seimund 566 (SING), Holttum 20707
(SING), Corner s.n. in Sept. 1937 (L); Cameron Highlands, Wyatt-Smith KFN 94594 (L); G. Rapis,
Symington & Kiah 28909 (SING). Trengganu, G. Tabu, Md. Shah et al. 3284 (SING); G. Pandan,
Whitmore FRI 12673 (L), Kiah SF 33358 (SING). Johore, G. Blumut, Holttum 10719 (SING), Md. Shah
& Samsuri 2227 (SING), Ng KFN 98043 (SING, L), Whitmore KFN 8727 (SING, L), Suppiah FRI 17839
(SING, L).
Sabah. Mt. Kinabalu, J. & M. S. Clemens 50720(L), Kokawa & Hotta 5628 (SAN), Hotta 3780 (SAN),
Meijer SAN 29197 (SAN).
EcoLoGy. In lower and upper montane forests, on ridges or in Leptospermum
scrub. Alt. 1000-2300 m; once recorded at 650 m (Henderson SFN 19745).
Fl.: Mar.-Sept., fr.: Sept.-Jan.
NOTE. This species is allied to G. vulcanica of W. & Central Sumatra. Both
species are characterized by the sessile flowers with woolly, imbricate bracteoles,
bracts and sepals. Both occur at high elevations. They differ from each other in
shape, size and venation of the leaves and in the detailed structures of the flowers
and fruit.
Original materials of G. imbricata King were collected from the northern Malayan
mountains (at about 1700-2200 m). Several recent collections from G. Blumut
(at about 1000-1100 m) in S. Johore match well with the original except that the
leaves are slightly thinner and the venations clearly visible, and the flowers slightly
larger in dimension (3.5 cm vs. 2.5 cm across). It also occurs in Mt. Kinabalu at
about 1400-2300 m.
8. Gordonia integerrima {T. & B., Cat. Hort. Bog. (1866) 204. nom. nud.] (Miq.)
H. Keng, comb. nov. Fig. 8
Laplacea integerrima Miq., Mus. Bot. Lugd. Bat. 4 (1869) 113; Back. &
Bakh.f., Fl. Java 1 (1963) 321.
Haemocharis integerrima (Miq.) K. & V. Med. ’S Lands PI. Tuin 16 (1896) 294;
BULK- Je sth. Br. Roy. AS. soc. 76. (1917) 151.
Haemocharis serrata K. & V., op. cit., 296, syn. nov.
Laplacea serrata (K. & V.) Melch. in E. & P., Pflanzenfam. ed. 2, 21 (1925) 136;
Back. & Bakh.f., l.c. syn. nov.
A tree, 18-30 m tall. Bark smooth, dark brown. Branches and branchlets terete,
glabrous. Leaf-blades thin-coriaceous, narrowly elliptic, or oblong-oblanceolate,
apex accuminate, obtuse, rounded or subemarginate, base acute, 7-14 (-16) cm long,
2.5-3.5 (-4.5) cm wide; margin entire, subentire or the upper half (or two-thirds)
finely serrulate, nerves (less clearly defined) about 9-11 pairs, anastomosing and
twining into submarginal network, glabrous on both surfaces; midrib slightly
sunken above, and elevated below; petioles 3-5 mm long, glabrous. Flowers in
upper axils, solitary or two to several together; peduncles 2-4 mm long. Bracts 2-3
on the peduncle slightly below the calyx, early caducous. Sepals 5-6, unequal,
broadly ovate to reniform, pubescent externally. Corolla 2-3 cm across, white;
20 Gard. Bull. Sing. 37(1) (1984)
petals 5-6, broadly oblong to rounded, concave, briefly joined below, varying from
0.6-1.4 cm long, thin-coriaceous. Silvery hairy on the outer surface except the upper
margin which is thin and glabrous. Androecium 5-8 cm long, the filaments shortly
united below. Gynoecium 6-8 mm long; style 3-3.5 mm long, the upper half to two-
thirds 5-branched, glabrous, the lower half of the style and the ovary densely
covered with long yellowish brown hairs; ovary globose, 3-5 mm long and across.
Capsule cylindric-ellipsoid, 5-grooved, sparsely pubescent 2.5-3 cm long, dehiscing
into S-valves; sepals persistent.
Fig. 8. Gordonia integerrima (Miq.) H. Keng comb. nov.
Java. Teysmann s.n. (Herb. Lugd. Bat. 908, 249-234 & 237).
Gordonia in Malesia 21
DISTRIBUTION. Malesia (Java, Bali, Celebes).
Java. Locality unknown, Teysmann s.n. (Herb. Lugd. Bat. 908, 249-237) (lectotype of Laplacea
integerrima Miq., L); G. Poelasari, Banten, Koorders 8202 (isotype of Haemocharis serrata K. & V., L);
Tjibodas, Preanger, Koorders 10018, 15760, 15761, 24352, 37271 (L); Bandung, Seekaria 180 (FRI Ja
7458) (L).
Bali. Collector unknown, Herb. Lugd. Bat. 980, 249-283 (L).
Celebes. Minahassa, Koorders 18099 (L); Menado, Neth. Ind. For. Serv. 18095 (L), 18100 (SING).
ECOLoGy. In primary forests, alt. 1000-1850 m. Fl.: Jan., Jun. & Oct., fl.: Oct.
(one collection).
Note. The name Gordonia integerrima T. & B. appeared first as a nomen
nudum in the unpublished Catalogue of the Botanic Gardens, Buitenzorg. This
name is thus invalid and without nomenclatural status. Miquel adopted the specific
epithet in describing his new species.
An isotype of Koorders & Valeton’s ‘Haemocharis serrata’ has been studied and
found indistinguishable from this species.
9. Gordonia luzonica Vidal, Rev. Pl. Vasc. Filip. (1886) 57; Merr. & Rolfe, Philip.
J. Sc. (Bot.) 3 (1908) 114; Burk., J. Str. Br. Roy. As. Soc. 76 (1917) 149,
f. 8; Philip. J. Sc. 15 (1919) 478; Merr., Enum. Philip. 3 (1923) 70. Fig. 9
Gordonia fragrans Merr. Philip. J. Sc. 1 (1906) Suppl. 95.
G. welbornei Elm. Leafl. Philip. Bot. 2 (1908) 500; Burk., J. St. Br. Roy. As.
Soc. 76 (1917) 150.
G. benguetica Burk. Philip. J. Sc. 15 (1919) 478; Merr. Enum. Philip. 3 (1923)
70. Syn. nov.
?G. subclavata Burk. Philip. J. Sc. 15 (1919) 478; Merr., op. cit. 71.
AGis fo, Wud, WAKES IIGor, Cos Cty le
Small to large tree, 10-50 m. tall. Bark thick, greyish, scaling in plates on old
trees. Branches spreading; branchlets puberulous. Leaf-blades thin-coriaceous or
membranous, elliptic, narrowly obovate or oblanceolate, 5-13 cm long, 2.5-5.5 cm
wide, apex acute or obtuse and abruptly acuminate, sometimes rounded, base
cuneate or attenuate; the upper half or two-thirds finely serrulate or crenulate, entire
near the base; nerves 9-12 pairs, intermingled with less distinct ones; dark green,
glabrous above, lighter, more or less puberulose beneath; petiole 2-5 mm long.
Flowers subterminal or in upper axils, solitary; peduncles 0.5-1 cm long, stout.
Bracts 2-3, caducuous. Sepals 5, broadly ovoid to deltoid, 0.5-1.2 cm long,
sericeous externally. Corolla 7-8 cm across, light yellow or creamy white; petals 5,
broadly oblong or suborbiculate, 3-3.8 cm long, united below, densely covered with
short yellowish hairs throughout the external surface and in the middle basal portion
on the internal surface. Androecium 1.2-1.5 cm long, the filaments puberulous.
Gynoecium 1.2-1.6 cm long, pubescent; ovary ovoid, 6-8 mm long; style approx-
22 Gard. Bull. Sing. 37(1) (1984)
imately the same length of ovary, rather thick, enlarged at the top, 5-lobed. Capsule
broadly cylindric, 3-4 cm long, tapering or nearly rounded above, dehiscing into 5
valves; calyx persistent. Seeds 2.5-3.2 cm long, including the wing.
DISTRIBUTION. Malesia (the Philippines).
Philippines. Luzon, Mt. Mariveles, Bataan, Borden 809 (SING); Baguio, Curran 5083 (isotype of
G. benguetica Burk., SING); Mt. Bulusan, Sorsogon, Edano & Gutierrez 38465 (L), Elmer 16373 )L);
Llavac, Real, Lagrimas 521 (L); Apunan, Meijer 2596 (SING); Panai, Benquet, Santos 31755 (L, SING);
Bontoc to Baguio, Steenis 17973 (L), Mt. Bulusan, Sorsogon, Sulit PNH 2754 (L, SING). Mt. Suiro,
Biliran Isl., Sulit PNH 21481 (L). Negros, Dumaguete, Negros oriental, Elmer 9584 (isotype of
Fig. 9. Gordonia luzonica Vidal
Philippines, Sulit PNH 21481 (fl.); Lagrimas EPRI 521 (fr.).
Gordonia in Malesia 23
G. welborni Elm., L). Mindanao, Mt. Apo, Davao, Elmer 11217 (L); Kulasihan, Lanao, Simajon FB
3063 (L).
EcoLoGy. In primary forests and on exposed ridges. Altitude 600-1300 m.
Fl.: Feb.-Aug., fr.: Apr.-Dec.
Norte. Burkill (1919) examined a loan of Gordonia specimens (under the name
G. luzonica Vidal) from the herbarium of the Bureau of Science, Manila, and found
five different forms which he called ‘species’ with a caution that ‘‘other authors may
perhaps not concede more than varietal rank to one, or possibly even two of them’’.
Most of the specimens he cited were probably destroyed during the War. At
present only one sheet, namely the isotype of G. benguetica Burk, is available to me.
Burkill emphasizes that in G. benguetica the leaves are rounded under the acumen,
about one-half as broad as long, and are broadest at the middle. It is in contrast
with ‘‘G. /uzonica’’ of which the leaves are narrowed to the acumen, about 4 as
broad as long. This does not hold. For example, in E/mer 11217 (L), the leaves are
both acute and rounded at the tip and the length-width ratio varies.
At present, I can only recognise Burkill’s G. polisana as a good species. As to
his G. subclavata (characterized by the subclavate capsule, about 4 cm long, with
the top domed) and Gordonia sp. (characterized by the small entire leaves), I tend
to think they are merely variants of G. /uzonica Vidal.
10. Gordonia maingayi Dyer in Hook.f., Fl. Brit. Ind. 1 (1872) 291; King, J. As.
Soc. Bengal 59, 2 (1890) 204, Ann. Bot. Gard. Calc. 5 (1896) 147, pl. 179A;
Burk., J. Str. Roy. As. Soc. 76 (1917) 147; Ridl., Fl. Mal. Pen. 1 (1922) 204;
H. Keng in Ng, Tr. Fl. Mal. 3 (1978) 285. Fig. 10
A small tree, to 13 m tall. Crown dense; bark brownish or light red brown,
smooth, finely fissured. Young twigs very slender, densely covered with short
yellow-brown hairs; older branches with rough bark glabrescent. Leaf-blades thin-
coriaceous, oblanceolate, ovate or narrowly obovate, apex short and bluntly acute,
base cuneate or attenuate, (4-) 6-8 cm long, 2-3.5 cm wide; drying yellow-brown;
the upper half or two-thirds remotely serrulate, the lower part entire; nerves 5-7
pairs, slightly elevated above, very obscure below; glabrous above, sparsely pubes-
cent beneath; petioles slender, 0.4-1 cm long, pubescent. Flowers subterminal or in
upper axils, solitary or sometimes 2-3 together; peduncles very short (1-2 mm long)
or almost sessile. Perules (bracteoles, bracts and sepals) persistent at anthesis and
forming an involucre about 1-1.5 cm high, densely greyish pubescent externally,
increasing in size from the lower bracts (deltoid, 2-3 mm wide) to the upper sepals
(suborbiculate, 5-7 mm long and wide). Corolla 1.5-2.2 cm across, white (fide van
Balgooy) or yellow (fide Cockburn); petals obovate to orbicular, concave, 0.6-1 cm
long and wide, often emarginate at apex and clawed at base, densely greyish hairy
on the external surface except the margin which is usually thinner and glabrous,
sparsely pubescent on the internal surface. Androecium 6-7 mm long, the filaments
glabrous, briefly connate below. Gynoecium 8-9 mm long; style slender; 4-5 mm
long, glabrous; ovary ovoid, 3-4 mm long, ridged, glabrescent. Capsule cylindric,
24 Gard. Bull. Sing. 37(1) (1984)
2.5-3 cm long, 1.2-1.5 cm across, dehiscing into 4, sometimes 5 valves; sepals
persistent. Seeds 2-2.5 cm long including the wing.
DISTRIBUTION. Malesia (the Malay Peninsula).
Malay Peninsula. Perak, Scortechini 1982 (L, SING), Wray 3766 (SING); Kuala Kangsar, Kochummen
FRI 2461 (L); Bintang Hijau F. R., Chan FRI 13328 (L, SING); nr. Fort Tapong, Whitmore FRI 15749
(L); G. Babu, Se/varaj FRI 6555 (L). Trengganu, Bt. Jebak Puyoh, Ulu Besut, Cockburn FRI 8335 (L).
Negri Sembilan, Nilai Jindaram Estate, Md. Shah 70 (SING). Pahang, Sg. Telom, Bt. Cheraya, Sohadi
FRI 14731 (L, SING); Taman Negara, van Balgooy 2577 (L). Kelantan, G. Babong, Soepadmo &
Mahmud 1172 (L). Malacca, A. C. Maingay 192 (L, isotype).
VERNACULAR NAMES: fitup, titup tiup, tiup, damak.
ECOLOGY. In primary forest and dipterocarp forest, on ridge-top or on hill side;
alt. 300-400 m. Fl.: Apr.—May, fr.: July-Jan.
11. Gordonia marginata (Korth.) Endl. ex Walp., Repert. 5 (1845) 134; Merr., J.
Str. Br. Roy. As. Soc. spec. no. (1921) 390; Masamune, Enum. Phan. Born.
(1942). 472. Fig. 11
Closaschima marginata Korth., Kruidk. (1842) 141 (incl. var. dasyophthalma
Korth.); Walp., op. cit. 1 (1842) 375.
Laplacea marginata (Korth.) Choisy, Mém. Soc, Phys. Hist. Nat. Geneve 14
(1855) 148; Miq., Fl. Ind. Bat. 2 (1857) 490.
Haemocharis marginata (Korth.) O. Ktze, Rev. Gen. Pl. (1891) 62; Burk., op.
Clits SSe
Tree, 25-40 m tall, buttressed. Bark grey, smooth, scaling off in large pieces;
living bark beefy red, sap wood reddish white. Young twigs reddish brown,
glabrous. Leaf-blades thin coriaceous, obovate or narrowly obovate, sometimes
rhomboid, obtuse or slightly emarginate, base cuneate, 4-7 (-9) cm long, 2-3
(-3.5) cm wide; margins for the most part undulate or remotely crenulate, entire near
the base; nerves 5-7 pairs, slightly impressed above, inconspicuous below, glabrous
on both surfaces, except the lower pilose midrib; petioles stout, 2-3 mm long.
Flowers in upper axils or subterminal, solitary or 2-3 in clusters; peduncles very
short, 1-3 mm long; bracteoles, bracts and sepals forming an involucre about
5-8 mm high, silvery puberulous on the back, increasing in size from the lower
bracts (broadly ovate, 1-1.5 mm long) to the upper sepals (broadly ovate or
subrounded, 4-5 mm long). Corolla 2-2.5 cm across, pale yellow; petals 5, obcor-
date to subrounded, 8-10 mm long, membranous, silky externally except the
margins which are thinner and glabrous; petals briefly joined at the base and adnate
to the filaments. Androecium 4-5 mm long, the filaments united below. Gynoecium
4-6 mm long; columnar, 1.5-2.5 mm long, ridged, the upper portion separating into
5 branches each bearing a terminal stigma; ovary ovoid, velutinous, tapering above
into the style. Capsule ellipsoid-cylindric, 2-2.5 cm long, 5-grooved, velutinous
along the grooves; sepals persistent. Seeds 1-1.4 cm long including the wing.
Gordonia in Malesia 25
DISTRIBUTION. SE. Borneo (Kalimantan).
SE. Borneo. Without precise locality, Korthals, s.n. (several collections, Herb. Lugd. Bat. 908,
249-245 selected as lectotype of C/osaschima marginata Korth.); G. Pamatton, Poelau Lampei, Korthals
s.n.; Martapoera, Korthal s.n.; G. Tirin, Korthals, s.n. (all from L.); Tanjong Bangko, near the mouth
of Mahakam River, alt. 20 m, Kostermans 7049 (BO, L); West of Samarinda, Kostermans 6702 (BO, L).
EcoLoGy. Sandy ridges, on loam soil; alt. 30-300 m. Fl. & fr.: Apr.-Aug.
VERNACULAR NAME: penagit.
NOTE. Over a dozen specimens collected by Korthals during his Borneo trip
(Oct.—Dec. 1836) are deposited in the Leiden Rijksherbarium. They were variously
10 {|
Fig. 10. Gordonia maingayi Dyer
Malaya, Wray 3766 (fr.); van Balgooy 2577 (fl.).
Fig. 11. Gordonia marginata (Korth.) Endl. ex Walp.
Kalimantan, Korthals s.n. (Herb. Lugd. Bat. No 925, 250-547); Kostermans 6702 (fl., fr.).
labelled as Closaschima (or as Closaschyma) marginata, C. obovata, C. lanceolata,
and C. marginata var. dasyopathalma, all in Korthals’ handwriting. Only the first
name was published. These specimens bear small flower buds or are sterile. Among
them there is only a single fruit (Herb. Lugd. Bat. 908, 249-238). The description
presented here is largely based on two collections of Dr. Kostermans.
12. Gordonia multinervis King, J. As. soc. Bengal 59, 2 (1890) 205; Burk., J. Str.
Br. Roy. As. Soc. 76 (1917) 154, f. 77; Ridl., Fl. Mal. Pen. 1 (1922) 203;
H. Keng in Ng, Tr. Fl. Mal. 3 (1978) 286. Fig. 12
Gordonia concentricicatrix Burk., op cit., 153, f. 1/0 & 11, op. cit., 78 (1918)
26
Gordonia in Malesia 27
49, pl. 3; Ridl., l.c.; H. Keng, op. cit., 285. Syn. nov.
Small to large tree, 10-30 m tall. Bark grey, smooth or reddish brown and with
large loose scales in several! layers, scallop-marked. Young twigs glabrous except
near the tip which is puberulous. Leaf-blades membranous or thin coriaceous,
obovate-spathulate, apex rounded, mucronate, or shortly acuminate, base gradually
attenuate and often narrowly winged below, varying from 10-15 cm long, 2-5.5 cm
wide (‘concentricicatrix’) to 12-20 cm long, 6-8.5 (-10) cm wide (‘multinervis’), the
upper two-thirds crenate, subentire or entire below; glabrous on both surfaces;
nerves 8-18 pairs (often with smaller, less conspicuous veins in between), very faint
above, barely visible beneath, fused near margin and forming intramarginal
reticulations; midrib sulcate above; petiole thickened, 2-5 mm long, glabrescent.
A
Fig. 12. Gordonia multinervis King
Malaya, Watson CF 878 & Abdul Rawi CF 878 (fl. & fr.); Scortechini 1968 (large leaf).
28 Gard. Bull. Sing. 37(1) (1984)
Flowers axillary, solitary; peduncles short, 2-5 (-10) mm long, stout. Bracts 2-3,
broadly deltoid, 3-5 mm long. Sepals 5-6, broadly ovate to suborbicular, thin
coriaceous, about 1 cm long and wide, greyish silky externally. Corolla 2-3.5 em
across, yellow; petals 5, broadly obovate or suborbicular, 1-1.7 cm long and wide,
thin coriaceous, greyish silky externally (indistinguishable from sepals from without
except by larger size and position) briefly clawed and joined at base, and adnate to
the filaments. Androecium 1.2-1.5 cm long, the filaments united below. Gynoecium
1-1.2 cm long; style solitary, columnar, 4-5 mm long, hispid, the tip discoid,
5-lobed into 5 stigmas; ovary ovoid, 3-4 mm long, densely covered with sericeous
hairs. Capsule bluntly 5 angulate, 3-4 cm long, 5-valved; sepals persistent. Seeds
2-2.5 cm long including the wing.
DISTRIBUTION. Malesia (the Malay Peninsula).
Malay Peninsula. Kelantan, G. Stong, Whitmore FRI 12490 (L); Kuala Kerbat, Whitmore FRI 20244
(L). Perak, Scortechini 1968 (isotype, SING). Pahang, Chini F.R., Cockburn FRI 11076 (L). Selangor,
Rantau Panjang, Watson CF 878 & Abdul Rawi CF 878 (Type of G. concentricicatrix Burk., SING);
Gading F.R., Loh FRI 13397 (L). Negri Sembilan, G. Angsi, Loh FRI 17314(L). Johore, Bukit Tingan
Laut, Corner SFN 37064 (L); Tenggaroh F.R., Ogata KEP 105152 (L), G. Ledang, Whitmore FRI 12303
(L). Singapore, Bukit Timah, Corner SFN 36435 (L); MacRitchie Res., Sinclair SFN 40231 (L).
EcCOoLoGy. In lowland forest, on sandstone ridge or on hill side; alt. 10-800 m.
Fl.: Apr.-June, fr.: July—Oct.
VERNACULAR NAMES: kelak merah. samak pulut. samak samak.
NOTE. This species, as observed by Steenis (Blumea 12 (1964) 319), is closely
allied to G. oblongifolia of Sumatra; both are characterized by the oblong or
obovate leaves with decurrent winged base, by the thin leathery petals with silky
outer surface, hardly distinguishable from the sepals before anthesis, and by the
single style with an enlarged discoid tip. It can be easily separated from the latter,
however, by its smaller leaves with more numerous nerves and more distinct
submarginal reticulation and by its much smaller flowers and fruits.
G. concentricicatrix Burk. was described from a small-leaved form of the species.
Of this the type specimen is a mixture of two specimens collected by two different
persons at the interval of one year but were mounted on one sheet and given the
same collection number.
The following two Bornean specimens are probably referable to this species:
E. Borneo, Mt. Palimasan alt. 500 m, Kostermans 13097 (L), and N. Borneo,
Beaufort Hill, alt. 350 m, Lajangah 44532 (L).
13. Gordonia oblongifolia (Miq.) Steenis, Blumea 12 (1964) 319. Fig. 13
Ploiarium? oblongifolium Miq., Fl. Ind. Bat. Suppl. (1861) 483.
Tree, 13 m tall. Branches stout, glabrous. Leaf-blades thin coriaceous, elliptic-
oblong, narrowly oblong or oblanceolate-spathulate, apex broadly acute or obtuse,
base acute or attenuate, decurrent and narrowly winged, 14-20 (-28) cm long, 6.5-7
Gordonia in Malesia 29
(-10) cm wide; margin finely crenulate-serrulate for the most part, nearly entire
toward the base; nerves 11-14 pairs, intermingled with less distinct ones and in-
tricately interlocked near the margin; glabrous on both surfaces; petiole 3-5 mm
long, thickened. Flowers solitary, in upper leaf-axils, sometimes several together
near the top of a top branchlet; peduncle 1.5-2 cm long, very stout. Bracts and
bracteoles 2-3, caducous. Sepals 5, broadly ovate, 1.8-2.2 cm long, thick cor-
iaceous, densely sericeous externally, yellowish brown. Corolla unknown.
Gynoecium about 1.8 cm long; style about 1.4 cm long, thinly hispid, minutely
5-lobed at the end; ovary ovoid, about 4 mm long, sulcate, densely sericeous.
Capsule cylindric, S-angulate, about 5 cm long; calyx persistent. Seed 3.5 cm long
including the wing.
Fig. 13. Gordonia oblongifolia (Miq.) Steenis
Sumatra, Neth. Ind. For. Serv. 2794 (fl.), 9571 (fr.).
30 Gard. Bull. Sing. 37(1) (1984)
DISTRIBUTION. Malesia (Central & W. Sumatra).
Sumatra. Benkoelen, Lebong, Neth. Ind. For. Serv. 2794, 957] (L); Res. Lum, Oud. Agam, Olivier
16 (B), Neth. Ind. For. Serv. 2946 (L). Res. Tapanoeli, Sibolga, Neth. Ind. For. Serv. 3780 (L). Fort
de Kock, nr. Bukit Silit, Teysmann HB 668 (L, type of Ploiarium oblongifolium Miq.)
EcOoLocy. In primary forest; alt. 50-1300 m. Fl.: Jan. (one collection), fr.: Dec.
(one collection).
Fig. 14. Gordonia ovalis (Korth.) Korth. ex Walp.
Sumatra, Korthals, s.n. (Herb. Lugd. Bat. No. 908, 249-324, fr); Jacobs 8311 (fl.).
Gordonia in Malesia 31
VERNACULAR NAMES: kajoe patjat, ubar lilim, djirok bantjoh (Sumatra).
Note. The original description of Ploiarium oblongifolium Miq. was based on
a sterile specimen (Teysmann HB 668). Miquel therefore was uncertain about the
generic status. In fertile material, van Steenis found the style bearing minute
stigmas, and therefore tranferred it to Gordonia. A well-preserved fruiting specimen
(Neth. Ind. For. Serv. bb. 9571) further confirms this transfer.
This species, although with large elliptic-oblong leaves, showy flowers and big
fruits, is poorly represented in herbaria. I failed to find any Sumatran specimens
collected later than 1925 referable to this plant.
14. Gordonia ovalis (Korth.) ex Walp. Repert. 5 (1845) 134 Fig. 14
Closaschima ovalis Korth., Kruidk. (1842) 140, ¢, 28.
Laplacea ovalis (Korth.) Choisy, Mém. Soc. Phys. Geneve 14 (1855) 148;
Melch. in E. & P., Pflanzenfam. ed. 2, 21 (1925) 136.
Haemocharis ovalis (Korth.) O. Ktze, Rev. Gen. 1 (1891) 63; Burk. J. Str. Roy.
As. Soc. 76 (1917) 158.
Laplacea aromatica Migq., Fl. Ind. Bat. Suppl. (1861) 482, (incl. var. minor, var.
longifolia), Mus. Bot. Lugd. Bat. 4 (1869) 114; Melch. in E. & P. Pflanzen-
fam. ed. 2, 21 (1925) 136. Syn. nov.
Haemocharis aromatica (Miq.) Szyszyl. in E. & P. Pflanzenfam. 3, 6 (1893) 185
(excl. syn. L. semierrata Miq.); Burk., op. cit., 151. Syn. nov.
Laplacea subintegerrima Miq. Fl. Ind. Bat. Suppl. (1861) 483. Syn. nov.
Haemocharis subintegerrima (Miq.) Burk., op. cit., 155. Syn. nov.
Tree, 10-25 m tall. Bark grey, peeling off in pieces. Young branches slender,
thinly covered with short silky hairs; older branches puberulent or glabrous. Leaf-
blades thin or thick membranous, narrowly elliptic, narrowly ovate or obovate,
sometimes slightly asymmetrical; apex acute, acuminate or subcaudate, sometimes
obtuse or rounded; base attenuate or cuneate, 4.5-9 (-14.5) cm long, 2-2.5 (-4) cm;
wide; margin finely serrulate-crenuate; nerves 10-13 pairs, faint on both surfaces,
shining dark green and glabrous above, light green, slightly glaucous and sericeous
below; petiole very short, 2-3 mm long. Flowers usually in upper axils, solitary,
peduncles 1-3 mm long, sericeous. Bracts and sepals 7-8, silver puberulous ex-
ternally, increasing in size from the lower bracts (deltoid or reniform, 1-1.2 mm
long) to upper sepals (reniform or subrounded, 2-3 mm long). Corolla 1.2-2.5 cm
across, white (fide Kostermans & Anta); petals usually 5, oblong, broadly oblong
to suborbicular, 5-8 mm long, thin coriaceous, silvery puberulous on the external
surface except the margins which are thin and glabrous, briefly joined below.
Androecium 4-5 mm long, the filaments in several whorls, united below and adnate
to the corolla. Gynoecium ovoid to globose, 2-2.5 mm across, sericeous; style
absent, only 5 very short, V-shaped protrusions (less than 1 mm long) representing
the stigmas; ovary shallowly longitudinally ridged. Capsule 2-2.5 cm long, ovoid or
narrowly ovoid, thinly puberulous, dehiscing into 5 valves; sepals caducous fully
ripe fruit; seeds 1.5-2 cm long including the narrow wing.
32 Gard. Bull. Sing. 37(1) (1984)
DISTRIBUTION. Malesia (Sumatra).
Sumatra. W. Sumatra, Korthals s.n. Herb. Lugd. Bat. 908, 249-247, -248, -249, -284, -305, -311, -312,
-313, -314, -315, -318, -319, -320, -321, -322 (holotype of Closaschima ovalis Korth., L.), -323, -324, -474
(L); Pariamen, W. Sumatra, Diepenhorst H.B. 3081 (isotype of Laplacea aromatica Miq., L), 2184
(lectotype of L. aromatica var. minor Miq.), 2492 (lectotype of L. aromatica var longifolia Miq., (L).
Lampung, NW. of Kota Agung, Jacobs 8311, 8439 (BO, L). Asahan, Krukoff 4224, 4235 (BO), Rahmat
Si Boeea 9346 (L). Bangka Lubok Besar, Kostermans & Anta 540 (BO, L). Palembang, Neth. Ind. For.
Serv. bb. 938, 32002, 31720 (L), Teysmann H.B. 3969 (holotype of Laplacea subintegerrima Miq., L).
ECOLOGY. In primary forest, on ‘red’ soil. Alt. 20-500 m. Fl.: Apr, May &
Sept., fr.: Apr.-May, & July.
Fig. 15. Gordonia penangensis Ridl.
Malaya, Curtis 834 (isotype).
Gordonia in Malesia 33
VERNACULAR NAME: palembang putih.
Note. This plant was first collected by P. W. Korthals from the forests of
Melintang Mountains (near Padang) between April and May, 1834 during his trip
to W. Sumatra. Over a dozen sheets of the same plant (including flowers buds,
flowers and a single fruit) are preserved in the Leiden Herbarium to-day. Korthals
was probably fascinated by the small flowers with an ovoid ovary totally devoid of
styles. In some of these specimens a tiny sheet of paper with hand-written ‘Gordonia’
was attached. This was possibly done in the field. Later he described this plant under
a new generic name Closaschima.
Miquel’s Laplacea aromatica was based on H. Diepenhorst’s collection from
Priaman, very near the area where the Korthals’ collection was made. Miquel
compared his ‘new species’ with Laplacea vulcanica (= G. vulcanica) but failed to
do so with Korthals’ Closaschima ovalis, with which it agrees in almost every aspect.
Miquel also proposed two new varieties of this species: var. /ongifolia has longer
(10-12.5 cm long) lanceolate-oblong leaves, and var. minor has smaller, elliptic-
lanceolate leaves. The leaves of var. minor match closer the type specimens of
Korthals.
Szyszylowicz (1893) transferred Laplacea aromatica Miq. to Haemocharis; at the
same time, he cited L. semiserrata Miq. as a synonym. This was obviously a mistake,
as Miquel never described a species under such a name which is in fact a New World
one, namely, Laplacea semiserrata Cambess. from Brazil.
There are two distinct forms (which probably warrant infraspecific status) in
Gordonia ovalis. In one form the leaves are generally smaller (4.5-7 cm long), thin
membranous and their leaf-apices usually gradually taper to a blunt point; this
includes the type specimens of Closaschima ovalis Korthals and Laplacea aromatica
Miquel, and most specimens were collected from Pariamen-Padang area in the
south. In another form the leaves are usually larger (6-9 cm long), thin-coriaceous
and their leaf-apices are obtuse or rounded; this includes the type specimens of
Laplacea subintegerrima Miq. and others from the Palembang area and Bangka
Island in the northeast. There are no obvious differences in their flower and fruit
structures. In both forms, in the styles are almost totally absent, with five stigmas
lying on top of the hairy ovary.
15. Gordonia penangensis Ridl. J. Str. Br. Roy. As. Soc. 73 (1916) 142, Fl. MI.
Pen. 1 (1922) 203; H. Keng in Ng, Tr. Fl. Mal. 3 (1978) 286, f. 4.
Fig. 15
Gordonia excelsa auct. non Blume: King J. As. Soc. Bengal 59, 2 (1890) 203.
A small, slender tree, 10-13 (-20) m tall. Bark smooth, pale grey brown; young
twigs very slender, short-silky pubescent, greyish; older branches less hairy,
brownish. Leaf-blades thin coriaceous, narrowly elliptic-lanceolate, apex acuminate
or more frequently obliquely caudate, base attenuate or cuneate, 6-10 cm long,
2.5-4 cm wide, the upper half or two-thirds remotely serrulate or almost entire;
midrib sulcate above and elevated below; nerves about 5-8 pairs, very faint above,
barely visible beneath; drying light olive green; glabrous above, somewhat greyish
34 Gard. Bull. Sing. 37(1) (1984)
and tomentose below; petiole slender, 0.5-1 cm long, silky, brownish. Flowers
solitary, terminal or subterminal on small branches; peduncle 2-3 mm long, silky
tomentose. Bracteoles 2, triangular-cordate, 2-3 mm long, caducous. Sepals 5,
ovate or suborbicular, thick coriaceous and with thinner and ciliate edge, silvery
pubescent at the back, 5-7 mm long and wide. Corolla 2.5-3 cm across, yellowish
to golden yellow; petals 5, suborbicular to spathulate, chartaceous, fused below.
Androecium 5-6 mm long, the filaments joined into a short tube and briefly adnate
to the corolla, more or less in 5 bundles. Gynoecium 9-11 mm long; style solitary,
slender glabrescent, 6-7 mm long, the upper portion enlarged into club-shaped
stigmas; ovary spherical, 3-4 mm across, woolly. Capsule cylindrical, 3.5-S5 cm
long, 1.5-2 cm across, glabrescent, 5-valved. Seeds 2.5-3 cm long including the
wing.
Fig. 16. Gordonia polisana Burk.
Philippines, Ce/estino 695 (f1.); Jacobs 7397 (tr.).
Gordonia in Malesia 35
DISTRIBUTION. Malesia (the Malay Peninsula).
Malay Peninsula. Penang, Penang Hill, Curtis 834 (isotype, SING), Ng FRI 1054 (L). Perak, Wray s.n.
(SING). Pahang, Cameron Highlands, Md. Nur SFN 32948 (SING); Whitmore FRI 15882 (L). Johore,
Tg Penawar, Cockburn FRI 7642 (L); Kota Tinggi, Stone & Anderson 8714 (L). Singapore, Seletar, Ridley
6214, 3913 (paratypes, SING); Sinclair SFN 39585 (L), Jumali 1048 (SING).
EcOLoGy. In open lowland forests, alt. 20-500 m. Fl.: Feb-May, fr.: May-June.
16. Gordonia polisana Burk., Philip. J. Sc. 15 (1919) 478; Merr., Enum. Philip. 3
(1923) 71. Fig. 16
Shrub or small tree, 8-15 m tall. Branchlets slender, glabrous. Leaf-blades thin-
coriaceous, obovate or narrowly obovate, apex obtuse or rounded, often shallowly
retuse, base acute or attenuate, 4-7 cm long, 2.5-3.5 cm wide, the upper part
remotely crenulate-serrulate, the lower entire, nerves 12-14 pairs, not distinct;
glabrous on both surfaces except a few scattered hairs on the midrib beneath;
petiole 2-5 mm long. Flowers in upper leaf-axils, solitary; peduncle about 0.5 cm
long. Bracteoles about 2. Sepals 5-6, broadly orbiculate or reniform, 0.8-1 cm long,
densely yellowish silky externally. Corolla 7-8 (-14?) cm across, white (fide
Celestino); petals 8-10, in two series, united below, densely covered with short
yellowish hairs over the external surface and in the middle-basal portion on the
internal. Androecium 1.2-1.5 cm long, the filaments puberulous. Gynoecium about
2 cm long, pubescent; ovary ovoid, about 1.5 cm long; style about 0.5 cm long, very
thick and stout, the top enlarged and shallowly lobed into stigmas. Capsule broadly
cylindric, 3.5-4 cm long.
DISTRIBUTION. Malesia (endemic to the Philipines; Luzon).
Luzon. Mt. Polis, Ifugao, Mountain Province, M. Celestino 695 (PNH 8021) (L). Mt. Pulog, Jacobs
7397, 7421 (L).
ECOLOGY. In mossy forest, above stream; alt. 1700-2300 m. Fl. Feb.-Mar. (two
collections), fr.: Feb. (one collection).
Note. Among the 4 new species described by Burkill from the Philippines, this
is undoubtedly the most outstanding. The type specimens (A/varez FB 18384 and
Sandkuhl 316), both collected on Mt. Polis in Bontoc Sub-province, were not
available for this study. Among a limited number of Philippino specimens at my
disposal, I was only able to identify the above-cited ones as belonging to this species.
Burkill pointed out that (1) the leaves of this plant are rounded under the acumen,
broadest above the middle; (2) the flowers are very large, about 9 cm (or according
to Sandkuhl, 11-14 cm) in diameter; and (3) the capsule is about 4 cm long, long-
tapering above.
The leaves of Celestino 695 fully agree with Burkill’s description. This bears two
fully expanded flowers, the larger one measures about 8 cm in diameter. One
unusual feature of the flowers, which Burkill failed to mention is that the petals are
36 Gard. Bull. Sing. 37(1) (1984)
8-10 in number and are arranged in 2 series. Petals of the outer series are
intermediate between the largest sepals of the calyx and the smaller petals of the
inner series.
17. Gordonia sarawakensis H. Keng sp. nov. Fig. 17
Arbor ca. 30 m alta. Folia angusto-lanceolata vel angusto elliptica, 12-15 cm long, acuminata vel
breviter obtusa, basi cuneata, coriacea, nervis lateralibus 20-35 bene intra marginem anastomosantibus,
pagina utrinque suborscuris, petiolo 3-5 mm longo. Flores flavi (fide Banyeng & Benang), axillares,
solitarii, 5-6 cm diametro; bracteae 2-3, decicuae. Sepala coriacea, ovato-rotundata, 1-1.3 cm longa,
dorso adpresse pubescentia. Petala late obovata vel lineario-lanceolata, 1.5-2.4 cm longa. Gynoecium 4.5
cm longum; stylo 0; stigmate 8-10. Capsula late ovoidea, 3-3.5 cm longa, valvis 8-10. — Typus,
Sarawak, Banyeng & Benang S252/8, in L.
A large tree, 30 m tall. Bark flaky. Young twigs with scattered short hairs; older
branches stout, greyish brown glabrous. Leaf-blades coriaceous, narrowly ovate or
narrowly elliptic, acuminate or shortly obtuse, base attenuate, very narrowly
winged, 12-15 cm long, 5-6 cm wide; margin crenulate or undulate; midrib elevated
beneath; nerves 18-25 pairs (often with lesser ones between) fused near the margin and
forming a submarginal vein, faint on both surfaces; drying blade dark green or
brown, with scattered short hairs on both surfaces; petiole very thick, 2-3 mm long.
Flowers axillary, solitary, peduncle very stout, 4-5 mm long; bracts 2-3, deltoid-
caudate, 4-5 mm long, caducous. Sepals 5, broadly ovate or suborbicular, 1-1.3 cm
long and wide, thin-coriaceous, greyish silvery externally. Corolla 5-6 cm across
yellow (fide Banyeng & Benang) or yellowish white (fide Bujang); petals 7-8,
arranged in one series, varying from broadly obovate or obovate (1.8-2.0 x 1.4-1.8
cm) to narrowly obovate or linear lanceolate (1.8-2.4 x 0.8-1.2 cm), the exposed
part in bud thick and silvery hairy externally, other parts thinner and glabrous.
Androecium 5 mm long, the filaments glabrous, connate below and adnate to
corolla. Gynoecium 4-6 mm long; ovary subglobose, hirsute; style absent or very
short; stigmas 8-10, lying on top of the ovary. Capsule broadly ovoid or subglobose,
3-3.5 cm long, 2.5-3 cm across, dehiscing into 8-10 valves; short septicidal lines also
developed near the base of the fruit. Seeds 2-2.2 cm long including the wing.
DISTRIBUTION. Malesia (Borneo: Sarawak & Sabah).
Sarawak. Kuching, 12th mile, Penrissen Road, Banyeng & Benang S 25218 (holotype, L); Semengoh
F.R., Rosli S 14983 (L, SING), S 15192 (L), Hj. Bujang S 32444 (L), S 32957 (L, SING), Galau S 15738
(L), Zen 10031 (SING).
Sabah. Bukit Hampuan alt. 1300 m., Ranau Aban Gibot 61804; G. Lotung Inarat alt. 1700 m. Lamag,
Aban Gibot SAN 83233; Mt. Silam, Lahad Datu, Meijer & Anak SAN 37489, Mujin 37816; along Labuk
Road, Sandakan, Mikil 46699 (all in SAN).
ECOLOGY. In lowland dipterocarp forest, alt. 10-100 m. or in hill forest, alt.
1300-1700 m. Fl.: July-Oct., fr.: Oct.-Nov.
VERNACULAR NAME: entuyut.
Note. Superficially this species resembles G. multinervis King of Malaya, but
differs from the latter in detailed structures of the flower and fruit. This species
Gordonia in Malesia 37
possesses 7-8 petals, more or less arranged in one whorl, (at least as seen in a fully
expanded flower); the stamens are comparatively short and fewer in number; the
gynoecium is devoid of styles, thus the 8-10 V-shaped stigmas are lying on top of
the ovary which is 8-10 locular. The fully developed fruit is a nearly spherical capsule
dehiscing loculicidally into 8-10 valves. Short septicidal lines (one each along the
dorsal line of the capsule-valves) present near the base of the fruit. The combination
of these features makes this species one of the most outstanding in the genus.
At first, all the available specimens referable to this species were collected from
the lowland forest near Kuching, Sarawak. Later five specimens were found in a
loan from Sabah Forest Department; they match well with the Sarawak material,
except Aban Gibot 61804 & Mujin 37816, both flowering; in these, a short style (2-3
mm long) is present.
Vi
BOE.
RSNA
Fig. 17. Gordonia sarawakensis H. Keng sp. nov.
Sarawak, Banyeng & Benang S 25218 (holotype) (fl.); Galau S 15738 (fr.).
38 Gard. Bull. Sing. 37(1) (1984)
18. Gordonia scortechinii King, J. As. Soc. Bengal 59, 2 (1890) 204; Burk., J. Str.
Br. Roy. As. Soc. 76 (1917) 158; Ridl., Fl. Mal. Pen. 1 (1922) 204; H. Keng in
Ng, Tr. Fl. Mal. 3 (1978) 286. Fig. 18
A tree to 18 m tall; bark smooth, brown. Young twigs slender, dark brown or
greyish, puberulent; older branches glabrous. Leaf-blades thin-coriaceous, elliptic
or narrowly elliptic, or sometimes narrowly obovate, apex obtuse or bluntly acute,
base cuneate or obtuse, 6-9 cm long, 2.5-3.5 cm drying olive green; margin of the
upper half or two-thirds remotely undulate or subentire; nerves 6-8 pairs, faint
above, almost invisible below; glabrous on both surfaces, shining green above,
subglaucous beneath; petiole 3-5 mm long, puberulent. Flowers subterminal and
Fig. 18. Gordonia scortechinii King
Malaya, Scortechini 362 b (isotype) (fl.); Chelliah FRI 6545 (fr.).
Gordonia in Malesia 39
axillary, solitary; peduncles subsessile or very short (2-3 mm long). Bracteoles 2.
Sepals about 5, cordate-deltoid, 2-3 mm long and wide. Corolla 1.5-1.8 cm across,
butter yellow; petals 5, ovate or suborbicular, strongly concave, 8-9 mm long and
wide, thin-chartaceous, sparsely silvery puberulous externally, briefly joined at base
and adnate to the filaments. Androecium 6-8 mm long; the filaments glabrous,
briefly united below. Gynoecium 5-6 mm long; styles 3 (or 4?), free to base,
spreading; stigmatic surface inside the tip of styles; ovary ovoid densely strigose.
Capsule 1.8-2 cm long, glabrous, dehiscing into 3 (or 4?) valves; sepals caducous.
Seeds 1.2-1.4 cm long including the wing.
DISTRIBUTION. Endemic to the Malay Peninsula. (Perak, Kelantan Trengganu,
Selangor?)
Malay Peninsula. Perak, without locality, Scortechini 362 b (isotype, SING) Kelantan, Whitmore FRI
20647 (SAN). Trengganu, Bukit Rauk F.R. Dungun, S. Chelliah FRI 6545 (L).
EcoLoay. In hillside primary forest. Fl.: Mar. (one collection), fr.: Nov. (one
collection with two specimens).
NOTE. King (I.c) points out that this species superficially resembles G. maingayi,
but it has smaller flowers with fewer stamens (about 30) and a 3-locular ovary with
3 free styles. Burkill (l.c.) states that this species ‘‘might be described a G. imbricata
with an admixture of G. maingayi. Its branches however are more slender than the
first of these two and its flowers are recorded as remarkably small, the stamens
being not more than 30.’’
For years, only the original collection (Scortechini 362 b) was known. During the
course of this study, the other specimen which can positively be identified as this
species in Chelliah FRI 6545 from Trengganu. It has a small 3-valved capsule and
agrees closely with the type specimen in all aspects of vegetative characters except
that the texture of leaves appears slightly thicker. Besides, two other collections
possibly also represent this species: (1) Perak, Piah F.R. Ja’mat FMS 3931 (SING),
several small flower buds being present (dissection of one large bud revealed
numerous perianth-lobes without any sign of reproductive organs in the centre and
is likely to be a gall flower); (2) Selangor, Klang Gates quartz Ridge, Stone 12094
(L), the small capsules being 4-valved.
19. Gordonia singaporeana Wall. [Cat. no. 1457 (lith. 1829) nom. nud.; Ridl. J.
Sabre. As. Soc. 73(1916) 141, nom., Burk., op: cit., 154, f. 1, 2,3, 12,
& 13, nom.] ex Ridl. Fl. Mal. Pen. 1 (1922) 202; Corner. Ways. Tr. 2nd ed.
(1952) 629, pl. 186, f. 236; H. Keng in Ng, Fl. Mal. 3 (1978) op. cit., 286.
Fig. 19
Gordonia grandis King J. As. Soc. Beng. 59, (1890) 203, non André (1880).
Gordonia excelsa Bl. var. sincapuriana Dyer in Hook. f., Fl. Brit. Ind. 1 (1872)
291.
A tree up to 32 m tall. Bark black, scaly, Young twigs puberulous or glabrous.
Leaf-blades thin coriaceous, elliptic or oblong-lanceolate 7-12 (-15) cm long, 2.5-5
40 Gard. Bull. Sing. 37(1) (1984)
(-8) cm wide, apex subrounded or abruptly acuminate, base acute or attenuate,
slightly decurrent or not; margin crenulate-serrate or undulate, usually entire pear
the base; nerves 10-12 pairs, merged and looped near the margin into submarginal
reticulations, barely visible above, even less conspicuous below; glabrous on the
upper surface, short appressed pubescent beneath; petiole 2-5 mm _ long,
puberulous. Flowers in upper leaf-axils, usually solitary; peduncles 0.3-5 (-0.75) cm
long, stout, pubescent. Bracts and bracteoles about 3, caducous. Sepals 5-6,
cordate, subrounded or reniform, often notched above, 6-8 mm long, densely
covered with greyish hairs externally and also near the base on the inner surface.
Corolla 4-5 cm across, cream white, scentless; petals 5-6, broadly oblong or obcor-
date, 2-2.4 cm long, sericeous on the external surface, scattered puberulous on the
internal. Androecium about 1 cm long, the filaments glabrous, connate below.
Fig. 19. Gordonia singaporeana Wall. ex Ridl.
Singapore, fresh material.
ne
Gordonia in Malesia 4]
Gynoecium 6-8 mm long, the ovary ovoid, 4-5 mm long, densely covered with long,
yellowish grey hairs, tapering above to form a stout style which branches into five
for about half its length. Capsule ovoid cylindric, 3-3.5 cm long, dehiscing into 5
valves; seeds 2-2.5 cm long including the wing.
DISTRIBUTION. Malesia (the Malay Peninsula).
Malay Peninsula. Penang, Government Hill, Burkill 2891, Curtis 2281, Philip CF 1002, Ridley 7963.
Selangor, Fraser’s Hill, Md. Nur 11174, 11454. Malacca, Alvins s.n. in 1886, Curtis 3488, Derry 976,
Maingay 191, 1072, Ridley 976. Johore, Mawai, Corner 29253, Ngadiman SFN 34757. Singapore, Corner
33564, Ngadiman SFN 35000, 34923, Ridley 1946, 3812, 4564, 4801 (all in SING, except Maingay 1072,
Ridley 976, 481, L).
ECOLOGY. In primary and mature secondary forests; altitude 50-1300 m. FI.:
May, June & Oct.; fr.: Aug., Nov. & Dec.
VERNACULAR NAMES: kayu kelat asam, kayu kelat putih, sawak pulot.
NOTE. Corner (I.c.) observed that this tree is nocturnal, the flowers open at dusk
and fall next morning. He also noted that the flowering is seasonal and occurs more
than once in a year, perhaps after dry weather.
20. Gordonia taipingensis Burkill, J. Str. Br. Roy. As. Soc. 76 (1917) 148, f. 6;
Ridl., Fl. Mal. Pen. 1 (1922) 204; H. Keng in Ng, 288. Fig. 20
Small to medium-sized tree, 12-16 (-20) m tall. Young twigs covered with short
hairs. Leaf-blades membraneous or thin coriaceous, elliptic to elliptic-oblong, apex
acuminate, obtuse or briefly caudate, base cuneate or attenuate, (8-) 13-20 (-30) cm
long, (3.5-) 4.5-7.5 cm wide; margin of the upper two-thirds remotely crenulate,
entire below; nerves 10-11 pairs on each side, slightly elevated above, faint and
inconspicous below; dark green and glabrous above, with scattered strigose hairs on
the midrib beneath, otherwise glabrous; petiole 1.5-2 cm long, with scattered short
hairs. Flowers axillary, solitary; bracteoles and bracts about 3, caducous; peduncle
0.2-0.5 cm long. Sepals ovate, broadly ovate to suborbicular, 1.2-1.7 cm long and
broad, coriaceous, sericeous on both surfaces, but more densely hairy externally.
Corolla 5-6 cm across, yellowish; petals 5-6, broadly oblong, suborbicular to
reniform, 2.5-3 cm long, the exposed portion of the three outer petals thick and
densely sericeous, the covered portion and the two inner petals membranous and
puberulous. Androecium 1-1.2 cm long; stamens numerous, in 3 whorls and more
or less in 5 bundles, the filaments hairy below, connate briefly with the base of
corolla tube. Gynoecium about 1 cm long; ovary globose, densely sericeous, about
6 mm across, the upper tapering into a short style (about 4 mm long) with 5 branches
(about 2 mm long). Capsule (immature) ovoid, 2 cm long, subtended by the persis-
tent calyx.
DISTRIBUTION. Malesia (the Malay Peninsula: Perak and Pahang).
Malay Peninsula. Perak, Taiping Hill, Md. Haniff & Md. Nur SFN 2359 (isotype, SING); Caulfield’s
Hill, Md. Haniff & Md. Nur SFN 12734 (SING); Birch’s Hill, Wray 617 (SING). Pahang. Cameron
Highlands, Henderson 11191] (SING).
42 Gard. Bull. Sing. 37(1) (1984)
ECOLoGy. In montane forests, altitude 1200-1700m. Fl.; Feb. & June.
21. Gordonia vulcanica (Korth.) H. Keng, comb. nov.
Laplacea vulcanica Korth., Kruidk. (1842) 138, ¢. 26; Miq., Fl. Ind. Bat. 2
(1857) 490.
Haemocharis vulcanica O. Ktze, Rev. Gen. (1891) 62 (as ‘vulcania’); Burk., op.
Ciisg MSI.
Gordonia densiflora Ridl. J. Fed. Mal. St. Mus. 8 pt. 4 (1917) 17 Syn. nov.
Fig. 21
var. vulcanica
A stout shrub, 2-6 m tall, or sometimes a crooked tree to 20 m tall, densely
crowned. Young twigs densely covered with yellow or black hispid hairs; older
branches stout, less hairy. Leaf-blades coriaceous or thick-coriaceous, ovate or
broadly oblong, apex rounded or obtuse, sometimes retuse or emarginate, base
rounded or very shallowly cordate, 2.5-5 (-6) cm long, 2-2.5 cm wide; margin of
the upper half or two-thirds remotely serrulate or subentire; nerves 5-7 pairs, often
slightly impressed above, faint or inconspicuous beneath; drying green and glabrous
but often verrucous above, light green and somewhat faintly glaucous below, strigose
near and on the midrib especially at the base; petiole stout, 1-3 mm long, sometimes
almost sessile, hispid. Flowers in upper axils or subterminal, solitary or 2-3
together; peduncles almost absent or very short, 1-2 mm long, stout. Bracteoles,
bracts and sepals 8-10, forming an involucre about 1-1.2 cm high, silvery wooly on
the back, and increasing in size from the lower bracteoles (deltoid, 2-3 mm long)
to the upper sepals (broadly obovate or suborbicular, 1-1.2 cm long). Corolla
4-5 cm across, creamy white (fide de Wilde); petals 5-6, varying from reniform to
obcordate, 1.5-2.5 cm long, membranous, tapering below, glabrous except the
lower one or two which are silvery puberulous and thickened at the back in the
central portion, all petals are briefly joined at the base and adnate to the filaments.
Androecium 6-7 mm long, the filaments briefly united below, often in 5 less distinct
fascicles. Gynoecium 8-9 mm long; style columnar, 2-3 mm long, ridged, the upper
portion separating into 5 free branches; ovary ovoid, 4-5 mm long, strigose or
velutinous. Capsule 2.5-3 cm long, puberulous, dehiscing into 5 valves; sepals
caducous eventually. Seeds 1.2-1.5 cm long including the wing.
DISTRIBUTION. Malesia (W. & Central Sumatra.)
Sumatra. Without precise locality, Korthals Herb. Lugd. Bat. 908. 251-803,-804, -815, -824 (lec-
totype) (L); Beccari 207 (L). Atjeh, Jeswiet 6827 (L); G. Losir, Atjeh, Steenis 8473 (BO), 8493, 8636,
9653 (L), de Wilde & de Wilde-Duyfjes 15276, 15432, 16141, 16565 (L). Mt. Sinabung, Lorzing 8/82,
13681 (L). G. Singgalang, Bunnemeijer 2839 (L); Mt Kerintji, Jacobs 4417 (L); Korinchi Peak, Robinson
& Kloss s.n. 10 May 1914 (holotype of Gordonia densiflora Ridl., SING). Mt Tanggamus, Lampung,
Jacobs 8246 (L).
ECOLOGY. In montane forest, mostly in mossy elfin forests or in dense
Gleichenia scrubs, sometimes near streams; alt. 2000-3400 m. FIl.: Jan.—May; fr.:
April-July.
NOTE. Korthals described this species as having 5 free styles. This was because his
description was based on small flower-buds rather than fully developed flowers.
Gordonia in Malesia 43
Ridley’s Gordonia densiflora is clearly a synonym.
var. buxifolia (Mig.) H. Keng, stat. nov.
Laplacea buxifolia Miq., Fl. Ind. Bat. Suppl. (1861) 482.
Haemocharis buxifolia (Miq.) Szyszyl. in E. & P., Pflanzenfam. 3, 6 (1893) 185;
Burk., J. Str. Br. Roy. As. Soc. 76 (1917) 158.
It differs from the above in the smaller and narrower (3-4 x 1-1.5 cm, rarely to
6 x 2 cm) leaves with less conspicuous nerves, and in the smaller fruit (1.2-1.4 cm
long) (? immature).
Fig. 20. Gordonia taipingensis Burk.
Malaya, Md. Haniff & Md. Nur SFN 12734 (f1.).
44 Gard. Bull. Sing. 37(1) (1984)
DISTRIBUTION. Malesia (W. Sumatra.)
Sumatra. Paya Kombo, Herb. Teysmann H. B. 656 (isotype of Laplacea buxifolia Miq.); Mt. Sago,
near Pajakumbuh, Meijer 5525; Taram, Bukit Paku, Pajakumbuh, Meijer 7/71. Gajo Loeus G. Agosan,
Neth. Ind. For. Serv. bb. 22441 (L).
ECOLOGY. Meijer 5525 was collected from 1800-2000 m, and Meijer 7171, from
600 m. Fr.: July.
NOTE. This variety probably represents merely a diminutive form of the above.
The two taxa may have to be merged eventually.
Fig. 21. Gordonia vulcanica (Korth.) H. Keng comb. nov.
Sumatra, Steenis 8493 (habit); de Wilde & de Wilde-Duyfjes 16565 fl & fr).
2. Gordonia lanceifolia Burk., op. cit., 150.
Gordonia in Malesia
45
EXCLUDED AND DOUBTFUL SPECIES
1. Gordonia brevifolia Hook.f., Trans. Linn. Soc. 23 (1860) 162; Burk., J. Str. Br.
Roy. As. Soc. 76 (1917) 158.
= Schima brevifolia (Hook.f.) Stapf. in Hook. Ic. IV, 3 (1893) ¢. 2264, Trans.
Linn. Soc. Bot. 4 (1894) 135.
Fig. 22
Burkill writes: ‘‘G. /anceifolia, a new species, comes near to G. /uzonica. It has
leaves of the same outline but more nearly entire, and differently veined. Its
Fig. 22. Gordonia lanceifolia Burk.
Sarawak, Haviland 1010 (holotype).
46 Gard. Bull. Sing. 37(1) (1984)
capsule terminate leafless branchlets which may be 4 cm long. It occurs in Borneo
near Kuching’’. Only a single specimen, the holotype, Haviland 1010 (SING) was
available for this study, which is a fruiting one. A recent (here depicted in figure
22) collection from Sarawak, which is close to or probably referable to this
species is: P. Chai S§ 32128 (L, SAN), collected from Segan For. Res., Bintulu
on Sept. 19, 1972. It differs from the type specimen, however, in the smaller
leaves (to 11 vs. to 13 cm long) and fruits (to 2.5 vs. to 3 cm long). There is only
a single S-valved fruit in the type, but over twenty dry capsules in P. Chai’s
collection, mostly 6-valved, few are 7- or 5-valved.
3. Gordonia lobbii Hook. f., Trans. Linn. Soc 23 (1860) 162; Burk., op. cit., 156.
This species was based on T. Lobb’s collection from Sarawak. Mr L. L. Forman
of the Royal Botanic Gardens, Kew, kindly sent me a photo of the type specimen
and copied Mr Airy-Shaw’s annotation as follows: ‘‘On right, middle —’ leaves
crenate/ cf. S. superba Grdn. et Champ. The word ’Schima’ (Bottom right
corner) is in Pierre’s hand’’. Shaw’s note says ‘verisim a cl. Pierre scriptum’. The
photo appears to confirm J. B. L. Pierre’s and H. K. Airy-Shaw’s identification
that it is a specimen of Schima.
4. Laplacea sarasini Warb. MS.
Melchior [(in E. & P. Pflanzenfam. ed. 2, 21 (1925) 136)] mentioned this un-
published S. Celebes species in his key to the Malesian Laplacea. it differs from
other species, according to his key, in the leaves being more or less herbaceous,
glabrous, and with an acuminate (or cuspidate) apex. No specimens have been
seen. L. sarasini is probably a synonym of G. amboinensis.
IV. ACKNOWLEDGEMENTS
I am grateful to the Commissioner and staff of the Botanic Gardens, Singapore for
the herbarium and library facilities, to the Director and staff of the Rijksherbarium,
Leiden, for the loan of the entire collection of Gordonia, to the Directors and staff
of Herbarium Bogoriense, Bogor, Herbarium of the Forest Department, Sandakan,
for the loan of some critical specimens, and to the Directors of the Arnold Arboretum,
Cambridge, and Royal Botanic Gardens, Kew, for supplying photographs of the
type specimens of G. grandiflora Merr. and G. lobbii Hook. f., respectively.
I also would like to thank Professor Dr. C.G.G.J. van Steenis for patiently
answering my queries and for going through the manuscript of this paper, and for
his valuable comments; Dr. Ding Hou for his advice on certain nomenclatural
problems and for supplying the xerox and photocopies of literature; and my wife,
Mrs. Ro-siu Ling Keng for preparing the illustrations of this paper and for her
encouragement.
Gordonia in Malesia
INDEX TO SCIENTIFIC NAMES
Antheeischima excelsa (Bl\.) Korth.
= Gordonia excelsa
Closaschima marginata Korth.
= Gordonia marginata
Closaschima ovalis Korth.
= Gordonia ovalis
Gordonia acuminata Choisy
= Gordonia excelsa
Gordonia amboinensis (Miq.) Merr.
Gordonia berguetica Burk.
= Gordonia luzonica
Gordonia borneensis H. Keng
Gordonia brassii Kobuski
= Gordonia amboinensis
Gordonia brevifolia Hook. f.
= Schima wallichii (DC.) Korth. ssp.
brevifolia (Hook. f) Bloemb. ex Walp.
Gordonia concentricicatrix Burk.
= Gordonia multinervis
Gordonia densiflora Ridl.
= Gordonia vulcanica
Gordonia excelsa (BI.) BI.
Gordonia fragrans Merr.
= Gordonia luzonica
Gordonia grandiflora Merr.
Gordonia grandis King (non André)
= Gordonia singaporeana
Gordonia havilandii Burk.
Gordonia hirtella Ridl.
Gordonia imbricata King
Gordonia integerrima (Miq.) H Keng
Gordonia lanceifolia Burk.
Gordonia lobbii Hook. f.
= Schima wallichi (DC.) Korth.
ssp. noronhae Bloemb. var.
noronhae Bloemb.
Gordonia luzonica Vidal
Gordonia maingayi Dyer
Gordonia marginata (Korth.) Endl.
ex Walp.
Gordonia multinervis King
Gordonia oblongifolia (Miq.) Steenis
Gordonia ovalis (Korth.) Korth.
Gordonia papuana Kobuski
= Gordonia amboinensis
Gordonia penangensis Ridl.
Gordonia polisana Burk.
Gordonia rumphii Merr.
= Gordonia amboinensis
Gordonia sarawakensis H. Keng
Gordonia scortechinii King
Gordonia singaporeana Wall. ex Ridl.
Gordonia subclavata Burk.
?= Gordonia luzonica
Gordonia taipingensis Burk.
Gordonia vulcanica (Korth.) H. Keng
Gordonia vulcanica (Korth.) H. Keng
var. buxifolia (Miq.) H. Keng
Gordonia welbornei Elm.
= Gordonia luzonica
Haemocharis amboinensis (Miq.) Burk.
= Gordonia amboinensis
Haemocharis aromatica (Miq.) Szyszyl.
= Gordonia ovalis
Haemocharis buxifolia (Miq.) Szyszyl.
= Gordonia vulcanica var. buxifolia
Haemocharis integerrima (Miq.) K. & V.
= Gordonia integerrima
Haemocharis marginata (Korth.) O.K.
= Gordonia marginata
Haemocharis ovalis (Korth.) O.K.
= Gordonia ovalis
Haemocharis serrata K. & V.
= Gordonia integerrima
Haemocharis subintegerrima (Miq.) Burk.
= Gordonia ovalis
Haemocharis vulcanica (Korth.) O.K.
= Gordonia vulcanica
Laplacea amboinensis Miq.
= Gordonia amboinensis
Laplacea aromatica Miq.
= Gordonia ovalis
Laplacea buxifolia Miq.
= Gordonia vulcanica var. buxifolia
Laplacea integerrima Miq.
= Gordonia integerrima
Laplacea marginata (Korth.) Choisy
= Gordonia marginata
Laplacea ovalis (Korth.) Choisy
= Gordonia ovalis
Laplacea sarasini Warb.
?= Gordonia amboinensis
Laplacea serrata (K. & V.) Melch.
= Gordonia integerrima
Laplacea subintegerrima Miq.
= Gordonia ovalis
Laplacea vulcanica Korth.
= Gordonia vulcanica
Ploiarium ? oblongifolium Miq.
= Gordonia oblongifolia
Schima excelsa Bl.
= Gordonia excelsa
47
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A MONOGRAPH OF MELIA IN ASIA AND THE PACIFIC
The history of White Cedar and Persian Lilac
D.J. MABBERLEY
Departments of Botany and Forestry, University of Oxford, Oxford, England
EFFECTIVE-PUBLICATION DATE: 5 Sept 1984
Abstract
The wild forms of Melia in Asia are described and assigned to one species. The relationship of that
taxon to plants cultivated in Asia, Europe, America and Africa is examined and it is concluded that
selected forms have been long cultivated in both India and China. Groups of cultivars have been selected
from these two centres of domestication and introduced to other warm parts of the world. The
widespread Persian Lilac or mindi kechil seems to be of Indian stock and to have been widely introduced
in America and Africa: mutant forms of it include the Texas Umbrella Tree. The Chinese cultivars are
those widely grown in Japan and introduced thence to Europe as M. japonica. The whole complex is
treated as one species (M. azedarach) comprising the wild populations and the two major groups of
horticultural cultivars which are set out in a formal revision. All names in Melia and its synonyms applied
to Asiatic and Pacific plants are identified in an appendix, while the typification of M. azedarach is set
out in another.
WILD MELIA iN ASIA
In the Gunung Leuser Nature Reserve, Aceh, northern Sumatra, grow magnifi-
cent specimens of a species of Melia. Those collected by W. de Wilde and his wife
(15884, 16548 - L!) in the Lau Alas valley in 1975 were noted as being 40 m tall with
a diameter at breast height of some 60 cm; the flowers had white petals and the
staminal tube was yellow at the apex, turning purple-red as the flower matured; the
fruits were some 3 cm long, the twigs stout and heavily clothed with a fulvous
stellate indumentum when young, while the leaflets of the bipinnate leaves were
strongly acuminate with rather obtuse or even rounded bases and were up to some
6 cm long. These specimens are exceptionally fine examples of a forest species which
seems to grow wild throughout the more seasonal forests of Indomalesia, from
India, Burma and Malesia (absent from the Malay Peninsula and Borneo) to the
Solomon Islands and tropical Australia. In India, it has been known as M. superba
and M. robusta, names based on material collected from trees grown in the Calcutta
Botanic Garden from seeds gathered in Soonda (? Carnatic) and ‘Malabar’ respec-
tively by Andrew Berry, or as M. composita and more lately as M. dubia. In Burma,
it is M. birmanica, in Indochina M. composita var. cochinchinensis; in Java it was
described from cultivated material as M. bogoriensis while Timorese material has
been called M. candollei and seems not to differ substantially from M. azedarach
var. australasica, the White Cedar of tropical Australia. The specimens from Java
and further east seem to be somewhat more glabrous than those collected in
Sumatra, while material from Australia often has distinctly pink or even mauve
flowers. Nevertheless this wide distribution with such minor variation is typical of
Asiatic Meliaceae of drier forests and other more open vegetation, being very similar
to that of Turraea virens L. (incl. T. pubescens Hell.) for example, while Cipadessa
49
50 Gard. Bull. Sing. 37(1) (1984)
baccifera (Roth) Miq., which is found only in the western part of the range of those
two species, is again absent from the forests of the Malay Peninsula and Borneo.
MELIA IN CULTIVATION
Outside botanic gardens in Malesia, the common Melia cultivated is the Persian
Lilac or Bead-Tree (fig. 1). It is typical M. azedarach, the type species of the genus,
and is the only Melia to be seen in the Malay Peninsula and Borneo. In the first and
in Singapore, it is known as mindi kechil. It is one of the most widely cultivated of
all tropical trees and differs from the wild trees described above in its variably lobed
or serrated leaflets, which are smaller, its relative glabrousness and its large lilac,
blue, or occasionally white flowers. As it flowers precociously (even as seedlings
sometimes - see van Steenis in Flora Malesiana 1, 4 (1948) xxi - like a number of
other species in cultivation, where this phenomenon is most readily observed), forms
of it can be grown in conservatories in temperate countries and in bedding schemes.
Selected hardy forms of this plant, which, when fully grown, is straggly and of poor
form, may be grown outside in Europe, even in Great Britain where it was probably
first cultivated in the sixteenth century.
The source of the European plants was probably Iran or elsewhere in the Middle
East, where it has been long known and both its common English name as well as
its specific epithet commemorate this. Azedarach is a Persian word, or at least a
rendering of a@zadraxt or azadiraxt, which Laufer (Field Mus. Nat. Hist. Publ. 201,
Anthrop. ser. 15, 3 (1919) 583) takes as the name for Azadirachta indica A. Juss.
(M. azadirachta L.), the neem tree. The Arab philosopher and physician Avicenna
(979-1037) refers to the name, which probably first appeared in a European text in
the work of L’Obel (Nova stirpium adversaria. . .(1576)) as Azedaraeth. It has some
medicinal properties but has been widely cultivated in the Mediterranean for shade
and as a source of beads for rosaries and so forth, the fruits with a central channel
through them when dried being ideally suited to this. Indeed the tree became known
there as arbor sancta or arbor pareiso (i.e. Paradise) a name still common in
Spanish-speaking countries. The earliest reference I have found to it is as (is)zanzali-
qu on a stela of the Assyrian king, Assurnasirpal II, near the doorway of the palace
leading to the throne room at Nimrud, the ancient military capital. The inscription
is dated ca. 879 B.C. and describes the trees planted: Persian Lilac was one of the
42 species of tree in this Babylonian botanic garden (see D.J. Wiseman in Jraq 14
(1952) 24-44 and M. Levey in Isis 52 (1961) 94-95). It is very tough and is naturalized
in many warm countries, sometimes becoming, as in Madagascar for example, an
aggressive pioneer. By contrast, the White Cedar is rarely grown outside the tropics,
where a number of selected cultivars are used in forestry and, in Great Britain,
where it was introduced in the 1750s, it must be grown under glass.
According to Bailey (Stand. Cyclop. Hort. 2 (1933) 2024), M. azedarach is widely
naturalized in the southern United States. He reports that ‘Several forms have been
found, a white-flowering and one with the segms. of the Ifts. cut in narrow divisions.
These forms are not constant, the seedlings frequently reverting to the typical
species’. Such may be the forms described from cultivated plants and known as M.
sambucina and M. azedarach var. incisa (Java) and var. acuminatissima (Sulawesi)
Melia in Asia & Pacific 51
and var. subtripinnata (Japan), though the last may be a form of the ‘t6 sendan’
(see below).
A particularly precocious floriferous form has been named M. floribunda, though
perhaps the most striking of all is that known as the Texas Umbrella Tree, which
has a dense flattened crown. Bailey notes, ‘The first tree that came to notice is said
to have been found near the battlefield of San Jacinto, Texas, but with no record
of its intro. there. If the fls. are not crosspollinated with the common sort, the
percentage of seedlings which reproduce the exact umbrella shape seldom varies;
hence it is supposed by some to be a distinct species’. Linnaeus himself described
a variety sempervirens from cultivated Sinhalese material grown in Holland, a form
which retained its leaves rather more then does the typical plant. However, the text
to the plate (see fig. 1) in the Botanical Magazine (27 (1807) t. 1066) suggests that
even the deciduous form would retain its leaves more effectively under stovehouse
rather than greenhouse conditions in Europe. According to Watt (Dict. Econ. Prod.
India 5 (1891) 221) these Sinhalese plants were brought south from northwest India.
An apparently similar form seems to have arisen spontaneously in Argentina
(Balozet in Rev. Bot. App. Agr. Trop. 33 (1933) 461).
Fig.l. Persian Lilac as long grown in Europe.
Drawing by Sydenham Edwards from material cultivated in a nursery near London at the
beginning of the nineteenth century (Bot. Mag. 27 (1807) t. 1066).
Sy Gard. Bull. Sing. 37(1) (1984)
Melia, as circumscribed by Harms (1940), is an Old World genus*, and all
neotropical populations are derived from introduced stock of M. azedarach. Of
those in the West Indies (‘M. sempervirens’, though whether they are the form
Linnaeus named var. sempervirens is not clear), some were brought back to the Old
World, much as other Old World plants such as the African Parkia biglobosa
(Jacq.) R. Br. ex G. Don f. (Leguminosae) were. The Melia became naturalized,
notably in West Africa. Specimens collected there were duly named M. angustifolia
and living plants called M. guineensis, but there was never any doubt that these were
of cultivated origin.
In the nineteenth century, a new source of cultivated Melia for European gardens
was Japan: the trees were called M. japonica and had larger fruits and more entire
leaflets. In Japan, they were known as (6 sendan, i.e., Melia from China, and it was
soon realized that these, like so many Japanese garden plants, were anciently in-
troduced from China, where they had been long cultivated. The first mention of the
plant in Chinese literature is in the ancient Taoist classic Chuang-tze, where it is said
that the fruit was eaten by a fabulous bird: the text is c. 300 B.C. In Ch’t Ta-chtn’s
Kuang-tsung hsin-yu of 1700, there is an article on k’u-lian (k’u means bitter as in
k’u-li (coolie), bitter strength), p. 641, ‘It is very easy to grow. In the villages
whenever a girl is born they always plant many of them in order to make vessels at
the time when they marry. The fruit is bitter and should not be put in the mouth’.
He also notes that it was much prized in antiquity and that the flowers were used
for making incense, while in the late Ming and Ch’ing periods, the wood was used
for patten-making. It was early grown in Japan for it is mentioned several times
in the earliest extant anthology of Japanese poetry, the Manyoshu of the mid-eighth
century.
Forms from China were named M. toosendan and plants selected in Japan were
named, e.g., var semperflorens, f. albiflora, but Makino, who described many of
these at different ranks and under different specific names, records (Bot. Mag.
Tokyo 28 (1914) 35) that hybrids between ‘subvar. japonica’ and ‘subvar.
toosendan’, which he named subvar. intermedia, occur when these forms are grown
together.
THE STORY SO FAR
From the foregoing, it can be seen that Persian Lilac in cultivation is extremely
variable but that the selected forms which look very different have not lost the
capacity to form fertile hybrids. Furthermore, the stocks in Europe and North
America seem to have at least two distinct origins: India via the Middle East (and
via Sri Lanka and the Netherlands) and China via Japan. White Cedar is a
somewhat variable forest tree, tender in Europe and the bulk of specimens referred
to it are readily distinguishable from the commonly cultivated forms of Persian
Lilac. As Harms noted, however, it would be misleading to infer from this that all
specimens can be neatly pigeonholed, for there are specimens of White Cedar with
* See Pennington in Flora neotropica 28 (1981) 25 for fate of names published in Melia for native
neotropical plants.
Melia in Asia & Pacific 53
little indumentum or with rather serrate leaflets. Fully ripe fruits are rarely gathered
and collectors do not always note the colour of the flowers, which changes with time
in any case, while young leaves are often gathered with flowers, the mature ones
being neglected. Furthermore, I have been quite unable to correlate differences in
flower colour, leaf size, shape and indumentum with geography. Indeed, as far as
the Asiatic representatives of Melia are concerned, then, I cannot but agree with
Ramamoorthy (in Saldanha & Nicolson, F/. Hassan Dist. (1976) 395) who notes,
‘The taxa of this genus (species, varieties, etc.) apparently intergrade’.
In Africa, the matter is as yet not completely resolved in that there are apparently
truly wild trees besides the cultivated Persian Lilac. Those in Angola have been
included in M. dubia, i.e., White Cedar in the sense of this paper, by Exell and
Mendonga, Consp. Fl. Angol. 1 (1951) 318. However, the limited material I have
seen looks rather different and has been referred to the distinct species M. volkensii
Guerke (East Africa) and M. bombolo Welw. (Angola). These clearly require
further study but are something of a sideshow as far as the problem of the taxonomy
of the bulk of Melia is concerned.
POSSIBLE SYSTEMATIC ARRANGEMENTS
Miquel, who worked on the Japanese cultivars as well as those found in Malesia,
wrote (Ann. Mus. Bot. Lugd. - Bat. 3 (1868), ‘in universum Meliae species nondum
ab omni parte satis inter se comparatae novo indigent examine’. Harms, a reluctant
‘lumper’ elsewhere in Meliaceae, felt (1940, q.v. for detailed references of following
works) that there were indeed only two species in Asia and the Pacific, and these
were separable only with difficulty: M. azedarach (Persian Lilac) and M. dubia. The
earlier monographer of the family, Casimir de Candolle (in DC., Mon. Phan. 1
(1878)), in the meantime, had maintained a number of previously described species
as distinct and recognized three varieties of M. azedarach: var. glabrior (Persian
Lilac), a superfluous name for the type variety, var. australasica (White Cedar) and
var. squamulosa for forms of the wild tree with a rather heavy indumentum found
in mainland Asia and Java. Koorders & Valeton (in Med. ’s Lands Pl. Tuin 16
(1896)) recognized a number of species in the complex and designated the wild
Javanese tree var. javanica. Pierre (Fl. For. Cochinch. (1897)) had the wild Indo-
Chinese tree as var. glandulosa and Pellegrin (in Lecomte, F/. Gén. Indoch. 1
(1911)) gave the name var. cochinchinensis to such a plant.
The cultivated forms have also been variously reduced to varieties, subvarieties
or forms, though it would seem that if these are worth recognizing, a cultivar
nomenclature would be more appropriate as it is in the provenances used in forestry.
Indeed, two extreme types have already been thus designated (see below) but as for
any further naming, particularly of the Chinese cultivars, this can be largely left to
those who deal with the plants readily available in the trade. The problem from the
botanist’s standpoint is what to do with White Cedar, given that it is not apparently
specifically distinct from the type, which is Persian Lilac (see Appendix II).
Of possible solutions, the simplest would be to amalgamate all the Asiatic plants
and those introduced into the rest of the world as M. azedarach. This would have
54 Gard. Bull. Sing. 37(1) (1984)
the advantage of coping with the intermediate specimens and would leave open the
question of the origin of the original ‘Indian’ material and the Chinese plants.
Indeed, with the widespread cultivation of the tree and destruction of the original
forest, it may now be quite impossible to reconstruct this history anyway. It would
have the disadvantage, however, of throwing together cultivated and wild plants in
Asia, where the wild ones are manifestly rather different in form and so forth as
well as in value for horticulture. The opposite solution would be to maintain the wild
Asiatic trees as one species whilst maintaining the widespread cultivated Persian
Lilac as another, as Harms (1940) did. This solution conceals the apparent close
relationship of the two ‘species’ notably in not coping with the rather intermediate
position of the Chinese cultivars. In Malesia, it would be convenient to be able to
distinguish the truly wild trees from the introduced exotics but as the whole cannot
be honestly treated other than as a single species, some intraspecific system is
required. With some cultivated plants, it has been found appropriate to name the
presumed ancestral populations at the subspecific or varietal level, as in the case of
the carrot, sugar beet and teasel for example. In making such a category for the wild
plant, in this case, White Cedar, however, the cultivated plants would fall into the
nominal variety or subspecies. This is exactly paralleled by the hypothesized state
of affairs in the grape vine, Vitis vinifera L. (Burtt, Biol. J. Linn. Soc. 2 (1970)
233-238)), where Linnaeus is deemed to have made a type variety when actually
naming an atypical variant as a variety, while the same plant would be the typical
subspecies should an atypical subspecies be described in V. vinifera. In M.
azedarach, such a formal system would be undesirable in any case, as the Chinese
and the Indian cultivars would be drawn together in what would be, in effect, a
polyphyletic taxon, segregated from the wild forms, to certain groups of which
cultivar grouping is more closely allied than to one another.
In view of this, then, it seems preferable to take Burtt’s advice and to avoid the
artificiality of the formal infraspecific hierarchy and use an informal system, which,
in this case, is very straightforward. The whole complex is to be known by its oldest
name, M. azedarach, even though the type is a cultivated plant. Those cultivated
plants which merit recognition as distinct should be given cultivar names by those
who deal in the horticultural trade. No name seems to me to have been given at this
level to the Persian Lilac in its commonest Indian or ‘Japanese’ forms, though there
are a host of varietal names. The most precocious forms have already been called
‘Floribunda’, the Texas Umbrella Tree ‘Umbraculifera’. Here then, from a
botanist’s standpoint, the synonyms pertaining to the wild tree are grouped together
and those to the horticultural trees are tentatively grouped into Chinese and Indian
cultivars.
MELIA L.
Melia L., Sp. Pl. 1 (1753) 384; Pennington, Blumea 22 (1975) 463. - Type:
M. azedarach L.
Azedarach Mill., Gard. Dict. Abr. ed. 4 (1754) [170]. - Type: not indicated.
Zederachia Heist. ex. Fabr., Enum. Meth. PI. (1759) 221, nom. superfl. pro Melia L.
Antelaea Gaertn., Fruct. Sem. Pl. 1 (1788) 277. - Type: A. javanica Gaertn.
Melia in Asia & Pacific 55
Azedara Raf., Fl. Ludov. (1817) 135, nom. superfl. pro Melia L.
Azedaraca Raf., Med. FI. 2 (1830) 199, nom. superfl. pro Melia L.
Trees, occasionally flowering precociously as shrublets. Indumentum of simple
and stellate-tufted hairs. Leaves 2(3)-pinnate. Inflorescence thyrsoid, axillary.
Flowers hermaphrodite and male on same tree. Ca/yx 5(6)-lobed to near base, lobes
somewhat imbricate. Petals 5(6), free, imbricate. Staminal tube narrowly cylindrical,
slightly expanded at mouth, 10(12)-ribbed, with 10 or 12 truncate, bifid or 4-fid
filiform lobes; anthers 10(12), inserted at margin or just within tube, alternating
with or opposite lobes. Disk small, surrounding base of ovary. Ovary 4-8-locular,
each locule with 2 superposed ovules; stylehead capitate to coroniform with 4-8
short, erect or incurved stigmatic lobes. Drupe 3-8-locular; endocarp thick, bony,
deeply dimpled at both ends; loculi 1(2)-seeded. Seed oblong, laterally compressed;
testa leathery, sometimes slightly swollen and fleshy round hilum; embryo embedded
in oily endosperm; cotyledons flat. Germination phanerocotylar; eophylls opposite,
pinnatisect or trifoliolate. 2n = 28.
One species in Indomalesia with possibly two closely allied ones in south and east
tropical Africa. Forms of the Indomalesian species are widely cultivated and
naturalized throughout the warm parts of the world.
SYSTEMATIC ARRANGEMENT OF THE COMPLEX OF WILD (ASIATIC)
AND CULTIVATED FORMS
Melia azedarach L., Sp. Pl. 1 (1753) 384. - Type: Holland, de Hartecamp, cult.
Hort. Cliff. 161.1 (BM! lecto; see Appendix II).
Azederach deleteria Medik., Geschl. Malv. (1787) 116.
M. florida Salisb., Prodr. Hort. Chapel Allert. (1796) 317, nom. superfl.
Azedara speciosa Raf., Fl. Ludov. (1817) 30, nom. superfl.
Azedaraca amena Raf., Med. Fl. 2 (1830) 199, nom. superfl.
M. azedarach var. glabrior C. DC. in DC., Mon. Phan. 1 (1878) 452, nom. superfl.
— Azedarach sempervirens (L.) O. Ktze var. glabrior O. Ktze, Rev. Gen. 1 (1891)
109.
A. vulgaris Gomez de la Maza in Repert. Med.-farm. Havana 5 (1894) 296 (n.v.).
Tree to 40 m, + deciduous: bole fluted when old, to 60 (180) cm diam. Bark grey-
brown, smooth, lenticellate, becoming lightly fissured or scaling with age; inner
bark yellowish; sapwood whitish, soft; heartwood rusty brown. Crown of widely
spread but sparsely branched limbs. Twigs upturned at ends of drooping branches,
smooth, brown, lenticellate, with raised cicatrices. Leafy twigs ca. 6-8 mm diam.,
+ clothed with fulvous stellate hairs. Leaves 15-80 cm long with 3-7 pairs of lateral
rachides, each with 3-7 leaflets, the most proximal of which sometimes replaced by
short rachides with a few pairs of leaflets, + weakly pubescent but usually
subglabrous; petiole 8-30 cm, to 6 mm diam., terete, lenticellate, swollen at base;
lateral rachides to 25 cm long, weakly ascendant, articulated with jointed main
rachis and weakly swollen there; leaflets 3-6 (— 10) x 1-2.5 (—3) cm, ovate or
oblong-lanceolate to elliptic, base acute to rounded, apex acuminate, margin entire
56 Gard. Bull. Sing. 37(1) (1984)
to variously serrate, costae ca. 7-10 on each side, subsquarrose to weakly ascendant
and arcuate, looped at margin; petiolules 3-7 mm. Thyrses 10-12 cm, axillary or
borne on shoots with terminal bud (see Corner, Wayside Trees 1 (1940) 464);
primary branches ca. 5-7.5 cm long, weakly ascendant, secondary to 2 cm, bearing
fascicles of scented flowers; axes + mealy pubescent; bracts 3-10 mm long,
filiform, pubescent, caducous; bracteoles similar but smaller; pedicels ca. 2-3 mm
long. Calyx ca. 2 mm diam.; lobes ca. 2 mm long, ovate, stellate- and simple-hairy
without, margin ciliate. Petals 6-10 x 2 mm, narrowly oblong, white to lilac or
bluish, stellate- and simple-hairy without, sometimes simple-hairy within, midvein
conspicuous. Staminal tube subglabrous without, + densely simple hairy within,
lobes bifid, or 4-fid, sometimes irregularly so; anthers ca. 1.5 mm long, apiculate,
+ hairy, inserted opposite lobes. Disk obscure and closely enveloping ovary. Pistil
glabrous; stylehead ca. 0.75 mm diam. Drupe 2-4 cm long, 1-2 cm diam., plum-
shaped, glabrous, yellow-brown when ripe; endocarp very hard. Seed ca. 3.5 X 1.6
mm, oblong, smooth, brown.
Wild trees are known from India, Nepal, Sri Lanka and tropical China south and
east through Malesia (Sumatra, Java, Philippines (Luzon, Negros, Mindanao),
Lesser Sunda Isls. (Flores, Timor, Wetar), New Guinea to tropical Australia and
Solomon Isls. to 1200 m (1800 m in Himalayan tract) in forests, particularly seasonal
ones including bamboo thickets (Thailand) and those on limestone, Tamarindus
woodland and Eucalyptus savanna, where it may coppice. Cultivated forms persist
and may become naturalized in secondary vegetation in warm parts of the world.
Uses
The wood of different forms of the species has been used for furniture and light
construction, notably for ceilings, boats and tea boxes while, during the American
Civil War, the trees were a commercial source of alcohol (10% by weight from the
fruits). In tropical America, it has been grown in plantation for the production of
fibreboard (Pennington in Flora neotropica 28 (1981) 24) and is important in the
sports goods industries of Pakistan (Amjad & Mohammed in Pak. J. For. 1 (1980)
39). If not grown too fast, it is good for paper making (Singh ef al., Ind. For. 103
(1977) 641). It has been used as fastgrowing coffee-shade and it is alleged that fruit
trees grown under it remain relatively free of aphids. Indeed a decoction of the fruits
has long been used as an insecticide for plants in India and China and fruits or leaves
have been placed with dried fruit, clothing and in books to keep insects away. An
extract has also been used as a fish poison.
Although there has been considerable confusion with neem, Azadirachta indica
A. Juss., M. azedarach has been widely attributed with medicinal qualtities. Its root
appears as Cortex Meliae azedarach in pharmacopoeias, but it is generally held that
the bark is most efficacious, particularly as a vermifuge. For details of its action and
other medicinal uses, see G.A. Stuart, Chinese materia medica: vegetable kingdom
(1911) 261 and L.M. Perry, Medicinal plants of East and Southeast Asia (1980) 262.
So valued are the fruits in the Malay Peninsula, that they have been imported from
Szechwan.
Melia in Asia & Pacific 57
The seeds also yield an oil and the trunk a gum but these have been little utilized,
though the first is suitable for soap and hair oil (Wealth of India 6 (1962) 323 et
seqq., q.v. for further uses of the tree).
Informal infraspecific classification
The names applied to the wild trees are set out below, followed by those applied
to the ‘Chinese’ and ‘Indian Cultivars’.
a. ‘Wild’ plants, White Cedar
?Antelaea javanica Gaertn., Fruct. Sem. Pl. 1 (1788) 277, ¢. 58. - type: ‘Java’
(Sri Lanka, KOnig ‘120’ (L, teste Hallier in Réc. Trav. Bot. 15 (1918) 33; lost
teste Jacobs in Gdns’ Bull. Sing. 18 (1961) 74; no duplicates found at BM
or TUB).
Melia dubia Cav., Dissert. 7 (1789) 364, ?nom. provis. (as with Malpighia dubia
Cav., Dissert. 8 (1789) 413 (= Heteropterys laurifolia (L.) A. Juss.,
Malpighiaceae), it could be argued that this name is not validly published for,
like the Malpighia, it is placed at the end of the generic account, here with
the words, ‘Exemplar unicum innominatum vidi in herb. D. de Lamarck,
quod fructibus orbatum facile ad Trichiliam aut ad Meliam Linnaei pertinere
poterit: itaque ut dubiam speciem hic adiungam quoadusque de fructu con-
stet, et ad debitum genus reducatur’. Elsewhere in the work, where he is
uncertain about the disposition of a species, as with Banisteria sericea
(Dissert. 9 (1790) 429), he uses a ‘?’, which is permissible (Int. Code Art.
34.2). Monographers, however, seem to have considered M. dubia valid even
though it is so close to the borderline between provisional and valid (J.F.
Veldkamp, pers. comm.) —- Azedarach sempervirens (L.) O. Ktze. var. dubia
(Cav.) O. Ktze, Rev. Gen. 1 (1891) 110. - Type: (?) Indonesia, (?) Java,
Sonnerat s.n. (P-LAM! holo).
M. composita Willd., Sp. Pl. 2 (1799) 559. - Type: India, 1785, Klein in Hb.
Willd. 8086 (B-WILLD!, holo).
M. robusta Roxb., Hort. Beng. (1814) 33, nom. nud.: Fl. Ind. ed. Carey 2 (1832)
397. - Type: India, Calcutta, cultivated in Bot. Gard. from seed collected
by Berry in Malabar (?CAL; specimen labelled in Roxb.’s hand at G(!);
specimen apparently collected from type tree by Carey in 1824 at E(’)).
M. superba Roxb., Hort. Beng. (1814) 33, nom. nud.; Fl. Ind. ed. Carey 2 (1832)
396. — M. argentea Hiern in Hook.f., Fl. Brit. Ind. 1 (1875) 545, nom. in
synon., sphalm. pro M. superba. - Type: India, Calcutta, cultivated in Bot.
Gard. from seed collected by Berry in Soonda (?CAL; EIC 1254 (K-W!) may
be from this tree).
?M. australis Sweet, Hort. Brit. ed. 2 (1830) 85, nom. nud.; G. Don f., Gen. Syst.
1 (1831) 680. - Type: England, cultivated from material from Australia (?not
preserved).
M. candollei A. Juss., Bull. Sci. Nat. Géol. 23 (1830) 239 - Type: Indonesia,
Timor, (prob.) Gaudichaud (P-JU!, holo; L!, P!).
58 Gard. Bull. Sing. 37(1) (1984)
M. australasica A. Juss., |.c. - M. azedarach var. australasica (A. Juss.) C. DC.
in DC., Mon. Phan. 1 (1878) 452 - A. sempervirens (L.) O. Ktze var.
australasica (A. Juss.) O. Ktze, Rev. Gen. 1 (1891) 110. - Types: Australia,
New South Wales, Port Jackson (P-JU!, syn; P!) & d’Entrecasteaux Isls. (P-
JU!, syn; P!)
[M. azedarach sensu Blanco, FI. Filip. (1837) 345 (‘acedarach’), non L., s.s. Cf.
Merr., Sp. Blanc. (1918) 209.]
[M. flaccida Zipp. ex Span., Linnaea 15 (1841) 182, nom. in synon.]
[M. tomentosa sensu Miq., Fl. Ind. Bat. 1* (1859) 532, non Roxb. (1832, i.e.,
Chisocheton tomentosus (Roxb.) Mabb.]
M. birmanica Kurz, J. Asiat. Soc. Bengal 43, 2 (1874) 183. - Type: Burma, Mar-
taban, Kurz (?CAL, holo).
M. azedarach var. squamulosa C. DC. in DC., Mon. Phan. 1 (1878) 452. - Type:
of de Candolle’s syntypes, I select Java, Zollinger 166 (G-DC!, lecto; L!).
M. bogoriensis Koord. & Val., Med. ’s Lands Plant. Tuin 16 (1896) 18. - Type:
Indonesia, Java, Bogor, cultivated sub III B 12, 6 Dec. 1895 (BO, holo; L!)
M. azedarach var. javanica Koord. & Val. ibid., 15. - Type: of sheets in
Koorders’s herbarium (BO) labelled ‘De hoc specimine agitur in libro ...’
I select Indonesia, Java, Besuki, Curamanis, 17 Oct. 1889, Koorders 51693
(BO, lecto; L!)
M. composita var. cochinchinensis Pierre, Fl. For. Cochinch. 5 (1897) t. 356A.
-— M. azedarach var. cochinchinensis (Pierre) Pellegr. in Lecomte, Fl. Gén.
Indoch. 1 (1911) 727. - Type: of Pierre’s syntypes, I select Vietnam, Pay
Ninh, Cai Cong, 30 Apr. 1866, Pierre 1737 (P!, lecto; L!). N.B. The other
syntype, Pierre 3366 (P!, L!), is a cultivated tree apparently referable to one
of the ‘Indian Cultivars’.
M. azedarach var. glandulosa Pierre, op. cit., t. 356B (“M. composita var. biglan-
dulosa’ in ic.). - Type: Vietnam, Saigon, cultivated in Bot. Garden, Feb.
1876, Pierre 1499 (P!, holo).
Tall forest trees to 40 m. Leaflets + entire, dark above, pale below, when young
densely stellate-tomentose like young shoots. Flowers sweetly scented, scentless or
malodorous. Petals white or pale mauve, often pubescent within. Staminal tube
creamish or pale mauve, darkening to purple with age. Drupe to 4 cm long.
Of the fastgrowing forms planted for forestry in the New World, one has been
named ‘Gigante’ (Cozzo, Rev. For. Argent. 3 (1959) 127) but it is unclear whether
this is intended to be a cultivar name or not, or, indeed, whether it is not more
closely allied to forms of Persian Lilac. Other important forestry provenances
include ‘var. gigantea’ (Ragonese & Garcia in IDIA 385-386 (1980) 110, nom. non
rite publ.), characterized by its general robustness, hairy anthers and large fruits
when compared with Persian Lilac, and treated as two cultivars: ‘Gotz’ and
‘Garrasino INTA, Castelar’, both widely planted in Argentina.
Melia in Asia & Pacific 59
b. ‘Chinese cultivars’, 16 sendan
?M. japonica G. Don. f., Gen. Syst. 1 (1831) 680. - 2M. azedarach var. japonica
(G. Don f.) Mak., Bot. Mag. Tokyo 28 (1914) 34, nom. superfl. - Type:
England, cultivated from Japanese material (?not preserved).
M. toosendan Sieb. & Zucc., Abh. Akad. Muench. 4? (1843) 159. - M. azedarach
subvar. toosendan (Sieb. & Zucc.) Mak., ibid., 35. - M. japonica var.
toosendan (Sieb. & Zucc.) Mak., l.c., mom. in synon. - M. azedarach var.
toosendan (Sieb. & Zucc.) Mak., J. Jap. Bot. 5 (1928) 20. - Type: Japan,
cultivated from Chinese material, Siebold s.n. (?1LE, holo; L! M!).
M. japonica Hassk., Cat. Hort. Bog. Alt. (1844) 219, non G. Don f. (1831) - M.
javanica M.J. Roem., Syn. Hesp. (1846) 96. - M. hasskarlii K. Koch, Hort.
Dendrol. (1853) 72. - Type: Indonesia, Java, cultivated from Japanese
material (?BO, holo).
[M. chinensis Sieb. ex Mig., Ann Mus. Bot. Lugd.-Bat. 3 (1867) 23, nom. in
synon.|
M. japonica G. Don f. var. albicans C. DC. in DC., Mon. Phan. 1 (1878) 457.
- Type: Japan, Nagasaki, cultivated, Oldham 159 (B7, holo; N.B. Oldham
139 at L!).
M. japonica var. semperflorens Mak., Bot. Mag. Tokyo 18 (1904) 67. - M.
azedarach subvar. semperflorens (Mak.) Mak., ibid., 28 (1914) 34. - Type:
Japan, Tokyo, cultivated, 20 July 1903, Makino s.n. (2MAK, holo).
M. azedarach f. albiflora Mak., ibid. - M. japonica var. albiflora Mak., |.c.,
nom. in synon. — Type: Japan, Tosa, Sakawa, cultivated, Makino s.n.
(2MAK, holo).
Small trees. Leaflets usually almost entire. Flowers sweetly scented. Peta/s mauve,
pink or blue, occasionally white. Staminal tube purple.
This is the form commonly cultivated in Japan, though Indian forms are
sometimes grown there. e.g. a white-flowered tree introduced from Lucknow in the
1960s or so (see Hisauchi in J. Jap. Bot. 45 (1970) 256). In Malesia it is rarely seen
outside Botanic Gardens and has larger fruits than the commonly cultivated Persian
Lilac does.
c. ‘Indian Cultivars’, Persian Lilac, Bead-tree, Chinaberry, mindi kechil.
N.B. the cultivars currently recognized as distinct are placed after the main
synonymy.
M. azedarach s.s.
M. azedarach var. sempervirens L., Sp. Pl. (1753) 384. - M. sempervirens (L.)
Sw., Prod. Veg. Ind. Occ. (1788) 67. - Azedarach sempervirens (L.) O. Ktze,
Rev. Gen. 1 (1891) 109. - Type: Holland, de Hartecamp, cultivated, Hort.
Cliff. 161/1a (BM!, lecto; see Wijnands Bot. Commelijns (1983) 145).
60 Gard. Bull. Sing. 37(1) (1984)
M. commelini Medik., Bot. Beobacht. 1782 (1783) 164. - Azederach commelini
Medik. ex Steud., Nomencl. ed. 2, 1 (1840) 175 ‘Moench’ & 2 (1841) 118,
nom. in synon. — Type (lecto): Commelijn, Horti Med. Amst. 1 (1697) 147,
t. 76, drawn from a plant raised from Sinhalese material in Holland.
[A. odorata Nor. in Verh. Bat. Genoot. ed. 1 (1791) art. 4,5, nom. nud.]
M. arguta DC., Prodr. 1 (1824) 622. - Type: Indonesia, ‘Moluccas’, ex Herb.
Lambert, 1815 (G-DC!).
M. sambucina Bl., Bijdr. (1825) 162. - M. azedarach var. sambucina (Bl.) Miq.
Ann. Mus. Bot. Lugd. - Bat. 4 (1868) 5. - A. sambucina (Bl.) O. Ktze, Rev.
Gen. 1 (1891) 110. - Type: Indonesia, Java, cultivated, Blume s.n. (L! holo; P!).
M. angustifolia Schum. & Thonn. in Schum., Beskr. Guin. Pl. (1827) 214. -
Type: Ghana, Elmina, cultivated, Thonning 59 (C (microfiche 67: 1, 3, 4!)).
M. guineensis G. Don f. in Loud., Hort. Brit. (1830) 168 & Gen. Syst. 1 (1831)
681. - Type: England, cultivated from seed from ‘Guinea’ (?not preserved).
?M. bukayun Royle, Illustr. Bot. Himal. (1835) 141, 144. - M. bukheim Griff.,
Itin. Notes (1848) 355, 403. - Type: India, Royle (?LIV).
M. cochinchinensis M.J. Roem., Syn. Hesp. (1846) 95. - Type: M. azedarach
sensu Lour., Fl. Cochinch. (1790) 269.
M. orientalis M.J. Roem., |.c. - Type: M. sempervirens sensu Roxb.; EIC 1252E
is inscribed M. sempervirens in Roxburgh’s hand (K-W!; lecto).
[(M. lobata [Hort. Donat. ex] Planch., Hort. Donat. (1858) 82, nom. in synon.]
[Jacaranda fraxinifolia (Hort. Donat. ex] Planch., l.c., nom. in synon.]
M. azedarach var. subtripinnata Miq., Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 24.
- Types: Japan, Burger s.n. (?U), Keiske s.n. (L!).
M. azedarach var. incisa Miq., ibid., 4 (1868) 5. - Type: Indonesia, Java,
cultivated (L (sheet 908. 133-712)!, holo).
M. azedarach var. acuminatissima Miq., |.c. - Type: Indonesia, Sulawesi,
Amura, cultivated, Dec. 1840, Forsten s.n. (L!, holo).
[M. composita var. cochinchinensis Pierre, Fl. For. Cochinch. 5 (1897) sub tf.
356A, quoad spec. Pierre 3366].
Small trees. Leaflets irregularly serrate, pale green. Flowers sweetly scented. Petals
mauve, pink or blue, occasionally white. Staminal tube purple.
Persian Lilac is one of the most widely cultivated of all tropical trees and indeed
is so readily grown as to be despised by the discriminating. Good forms with dark,
heavily-scented flowers are extremely desirable nevertheless. There is an almost con-
tinuous supply of flowers and of fruits, which have long been used for beads (see
Hoy & Catling in Davidsonia 12 (1981) 65-6 for illus.). The fruits are toxic to man,
some 6-8 being considered a fatal dose, and also to pigs, but apparently not to birds,
sheep or goats, (For a full account, see J.M. Watt and M.G. Breyer-Brandwijk,
Med. Pois. Pl. S. E. Africa (1962) 745-751). In Uganda, where it is almost certainly
naturalized, it is a particularly valuable source of timber for building poles, as these
Melia in Asia & Pacific 61
are reputed to be immune to the attacks of termites (see Styles in E. Afr. Agric. For.
J. 39 (1974) 416). Two cultivated forms have been given cultivar names:
i. cv. Floribunda
See L.H. Bailey Hort., Hortus Third (1976) 724. - M. floribunda Carr., Rev.
Hort. 44 (1872) 470 cum tab. - M. azedarach var. floribunda (Carr.) Morren,
Belg. Hort. 30 (1880) 176 cum tab. —- Type: France, Paris, cultivated at Jardin
des Piantes (?P).
A precocious form, flowering when only a few dm tall and used in bedding
schemes (and greenhouse decoration in Europe).
ii. cv. Umbraculifera, Texas Umbrella Tree
See L.H. Bailey Hort., Hortus Third (1976) 724. - M. azedarach var. um-
braculifera Knox in Gdner’s Monthly 27 (1885) 260 - M. azedarach f. um-
braculifera (Knox) Rehder, Bibliog. Cultivated. Trees, Shrubs (1949) 387. -
Type: United States, Texas, trees discussed by Knox (?not preserved).
M. azedarach var. umbraculiformis Hort. ex Berck. & Bailey, Cycl. Amer. Hort.
2 (1900) 1001, ¢. 1387, ?sphalm. vel mut. pro praec.
A mutant with a flattened crown of branches. It apparently arose in Texas in the
nineteenth century and is widely planted in the southern United States as a street
tree.
Acknowledgements
I am indebted to Frank White, Terry Pennington and Brian Styles for their
comments on a manuscript version of this paper and to Charlie Jarvis at the British
Museum (Natural History) for his help in the tortuous matter of the typification of
M. azedarach, set out in Apendix II. I am grateful to Professor A.F.L. Beeston and
to Mr. R. Dawson for their guidance in the Arabic and Chinese literature respective-
ly. The bulk of the work of this paper was carried out at the Rijksherbarium,
Leiden, where I was supported by ZWO*, the Flora Malesiana Foundation and
Greshoff’s Rumphius Fund.
Reference
Harms, H. (1940), Meliaceae. Jn A. Engler & K. Prantl, Nat. Pflanzenfam., ed. 2,
19b1: 1-172.
* Zuiver Wetenschappelijke Onderzoek (Netherlands Organization for the Advancement of Pure
Research).
62 Gard. Bull. Sing. 37(1) (1984)
APPENDIX I
NOMINA DUBIA VEL EXCLUDENDA
Antelaea azadirachta (L.) Adelb., Blumea 6 (1948) 315 = Azadirachta indica A. Juss.
A. canescens Cels ex Heynh., Nom. Bot. Hort. 1 (1840) 38 = sphalm. pro Anthelia (‘Anthelea’)
canescens Cels, Cat. Cult. Cels Prix Cour. 1842 (1842) 8, i.e., ? Epipremnum sp. (Araceae).
Azedarach edulis Nor., Verh. Bat. Gen. 5, ed. 1 (1791) art. 4, 5 ‘edule’, nom. nud. = Sandoricum koet-
Jape (Burm. f.) Merr.
A. elegans (Seem.) O. Ktze, Rev. Gen. 1 (1891) 110 = Koelreuteria elegans (Seem.) A.C. Sm.
(Sapindaceae).
A. excelsa (Jack) O. Ktze, l.c. = Azadirachta excelsa (Jack) Jacobs.
bh
. fraxinifolia Moench, Meth. Suppl. (1802) 58, nom. superfl. pro Melia azadirachta L., = Azadirachta
indica A. Juss.
A. nigra Nor., l.c. ‘nigrum’, nom. nud. = Dysoxylum excelsum BI. (fide Hassk., Cat. Hort. Bog. (1844)
221).
A. ramiflora Nor., l.c. ‘ramiflorum’, nom. nud. = D. parasiticum (Osb.) Kost.
A. recisa Nor., |.c. ‘recisum’, nom. nud. = Toona sureni (Bl.) Merr. (Norona drawing at BM).
A. tomentosa (Roxb.) O. Ktze, Rev. Gen. | (1891) 110 = Chisocheton tomentosus (Roxb.) Mabb.
Melia azadirachta L., Sp. Pl. (1753) 385 = Azadirachta indica A. Juss.
M. baccata Wall., Cat. (1829) 1256, nom. nud. = seq.
M. baccifera Roth, Nov. Pl. Sp. (1821) 215 = Cipadessa baccifera (Roth) Miq.
M. elegans Seem., Fl. Viti. (1865) 36 = Koelreuteria elegans.
M. excelsa Jack, Mal. Misc. 1 (1820) 12 = A. excelsa.
M. fraxinifolia Salisb., Prodr. (1796) 317, nom. superfl. pro M. azadirachta, = A. indica.
M. iloilo Blanco, Fl. Filip. ed. 2 (1845) 241 = Aglaia sp. (A. iloilo (Blanco) Merr.).
M. indica (A. Juss.) Brandis, For. Fl. Ind. (1874) 67, nom. superfl. pro M. azadirachta, = Azadirachta
indica.
M. integerrima Buch.-Ham. in Trans. Linn. Soc. 17 (1835) 231 = Trichilia connaroides (W. & A.) Bentv.
(Heynea trijuga Roxb. ex Sims). See Mabberley in Taxon 26 (1977) 530.
M. koetjape Burm.f., Fl. Ind. (1768) 101 = S. koetjape.
M. latifolia Griff., Itin. Notes (1848) 402, sphalm. pro Melica latifolia Roxb. ex Hornem., =
Thysanolaena latifolia (Roxb. ex Hornem.) Honda emend. Mabb. (7. maxima), Gramineae. See
Mabberley in Taxon 33 (1984) 437.
M. montana Herb. Madras ex Wall., Cat. (1832) 214, n. 1256D, nom. nud., = C. baccifera.
M. neilgherrica Walp., Rep. 4 (1857) 551, sphalm. pro Munronia neilgherrica Wight = Munronia pin-
nata (Wall.) Theob.
Melia parasitica Osb., Dagb. Ostind. Resa (1757) 278 = D. parasiticum (Osb.) Kost.
M. parviflora Moon, Cat. Pl. Ceyl. (1824) 35 = C. baccifera.
M. pendula Reinw. ex Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 29, nom. in synon., = Chisocheton
patens Bl.
M. penduliflora Wall., Cat. (1829) n. 1255 = C. penduliflorus Planch. ex Hiern.
M. pinnata Stokes, Bot. Mat. Med. 2 (1812) 482, nom. superfl. pro M. azadirachta, = A. indica.
M. pubescens Reinw. ex Koord. & Val., Bijdr. Boom. Java 3 (1896) 91, nom. in synon., = Dysoxylum
nutans (Bl.) Miq.
M. pumila Moon, Cat. Pl. Ceyl. (1824) 35 = Munronia pumila Wight.
Melia tomentosa Roxb., Hort. Beng. (1814) 90, nom. nud., Fl. Ind. ed. Carey 1 (1832) 394 =
Chisocheton tomentosus.
Melia in Asia & Pacific 63
APPENDIX II
THE TYPIFICATION OF MELIA AZEDARACH
During his stay in coastal Ceylon in the 1670s, the German-born, Dutch botanist
and physician, Paul Hermann (1646-1695) collected specimens referable to Melia
azedarach, the widely cultivated tree known as Persian Lilac, bead-tree or, in the
Malay Peninsula, mindi kechil. In Hermann’s posthumous Musaeum zeylanicum
(1717), based on the specimens mounted or loose in his book herbarium, is (p. 3)
Panukohumba (Azedarach fructu polypyreno), mounted in vol. 1, fol. 10, and
Kirikohomba (p. 67, Arbor fraxinifoliis flore caeruleo CBP, Azedarach Dod., i.e.,
the widespread form of Persian Lilac long known in Europe), unmounted at that
time.
Before Linnaeus was to deal with them in his Flora zeylanica (1747), based on the
Hermann collection, he replaced the long established Azedarach by Melia, a Greek
word for the Manna Ash, Fraxinus ornus L. (Oleaceae), which has leaves reminis-
cent of Azadirachta indica A. Juss. (Melia azadirachta .) for they are simply
pinnate whereas those of M azedarach are doubly so. Melia was first published in
his Hortus cliffortianus (1738, p. 161), where the Persian Lilac is called Melia foliis
decompositis. As a variety of this he cites another cultivated plant, from Sri Lanka,
illustrated in Jan Commelijn’s Horti medici amstelodamensis | (1697) 147, t. 76 and
a name quoted by Commelijn, Jacob Breyne’s Azadirachia indica. . . flore albo sub-
caeruleo purpurascente majore (Prodromus fasciculi rariorum plantarum 2 (1689)
21). Indeed it was Breyne who was the first to separate these two varieties in print,
giving the widespread ‘Syrian’ plant, which he considered to be Avicenna’s (i.e.
Husain Ibn ’Abd Allah’s) Azadiracht, the name ending ‘.. .flore caeruleo majore’.
In the synonymy of the Ceylon plant, Breyne included Azedarach floribus albis
sempervirens of Hermann, based on his examination of garden material, Hermann
had shown him in the Hortus Botanicus at Leiden, and citing Hermann, Horti
academici Lugduno-batavi catalogus (1687) 652 but without the synonyms in
Hermann’s book, for those are referable to neem, Azadirachta indica, for it is
possible that Hermann mistook the garden material from Sri Lanka, introduced into
the Netherlands in the 1680s, for Azadirachta which he had collected in Ceylon, his
specimen in his book herbarium preserved at Leiden being inscribed Azedarach
floribus albis (fol. 120). The neem was not to be grown in Europe for some decades
yet and Breyne seems right, therefore, in having shorn Hermann’s garden name of
its synonyms.
When Linnaeus gained access to Hermann’s main Ceylon herbarium, now
preserved at the British Museum (Natural History), he included Panukohumba, the
Ceylon plant, and not Kirikohomba, the Persian Lilac, in the typical form of his
species (number 162), Melia foliis duplicato-pinnatis in Flora zeylanica. Another
synonym of this typical form is his own Melia foliis decompositis from Hortus clif-
fortianus (i.e. the Persian Lilac) as well as Hermann’s garden plant, again without
synonyms, and the references to Breyne and Commelijn. In this case, the variety of
the species is the commonly cultivated Syrian plant in which is included Kirikohom-
ba. By the time of Species plantarum (1753), however, Linnaeus seems to have
i
64 Gard. Bull. Sing. 37(1) (1984)
changed his mind yet again, making Commelijn’s (and therefore Breyne’s and
Hermann’s garden plant) the atypical variety of the species under the name var.
sempervirens, which epithet is taken from the cited Azedarach sempervirens &
florens of Tournefort, who (/nst. 1 (1700) 616) included the earlier references too.
The typical plant is Persian Lilac, again with the Hortus cliffortianus name but the
phrase name next to the specific epithet azedarach is a new one, indicating a possible
rethink of the circumscription by Linnaeus. However, it is written curiously, ‘Melia
foliis bipinnatis Fl. Zeyl. 162’, though that name does not appear in Flora zeylanica
but may be a mere rephrasing. Had he merely used the Hortus cliffortianus
reference then the type specimen would clearly be that collected from live plants at
George Clifford’s garden at de Hartecamp near Haarlem in the Netherlands and
now preserved in the British Museum (Natural History). As he did not amend this
in later editions, it seems on the face of it that it would be wise to take Flora
zeylanica into account, particularly as the only specimen in Linnaeus’s own her-
barium (543/1), although the Persian Lilac, has no indication of its age or origin.
The only other ‘Melia’ specimens there are (2) Azadirachta indica and (3) ?San-
doricum koetjape (Burm. f.) Merr. Of the materials cited in Flora zeylanica, only
Hermann’s Kirikohomba i.e. the ‘Syrian’ plant collected by Hermann in Ceylon, was
a specimen and it would have seemed to have been wise to select it as lectotype. The
var. sempervirens is Hermann’s plant Panukohumba from Ceylon which was later
introduced to the Netherlands and confused by Hermann with Azadirachta. Indeed
the common name for the latter in Ceylon is still Kohumba. This was identified as
var. sempervirens by Trimen (J. Linn. Soc. Bot. 24 (1888) 141), though Rechinger
in Flora iranica 133 (1978) 2 has this plant as the type of the species. This lecto-
typification seems not have taken the complicated history of the cultivated plants
into account. Not surprisingly then, the confusing actions of Linnaeus have misled
several workers and, sadly, Hermann’s unmounted Kirikohomba cannot be found
and there is no duplicate at Leiden or in the Sherard or DuBois herbaria at Oxford,
both herbaria with a number of Hermann specimens. Of the elements in the pro-
tologue of M. azedarach, that referring to Hortus cliffortianus is supported by
specimen 161/1 seen by Linnaeus and I therefore propose that it be the lectotype.
Axillary Buds of Some Tropical Trees
S. K. LEE AND A. N. RAO
Botany Department, National University of Singapore
Abstract
The presence and development of axillary buds are important in shoot and plant growth, especially
in trees. As very little information is available on the axillary buds of tropical trees, about 100 local tree
species, growing in the Botanic Gardens, Nature Reserves and along waysides, were investigated. The
occurrence, morphology and size variations of the axillary buds are discussed.
Introduction
In a growing shoot system, the relationship between the terminal and axillary buds
is important in terms of origin, location, number and the relative role of apical
dominance. The general information regarding the origin and development of
axillary buds is available in the basic reference works on plant development and
anatomy (Cutter, 1972; Esau, 1965; Fahn, 1967). While analysing the architecture
of tropical trees, Hallé et a/. (1978) described the developmental variations noticed
in the axillary buds of a few dicotyledonous members. The available data on
tropical trees is very scanty or almost nil, considering the large number of tropical
tree species available in SE. Asia (Corner, 1952). It is said that the axillary buds vary
very widely among the species of a genus or sometimes even among the individuals
of the same species or, very rarely even within a tree (Hallé ef a/, 1978). It is also
well known that the development and the growth behaviour of the apical and lateral
buds determine the shape and the architecture of the shoots and eventually of the
tree (Koriba, 1958). The present paper summarises the morphological characteristics
of the axillary buds of some 100 species of tropical trees, including their occurrence,
number, position, prominence and other related characters.
Materials and Methods
Young branches, up to a length of ten nodes, were collected at random from
common and easily available trees growing along roadsides, the Botanic Gardens,
primary and secondary forests in Singapore. The leaves were removed to expose the
buds and these were examined under the binocular microscope to determine their
shapes and sizes. The node that was visible, distinctive and nearest to the apex was
considered as the first node and the others counted in basipetal order (fig. 1). The
relative prominence of the buds at different nodes on the axis was noted. The
hundred species studied belong to 31 dicotyledonous families, Podocarpaceae and
Gnetaceae.
Observations and Discussions
Buds were present at the axils of most of the species studied except in the case
of Brownea grandiceps Jacq. and Plumeria sp. where they were indistinct or absent.
65
66 Gard. Bull. Sing. 37(1) (1984)
The unit of study for each species was the shoot with 10 nodes. The relative
prominence of buds at different nodes varied. In the majority of them, almost 70%,
the buds were present up to the ninth node. Their absence at other nodes could be
due to early bud abscission instead of non-formation, since the bud scars were
obvious in most of the shoots studied (table 3). The sequential development of buds
in relation to the total shoot and individual internodal length should be interesting
(Hallé et al., 1978).
In all the standard works referred, very little or no mention was made on the
morphology and characteristic features of axillary buds (Goebel, 1900; Corner,
1952; Symonds, 1958; Clowes, 1961; Menninger, 1962; Symington, 1974; Palmer
and Pitman, 1972; Bernatzky, 1978; Hallé et a/., 1978; Kunkel, 1978; Hora, 1981).
It was also revealing that there is no well established terminology to describe the
axillary buds, even though other factors that affect shoot growth like mineral
nutrition, water availability, soil conditions and others are well considered.
Bud types
Among the tree species studied presently, buds with different sizes and shapes
were encountered. They were broadly classified into eight morphological types. The
relative shape and outline of the bud was the only criterion used in recognising
the eight bud types. The flattened, bulbous, pear-shaped and round ones were
omewhat radially symmetrical with almost a circular outline whereas the linear,
oblong and triangular ones were somewhat bilaterally symmetrical (fig. 2). All the
ny) R
‘__ node 1 EA a
node 10
Fig. 1. Young branch with opposite (/efr) and alternate leaf position (right) showing the first 10 nodes
labelled sequentially (diagrammatic).
Flattened Linear Buibous
Pear Shape
i
tas {
‘ ¢ 4
eG A: Frontal View
Vv ie
B: Side View
Fig. 2. Diagrammatic representation of the different shapes of axillary buds.
others that did not fit into the above types were grouped under the irregular type.
The various types present and the frequency of their occurrence were also noted.
The triangular buds (fig. 2; plate 1, h and k) were of most common occurrence,
followed by linear (plate 1, b and c), oblong (plate 1, f), buibous (plate 1, d) and
other types. All of them are diagrammatically illustrated in figure 2 and the relative
frequencies are shown in table 1.’
67
68 Gard. Bull. Sing. 37(1) (1984)
The bud morphology, quantitative and qualitative aspects are not studied so far
in any great detail, especially of tropical trees (Corner, 1952; Opeke, 1982). Lubbock
(1899) considered the structure of buds of about 25 temperate trees and shrubs
including three Gymnosperms. The buds of each species were described but no bud
types were recognised. Greater emphasis was laid on the structures that protect the
bud and the bud emergence after wintering. Other papers published occasionally
described the buds in individual plants like cotton, sweetgum, bamboos and others
(Mannery and Ball, 1959; Kormanik and Brown, 1967; McClure, 1976).
Single and Multiple Buds
In the majority of the trees or woody plants, occurrence of a single axillary bud
is common. Of the 100 species presently studied, 17 of them had multiple buds at
the nodes (table 2).
The species with multiple buds are grouped in the increasing order of their
numbers.
a) Species with 1-2 axillary buds per node
Annona muricata L. Hibiscus tiliaceus L.
Annona reticulata L. Mimusops elengi L.
Annona squamosa L. Lithocarpus urceolaris (Jacq.) Merr.
Delonix regia (Boj. ex Hk.) Raf. Samanea saman (Jacq.) Merr.
Gliricidia sepium (Jacq.) Kunth ex Walp.
b) Species with 2-3 axillary buds per node
Jacaranda obtusifolia HBK ssp. rhombifolia (Meij.) Gent.
c) Species with 3-4 axillary buds per node
Spathodea campanulata Beauv.
Peltophorum pterocarpum (DC.) Back.
Nephelium lappaceum L.
d) Species with 4-5 axillary buds per node
Tabebuia pallida Miers Erythrophleum suaveolens (Guill. & Perr.) Bren.
Acacia auriculiformis A. Millettia atropurpurea Bth.
Cunn. ex Bth.
The 100 species examined belonged to 74 genera of 33 families. Four or more
genera were included in Bignoniaceae, Dipterocarpaceae, Leguminosae and Myrtaceae
and in others, 1, 2, or 3 genera were involved (table 2). The family of Leguminosae
had the largest representation with about 16 genera and 23 species. In the members
of these families, the shape of the buds varied among the different genera, the
different species and sometimes even between the allied species in genera such as
Annona, Cassia, Shorea, Nephelium, Tabebuia and Podocarpus. \n contrast, the
Axillary buds of some tropical trees 69
Table 1
The shapes of axillary buds in some tropical trees.
Bud Shape Number of Species
Bulbous 16
Flattened 4
Irregular 2
Linear 18
Oblong 16
Pear 7
Round 5
Triangular 30
Buds indistinct 2
100
Note. The above observations were made on the first 3 nodes where the shape of the distinct buds was
relatively consistent. Owing to age, the buds in the older nodes in many species were either
detached or the shapes had changed due to emergence. Plate 1 a to q illustrate some of the shapes
described.
buds were of the same shape among the species of Eugenia, Ficus, Artocarpus and
Bauhinia.
Hallé et al. (1978) also observed that multiple buds are quite common with
tropical woody plants such as Coffea sp. The frequency of tropical trees species
having multiple buds was however not mentioned. The morphogenetic implication
of these buds has so far been little considered, especially by developmental
biologists. The buds at each node should be further studied as the differential
developmental potential of each number of the multiple bud complex will also
influence the ultimate architecture of the tree (Hallé et a/., 1978). Some preliminary
observations made with some tropical plants showed that the pattern of response by
the multiple buds, varied from species to species (Varossieau, 1940; Moens, 1963).
In Coffea, for example, the distal bud of each leaf pair on orthotropic shoots
usually grows out as a precocious branch, the others persist as reserve buds. In the
case of Dipterocarps, the dominant axillary buds would often grow in a plagiotropic
pattern and in some instances, the axillary buds which may be indistinct, would
grow to form orthotropic shoots (Ng, 1976).
The distinct variation in numbers, shapes and sizes of the axillary buds among the
species within a genus or, between the genera of a family, may have some taxonomic
importance. To-date, the morphological features of axillary buds are seldom or not
at all used for either identification or species classification (Goebel, 1900; Corner,
1952; Whitmore, 1972; 1973; Cockburn, 1976; Ng, 1978).
Very often, the identification of plants are confirmed through the study of flower
characters. However, as most tropical forest trees flower either rarely or infre-
quently, confirmation through flower features may be a problem. In fact, some
tropical trees do not even flower for years (McClure, 1966; Rao, 1973). Therefore,
HO) oe Gard. Bull. Sing. 37(1) (1984)
having at one’s disposal an additional vegetative character such as the bud mor-
phology may be immensely useful.
Buds, prior to their emergence, seem to be rigid and uniform in their shapes and
other morphological characteristics. They may not be so variable as the leaves in
sizes and shapes. Stace (1980) has already mentioned that vegetative characters such
as leaves of higher plants, are often looked upon as risky evidence because often
similar morphological features are found in quite unrelated plants. Therefore, as
many other vegetative characters as possible should be included to give an accurate
identification and, buds also can be used in terms of their shape, size and number.
In some instances, the number of buds varied from node to node on the same axis.
In Tabebuia pallida, multiple buds were seen at some of the younger nodes, and at
the older ones, the buds were indistinct. Where multiple buds were present, they
varied in number from 2 to 5}'but in some like Erythrophleum suaveolens, five buds
were present consistently at most leaf axils. The arrangement of buds varied among
the species with multiple buds. Clustered arrangement of buds was seen in Tabebuia
pallida and in Erythrophleum suaveolens; the buds were arranged in basipetal order.
The sizes of the multiple buds within a single axil differed greatly and the largest
of the lot was dominant. It developed into a shoot when conditions favoured. The
smaller ones remained either dormant or frizzled and dropped off.
Regarding their conspicuousness, the buds decreased in prominence in the lower
nodes and this was the general pattern for the majority of the species studied (table
3). In most species the buds became detached following leaf abscission and a few
species were exceptional like Bixa orellana L. and Michelia alba DC. where the buds
were prominent even at the older nodes.
The largest axillary bud measured up to | cm in length was in Erythrophleum
suaveolens and the smallest, about 0.1 mm, found in Fagraea fragrans Roxb., was
barely visible even under the binocular microscope. The larger buds were found in
the younger leaf axils and this perhaps was due to the vigorous meristematic tissue
subtending the bud and the dominant nature of the younger bud itself. In
Peltophorum pterocarpum and Fagraea fragrans the dominant buds at nodes 1, 2
or 3 gave rise to lateral branches.
Apart from shape and size, the buds varied with regard to the number of scale
leaves that covered each bud, but on an average, 4-5 bud scales were present in most
of them. The bud scales were smooth in certain species as in Eugenia grandis Wight
$$
Plate 1. (facing page):
a, lateral view of a flattened bud of Pferocarpus indicus Willd.; b, frontal view of a linear bud
of Coccoloba uvifera; c, lateral view of a linear bud of Michelia champaca L.; d, frontal view
of a bulbous bud of Mangifera indica L.; e, lateral view of a bulbous bud of Eugenia grandis; f,
lateral view of oblong to triangular buds of Cinnamomum iners Reinw. ex BI.; g, frontal view
of a pear-like bud of Samanea saman; h, frontal view of a triangular bud of Bixa orellana.
Scale: I division = Imm.
71
Plate 1 cont
A.
Axillary buds of some tropical trees 73
(plate 1d) or covered with hairs as in Muntingia calabura L. Most of the buds were
brown in colour and some had the same colour as the bark of the tree. Some buds
were distinctly pink as in Coccoloba uvifera (L.) L. (plate 1b), and covered by very
broad stipules.
The architecture and form of a tree is heavily influenced by the growth response of
axillary buds (Corner, 1952; Hallé et a/, 1978; Koriba, 1958). The geometry of the
arrangement of the buds and the manner of the response determine the ultimate
form of the tree. In monopodial trees, the axillary buds are normally arranged in
regular whorls around the orthotropic shoot and the uniform response and growth
of these buds give rise to the monopodial pattern of branching. In contrast with the
monopodial trees, the response of the axillary buds of sympodial trees varies
tremendously. Hallé et a/, (1978) stated that the pattern of the nodes on which the
axillary buds are located, is extremely complex and diverse, thus resulting in a
variable response. Furthermore, the distribution and the kinds of lateral or axillary
buds can vary widely on the different parts of one plant. Because of these variations,
different patterns of branching are observed in sympodial trees (Koriba, 1958).
Buds are also of great value in vegetative propagation as they can serve as starting
units for mass propagation. De Fossard (1980) had shown that when nodes of
Eucalyptus ficifolia F. Muell. were cultured in culture media containing 5 »M IBA
and 2 uM BAP, several shoots were obtained. As a result, young nodal segments
containing the axillary buds were used widely as explants to induce multiple shoot
formation in vitro condition for many species of trees (Hutchinson, 1981; Lee and
Rao, 1981; Mascarenhas ef a/, 1981). It will therefore be beneficial for further
studies to be carried out to determine whether there is any correlation between the
prominence or the morphology of these buds and the ease with which they develop
into multiple shoots.
Plate 1. cont. (facing page):
i, frontal view of a rounded bud of Erythrina variegata L.; j, lateral view of an irregular bud
of Nephelium lappaceum; k, lateral view of a triangular bud of Citrus microcarpa Bunge; notice
the sharp thorn structure enclosing the axillary bud; /, m, lateral and frontal view of multiple
buds of Erythrophleum suaveolens; n, lateral view of multiple buds of Gliricidia sepium; buds
were mainly pear-shaped; o, frontal view of multiple buds of Acacia auriculiformis; buds were
linear to triangular in shape; p, lateral view of multiple buds of Peltophorum pterocarpum;
buds were mainly linear to pear-shaped; notice the leader bud being he largest of the 3 to 4 buds
present; q, lateral view of multiple buds of Millettia atropurpurea; buds were oblong to
triangular in shape; notice the larger leader bud of the 2 present.
Scale: ] division = Imm.
74 Gard. Bull. Sing. 37(1) (1984)
Table 2
Number, shape and sizes of the axillary buds in the different taxa.
* The numbers are mentioned for those taxa which had multiple buds. All others had a single bud at
each leaf axil.
+ The sizes given for each species in this table refer to the smallest and the largest buds (length of)
the buds only), distinct at the nodes 1 to 10. All measurements were made only on buds before
emergence. Mean lengths of the buds were derived from nodes | to 4. Buds were somewhat indistinct
in certain species at subsequent nodes. Hence measurements were variable, e.g. Tabebuia pallida.
Family/Species Shape Sizes (mm)+ Mean (mm)
Anacardiaceae
1. Anacardium occidentale L. bulbous 0.5-1.0 0.7+0.2
2. Mangifera indica L. bulbous 0.5-1.0 0.8+0.2
3. Rhus succedanea L. triangular 2.0-3.0 2.5+0.5
Annonaceae
4. Annona muricata (\-2)* linear 1.0-4.0 2.441.3
5. Annona reticulata (1-2)* oblong 1.0-2.0 1.5+0.5
6. Annona squamosa (1-2)* irregular 1.0-2.0 1.5+0.5
7. Polyalthia longifolia (Sonn.) Thw. linear 4.0-8.0 $.8+1.7
Apocynaceae
8. Ervatamia dichotoma (Roxb.) Burk. round 0.1-0.5 0.2+0.1
9. Plumeria acuminata W.T. Ait. - - -
Bignoniaceae
10. Jacaranda (2-3)* linear to oblong 1.0-5.0 3.6+1.8
11. Spathodea campanulata (3-4)* linear to triangular 1.0-5.0 3.44+1.7
12. Tabebuia pallida (4-5)* irregular to linear 1.0-3.0 1.6+0.9
13. Tabebuia rosea (Bertol.) DC. bulbous 0.5-1.0 0.7+0.3
14. Tabebuia spectabilis (Planch. & round 0.5-2.0 N-Syse(0s7/
Lindl. ex Planch.) Nichols.
Bixaceae
15. Bixa orellana triangular 1.0-3.0 2.2+0.8
16. Cochlospermum religiosum (L.) Alston linear 1.0-4.0 iloi/ ae 8)
Capparidaceae
17. Crataeva religiosa Forst. f. triangular 2.0-4.0 Sil se)
18. Cratoxylon formosum (Jack) Dyer triangular 1.0-4.0 AN seit)
19. Cratoxylon pruniflorum Kurz bulbous 1.0-3.0 1.8+0.9
Celastraceae
20. Elaeodendron quadrangulatum Reiss. oblong 1.0-2.0 1.4+0.5
Combretaceae
21. Terminalia catappa L. triangular 3.0-6.0 Ae ele
Axillary buds of some tropical trees 75
Table 2 Continued
Family/Species Shape Sizes (mm)+ Mean (mm)
Dipterocarpaceae
22. Dryobalanops aromatica Gaertn. f. triangular 1.0-3.0 1.6+0.8
23. Hopea mengarawan Miq. bulbous 0.1-0.5 0.3+0.1
24. Shorea curtisii Dyer ex King bulbous 0.1-0.5 0.3+0.2
25. Shorea leprosula Miq. triangular 0.5-1.5 0.3+0.1
26. Shorea sumatrana (V. Sl. ex Foxw.) Sym. oblong 4.0-6.0 5.3+1.0
27. Vatica pallida Dyer triangular 0.1-1.0 0.6+0.4
Ebenaceae
28. Diospyros discolor Willd. triangular 2.0-4.0 oye) bees IAI
Euphorbiaceae
29. Antidesma bunius (L.) Spreng. triangular 0.5-1.0 0.6+0.3
30. Elateriospermum tapos Bi. bulbous 0.1-0.5 0.4+0.2
31. Macaranga triloba (Bl.) M.A. pear 3.0-6.0 4.4+1.5
Fagaceae
32. Lithocarpus urceolaris (1-2)* pear 1.0-2.0 1.4+0.5
Gnetaceae
33. Gnetum gnemon L. flattened 0.5-1.0 0.8+0.2
Guttiferae
34. Calophyllum inophyllum L. triangular 1.0-2.0 Hef/S=(l as
Lauraceae
35. Cinnamomum iners Reinw. ex BI. oblong to triangular 2.0-3.0 2.6+0.5
Leguminosae
36. Acacia auriculiformis (2-5)* oblong to linear 0.5-2.0 1.3+0.7
37. Albizzia falcata (L.) Back. linear 1.0-7.0 -
38. Andira inermis (W. Wight) HBK ex DC. linear 3.0-4.0 3.7+0.5
39. Bauhinia acuminata L. triangular 4.0-6.0 5.0+1.0
40. Bauhinia blakeana Dunn triangular 4.0-6.0 4.5+0.9
41. Bauhinia purpurea L. pear 2.0-5.0. 4.6+1.0
42. Brownea capitella Jacq. linear 2.0-6.0 3.4+2.1
43. Brownea grandiceps - = =
44. Cassia bakeriana Craib flattened 1.0-2.0 5-05
45. Cassia fistula L. triangular 2.0-6.0 4.6+0.8
46. Cassia multijuga Rich. linear 1.0-5.0 3321.6
47. Cassia sp. (hort. variety) triangular 2.0-4.0 3.2+1.0
48. Cassia spectabilis DC. oblong 1.0-3.0 1.9+0.9
49. Dalvergia oliveri Gamble & Prain flattened 1.0-2.0 1.3+0.5
50. Delonix regia (2)* pear 1.0-2.0 1.4+0.5
51. Erythrophleum suaveolens (3-5)* triangular to linear 1.0-10.0 G29 5s 4
52. Erythrina fusca Lour. bulbous to pear 2.0-4.0 3.0+0.8
53. Erythrina variegata L. round to triangular 1.0-3.0 Desj-et)
54. Gliricidia sepium (2)* pear to round 1.0-3.0 Posse
55. Millettia atropurpurea (3-5)* oblong 1.0-6.0 4.0+1.9
56. Peltophorum pterocarpum (2-4)* linear 1.0-5.0 3-3-5109
76
Table 2 Continued
Gard. Bull. Sing. 37(1) (1984)
Family/Species
Leguminosae cont.
57. Pongamia pinnata (L.) Pierre
58. Pterocarpus indicus Willd.
59. Samanea saman (1-2)*
Lecythidaceae
60. Barringtonia asiatica (L.) Kurz
Loganiaceae
61. Fagraea fragrans
Lythraceae
62. Lagerstroemia speciosa (L.) Pers.
Magnoliaceae
63. Michelia champaca L.
Malvaceae
64. Hibiscus tiliaceus (1-2)*
Meliaceae
65. Khaya grandiflora C. DC.
66. Sandoricum koetjape (Burm. f) Merr.
Myrsinaceae
67. Ardisia elliptica Thunb.
Myrtaceae
68. Callistemon citrinus (Curtis) Stapf
69. Eugenia aquea Burm. f.
70. Eugenia grandis
71. Eugenia javanica Lmk
72. Eugenia longiflora (Presl.) F. Vill.
73. Eugenia malaccensis L.
74. Eugenia michelii Lmk
75. Melaleuca cajuputi Powell
76. Psidium guajava L.
77. Rhodamnia cineria Jack
Podocarpaceae
78. Podocarpus koordersii Pilg.
79. Podocarpus neriifolius D. Don
80. Podocarpus polystachyus R. Br. ex Mirb.
Polygonaceae
81. Coccoloba uvifera
82. Triplaris americana L.
Sizes (mm)+ Mean (mm)
Shape
flattened 1.0-2.0
flattened 1.0-3.0
pear to irregular 1.0-6.0
bulbous 2.0-3.0
bulbous 0.1-0.5
linear 2.0-6.0
linear 5.0-15.0
oblong 1.0-4.0
round to oblong 0-1.0
round to oblong 1.0-3.0
round 3.0-6.0
triangular 5.0-8.0
bulbous 0.5-1.0
bulbous 0-1.0
bulbous 0.5-1.0
bulbous 0.5-2.0
bulbous 0.5-1.0
linear 2.0-6.0
pear 4.0-6.0
oblong 2.0-3.0
triangular 2.0-4.0
bulbous 1.0-2.0
round 1.0-3.0
oblong 1.0-2.0
triangular to linear 4.0-8.0
oblong 1.0-3.0
1.4+0.5
2.3+0.5
4.0+2.1
2.4+0.5
0.4+0.2
4.2+1.5
10.1+3.0
2.34+1.4
0.9+0.3
1.9+0.9
4.5+1.2
Axillary buds of some tropical trees
Table 2 Continued
77
Family/Species Shape Sizes (mm)+ Mean (mm)
Rutaceae
83. Atalantia spinosa (Willd.) Tanaka linear 1.0-3.0 2.0+1.
84. Citrus grandis (L.) Osb. triangular 2.0-6.0 a2 ae
Salicaceae
85. Salix sp. linear to triangular 5.0-9.0 $.6+1.7
Sapindaceae
86. Arfeuillea arborescens Pierre oblong 2.0-3.0 2.6+0.5
87. Filicium decipiens (Wright & Ann.) Thw. oblong 2.0-4.0 35-120
88. Nephelium lappaceum (1-4)* irregular 1.0-3.0 2.0+1.0
89. Dimocarpus longan Lour. var. oblong 1.0-2.0 1.3+0.6
malesianus Leenh.
Sapotaceae
90. Achras sapota L. oblong to irregular 1.0-2.0 1.2+0.4
91. Mimusops elengi L. (2)* triangular 2.0-5.0 35°213
92. Palaquium obovatum (Griff.) Engl. bulbous 0.5-1.0 0.8+0.2
Saxifragaceae
93. Brexia madagascariensis (Lmk) Thou. triangular 2.0-4.0 2.8+0.8
Tiliaceae
94. Muntingia calabura linear 0.5-1.0 0.8+0.2
Urticaceae
95. Artocarpus integer (Thunb.) Merr. linear 2.0-3.0 2.4+0.5
96. Artocarpus gomeziana Wall. linear 2.0-5.0 3.7+1.4
97. Artocarpus heterophyilus Lamk. oblong 1.0-3.0 1.5+0.8
98. Ficus benjamina L. triangular 2.0-4.0 3.0+1.0
99. Ficus elastica Roxb. ex Hornem. triangular 2.0-6.0 2.8+0.8
100. Ficus retusa L. triangular 1.0-3.0 1.6+0.8
Table 3
The prominence of the axillary buds in relation to their positions at different nodes.
Relative prominence of buds
Buds prominent in all 10 nodes
Buds prominent up to 7th to 9th node
Buds prominent up to 4th to 6th node
Buds prominent only in first 3 nodes
Buds not visible
Number of species exhibiting
the pattern of prominence
Note. All buds above the size of 2mm, which were visible to the naked eye,
29
39
18
12
2
100
were Classified as prominent.
78 Gard. Bull. Sing. 37(1) (1984)
Literature Cited
Bernatzky, A. (1978). Tree Ecology and Preservation. Elsevier & Co., Amsterdam.
Clowes, F. A. L. (1961). Apical Meristems. Blackwell, Oxford.
Cockburn, P. F. (1976). Trees of Sabah, Vol. 1. Sabah Forest Record no. 10. Forest
Department, Sabah.
Corner, E. J. H. (1952). Wayside Trees of Malaya. The Government Printer,
Singapore.
Cutter, E. G. (1972). Regulation of branching in decussate species with unequal
lateral buds. Ann. Bot. 36: 207-220.
Esau, K. (1965). Plant Anatomy. John Wiley & Sons, New York.
Fahn, A. (1967). Plant Anatomy. Pergamon Pubs., New York.
Fossard, R. A. de (1980). Tissue Culture Propagation No. 10. University of Hawaii,
Harold L. Lyon Arboretum, Hawaii.
Goebel, K. (1900). Organography of Plants, especially the Archegoniate and Sper-
matophytes. Part I: General Organography. Clarendon Press, Oxford.
Hallé, F., R. A. A. Oldeman and P. B. Tomlinson (1978). Tropical Trees and
Forests, an architectural analysis. Springer-Verlag, Berlin.
Hora, B. (ed.) (1981). The Oxford Encyclopaedia of Trees of the World. Univ.
Press, Oxford.
Hutchinson, J. F. (1981). Fruit tree propagation in-vitro, pp. 113-120. Jn Proc.
COSTED Symp. on Tissue Culture of Economically Important Plants, Ed. A. N.
Rao, Singapore.
Koriba, K. (1958). On the periodicity of tree-growth in the tropics, with reference
to the mode of branching, the leaf-fall, and the formation of resting buds. Gard.
Bull. Sing. 17: 11-81.
Kormanik, P. P. and C. L. Brown (1967). Root buds and the development of root
suckers in the sweetgum. Forest Sci. 13: 338-345.
Kunkel, G. (1978). Flowering Trees in Subtropical Gardens. Dr. W. Junk. b.v., The
Hague.
Lee, S. K. and A. N. Rao (1981). In-vitro plantlet development in tropical trees —
Calophyllum inophyllum and Eugenia grandis, pp. 185-190. In COSTED Symp.
on Tissue Culture of Economically Important Plants, Ed. A. N. Rao, Singapore.
Lubbock, J. (1899). Buds and Stipules. Kegan Paul, Trench, Trubner & Co. Ltd.,
London.
Mauney, J. R. and E. Ball (1959), The axillary buds of Gossypium. Bull. Torrey
Bot. Club 86: 236-244.
Mascarenhas, A. F., P. K. Gupta, V. M. Kulkarni, U. Melta, R. S. Jyer, S. S.
Khuspe, and V. Jagannathan (1981). Propagation of trees by tissue culture, pp.
Axillary buds of some tropical trees 79
175-179. In Proc. COSTED Symp. on Tissue Culture of Economically Important
Plants, Ed. A. N. Rao, Singapore.
McClure, F. A. (1976). The Bamboos, a Fresh Perspective. Harvard Univ. Press,
Cambridge, U.S.A.
McClure, H. E. (1966). Flowering, fruiting and animals in the canopy of a tropical
rain forest. Malay. Forester 29: 182-203.
Menninger, E. A. (1962). Flowering Trees of the World for Tropics and Warm
Climates. Hearthside Press Incorp., New York.
Moens, P. (1963). Les bourgeons végétatifs et génératifs de Coffea canephora
Pierre. Etude morphologique et morphogénétique. La Cellulare 63: 165-244.
Ng, F. S. P. (1976). Responses to leader-shoot injury in Shorea platyclados. Malay.
Forester 39: 91-100.
. (ed.) (1978). Tree Flora of Malaya, Vol. 3. Longmans, Kuala Lumpur.
Opeke, L. K. (1982). Tropical Tree Crops. John Wiley & Sons, New York.
Palmer, E. and N. Pitman (1972). Trees of Southern Africa, covering all known
indigenous species in the Republic of South Africa, South West Africa, Botswana,
Lesotho and Swaziland. A. A. Balkema, Cape Town.
Rao, A. N. (1973). Studies on growth of certain trees in Singapore, pp. 87-101. Jn
Proc. of the Pacific Science Association, Proc. of the Conf. on Planned Utilisa-
tion of the Lowland Tropical Forests, Ed. P. Soeratno, NBI and Biotrop, Bogor.
Varossieau, W. W. (1940). On the development of the stem and the formation of
leaves in Coffea species. Ann. Jard. Bot. Buitenzorg 50: 115-198.
Whitmore, T. C. (1972). (ed.) Tree Flora of Malaya, Vol 1. Longmans, Kuala
Lumpur.
. (1973). (ed.) Tree Flora of Malaya, Vol. 2. Longman, Kuala Lumpur.
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Physical Effects of Soil Compaction and Initial Growth of
Acacia pycnantha (Leguminosae) in a Clay-loam Soil
THAI WU FOONG'! & NORMAN WELLS?
‘Botanic Gardens, Parks & Recreation Department, Singapore
“Soil Bureau, D.S.I.R., Private Bag, Lower Hutt, New Zealand
Abstract
An ornamental plant, Golden Wattle (Acacia pycnantha) was grown in a clay loam subsoil sup-
plemented with adequate amounts of inorganic fertilizer and then compacted by four methods, namely,
light finger pressure, rubber hammer, steel bar and machine pressure. Dry matter accumulation and
rooting behaviour after four months growth in pots under glasshouse conditions were studied in relation
to the degree of soil compaction. Influences of compaction on such physical properties as dry bulk
density, penetration resistance, total porosity, oxygen diffusion rate and moisture content were also
investigated. Plant growth and soil penetration resistance were significantly related to the level of
compaction. Amelioration by addition of a medium-size grade of sand on a 50% volume basis before
applying compaction reduced the soil strength substantially and allowed the plant to grow normally.
Introduction
Soil compaction is a problem confronting urban horticulture in the Republic of
Singapore. Construction involving standard engineering procedures for building
roads and buildings, reclamation from the sea using clayey subsoils that lack organic
matter and the effects of recreational activities on school fields and in parks, often
lead to excessive soil compaction. In line with the Singapore Government’s ‘greening’
policy, the Parks & Recreation Department is required to establish vegetation in
planting holes frequently impounded by compact subsoil, typified by a myriad of
adverse physical conditions such as impeded drainage and high soil strength.
Consequently, root-bound vegetation is susceptible to root-rot during the wet
seasons (Raghavan ef a/., 1982 and Vigier et a/l., 1983).
The adverse effects of compaction on soil physical properties have been extensively
reviewed (Barley & Greacen, 1967; Cannell, 1977; Drew & Goss, 1973; Greacen &
Sands, 1980; Lal & Greenland, 1970 and Ruark ef a/., 1982).
The subject of this trial, Acacia pycnantha Benth., is much valued as an ornamental
plant in New Zealand. In addition, its bark is a source of tannins and gum. This plant
is deep-rooting and hardy, and can thrive under severe natural conditions (Audus,
1934). The objectives of this glasshouse trial were to study the rooting configuration
of Golden Wattle under various levels of soil compaction, to determine the growth
limiting levels of soil compaction and to evaluate the ameliorative effects of different
levels of two anti-compaction materials, i.e. sand and sewage sludge, incorporated
in the potting medium prior to artificial compaction. If the findings prove signifi-
cant and beneficial, then they could be adapted for practice in Singapore.
81
82 Gard. Bull. Sing. 37(1) (1984)
Table 1
Summary of treatments.
*Treatment Soil medium Mode of compaction
1 Clay loam Light aes pressure (uncompacted
contro
2 Clay loam Tapping with a rubber hammer
3 Clay loam Hitting with a steel bar
4 Clay loam Mechanical pressure of 5 tonnes applied
intermittently with a MTS stiff loading
frame
5 Pure sand Hitting with a steel bar
6 Clay loam + sand Hitting with a steel bar
(80:20 v/v)
7 Clay loam + sand Hitting with a steel bar
(50:50 v/v)
8 Clay loam + sludge Hitting with a steel bar
(80:20 v/v)
9 Silty pan material No treatment — Natural reconstitution
back into a hard pan
* There were 6 replicates in each treatment except for treatment 9 which was replicated twice.
Table 2
Effects of compaction on dry bulk density, total porosity, soil strength and plant dry matter.
Treatment 1 2 3 4 5 6 7 8
(Control)
Parameters
+Dry bulk density (g cm7’) 1.07 1.13 1.29 32) 1.22 1.31 1.22 1.29
+ Total porosity (%V) 60.7 58.6 52.6 51.4 54.0 51.5 54.8 Biles
Shoot dry weight (g plant') 1.46a 0.65be 0.72b 0.07**c O* 0.38be 1.44a O*
+ Root dry weight (g plant) 0.54 0.27 0.24 0.03 O* 0.15 0.37 O*
Penetration resistance (bar)
At 0-1 cm 2.46e 21.14d — 39.20b 54.80a — 34.69be 5.4le 29.76¢
At 1-2 cm 3.69e 31.61d 52.52b >100.00a — 40.02c 6.74e 32.02d
At approximately 10 cm 5.752 24.63e 75.54b >100.00a — SSS 15.48f 47.62d
Treatment 1-8 have reference in Table 1.
+ Average of two measurements.
* Complete mortality.
— Very low, not measured
**Low value due to mortality of 3 plant replicates.
Values in each row if not followed by the same letter are significantly different as judged by the DMR test at P < 0.05.
Growth of Acacia pycnantha in compacted soil 83
Materials and Methods
The soil compacted in this trial was taken from a buried reddish clay-loam layer
at a depth of 5 to 6 m from the surface of Paremata clay. It was collected from a
recently excavated area of quarry used for brick and pipe manufacture (Grid. Ref.
NZMS1 N160 473462). The undisturbed material had a dry bulk density of
1.44 g cm °, particle density of 2.72 g cm °, native moisture content of 23.7% and
a natural penetration resistance of 86 bar measured with a Chatillon Guage-R
penetrometer. On mechanical analysis, it was found to contain 52% clay, 45% silt
and 3% sand-size particles. Its pH was 5.6 and liming was not necessary. For this
study, a pan material was also collected from the natural B horizon and this had
an innate mechanical impedance of greater than 100 bar, particle density of
2.66 gcm © and a native moisture content of 12.8%. Mechanical analysis revealed
a composition of 9% sand, 53% silt and 38% clay. This material reconstituted
rapidly into a pan after initial disintegration for potting.
For anti-compaction purposes, a medium-size grade of sand, containing 3% silt-
size and 2% clay-size particles, was collected from the C horizon of a coastal aeolian
dune system at Waikanae. It had a natural moisture content of 4.4% and particle
density of 2.64 g cm °. Digested and dried sewage sludge was collected from a
nearby treatment plant and pulverized before mixing with the trial material.
The containers used were sections of PVC drain pipes of dimensions 150 x 300 mm
(diameter < height) with one end closed by a black perforated planting bag. The
composition of potting media and methods of compaction are summarized in table 1.
Each medium, in its naturally moist state, was packed layer by layer by one of 4
methods (table 1).
Seeds of A. pycnantha were germinated on moistened filter paper. When 5 mm
of root had developed, uniform seedlings were selected and implanted into a 1 cm’*
scarified zone in the middle of each pot. The replicates were arranged in randomized
blocks. The plants were watered by spraying twice daily according to their growth
requirements, and harvested after 4 months.
For treatments 1 to 8 (table 1), in two of the replicate pots an oxygen platinum
electrode was inserted to a depth of 3 cm to determine the oxygen diffusion rate
(Gradwell, 1972) and for measurements of penetration resistance using a probe of
5 mm in diameter. A second lot of two pots from each set were cored with the core
sampler to a depth of 1 to 4 cm in order to determine the amounts of water held
at a range of moisture tensions. Equilibrium times for these cores on the tension
plates varied as follows: 10 cm for 1 day; 25, 50 and 75 cm for 2 days and 100 cm
for 3 days. Dry bulk densities and porosities were determined from the same cores
according to Gradwell (1972). The particle densities were determined separately. The
remaining two pots per treatment were washed free of soil on a sieve to retain roots.
Plant tops were harvested from all replicates of each treatment. Both the root and
the shoot were subjected to dry matter determination after 4 months.
84 Gard. Bull. Sing. 37(1) (1984)
Results and Discussions
Results are presented in table 2, plates 1-4 and figures 1 and 2. Where possible,
data were analysed with the Duncan Multiple Range Test for significance.
The highest dry bulk density attained by using machine pressure on the clay-loam
soil was 1.32 g cm (table 2). This value was less than the native value of 1.44 g¢cm °
measured of the material in situ. A still higher value of 1.60 g cm °* had been recorded
for the B horizon of Paremata clay developed naturally over many wet and dry
seasons (New Zealand Soil Bureau, 1968). Repeated attempts to establish A. pyc-
nantha on a scarified zone on this pan material supplemented with inorganic
fertilizer were in vain (plate 1). The inability of the seedling to root in this pan
material guided the limits required for maximum dry bulk density in the pot trial.
The soil strength was directly and significantly proportional to the degree of
compaction at the 3 depths concerned (table 2). Such a relationship has been
confirmed by other researchers (Cannell, 1977 & 1982; Drew & Goss, 1973 & 1974;
Eavis, 1972 a&b; Heilman, 1981; Russell & Goss, 1974; Sands & Bowen, 1978 and
Zisa et al., 1980) It was noted that the penetration resistance generally increased with
depth (table 2). A soil strength of 25-55 bar in the rooting zone (0-10 cm) resulted
in a 50% reduction in shoot and/or root growth (table 2). Compaction with the
rubber hammer produced values of soil strength similar to those that were measured
in the field, 18-19 bar, on one occasion (New Zealand Soil Bureau, 1968). Data on
dry bulk density and total porosity were not statistically analysed. However, as
expected, dry bulk density increased and total porosity decreased with increasing
rate of compaction (table 2). This is in accordance with Baligar ef a/., (1981) and
Boone ef al., (1978).
Amount of shoot dry matter from compacted pots differed significantly from
those of the control and an inverse relationship occurred between plant growth and
the level of compaction (cf Mitchell et a/., 1981). There was a 10-fold range in the
growth at extreme degrees of soil compaction. However, no difference in shoot dry
matter was found between the intermediate rates of compaction (table 2). Top
growth is shown in Plate 3: A-E. Dry root matter was not statistically analysed but
a downward trend with increasing rate of compaction was apparent. Typical
development of the root system in straight, compacted and uncompacted clay loam
is depicted in Plate 2: a, b, c & f. Plate 2g represents the root system of an
A. pycnantha developed in an uncompacted garden loam supplemented with
inorganic fertilizer. This was kept solely for observation. Plate 2 shows that the root
system became less fibrous and ramified with increasing level of compaction, and in
the case of machine compaction (plate 2a), root development was highly incompatible
with normal plant growth.
Addition of a medium-size grade of sand on a 20%V basis had no mitigating
effect on the mechanical impedance of clay loam compacted with the steel bar. The
presence of a small amount of coarse material could have actually promoted the
compactibility of the fine clay loam. Consequently, plant growth in treatment 6 did
not differ from that in straight clay loam soil compacted similarly (table 2 — treat-
ment 3). The ameliorative effects of sand were prominent on a 50% V basis (table 2).
—OO
Plate 1.
¢
cm
L
=
i
=
=
=
=
=
=
Plate 2.
Natural and remoulded pan material.
A, left, pots of remoulded and natural pan material (middle) from the B horizon of Paremata
clay; B, right, close-up of remoulded material.
Development of root systems in straight and amended clay loam under different levels of
compaction.
Right to left, a, treatment 4; b, treatment 2; c, treatment 3; d, treatment 6; e, treatment 7; f,
treatment 1 (uncompacted soil); g, root system developed in an uncompacted garden loam.
86 Gard. Bull. Sing. 37(1) (1984)
The penetration resistance was reduced to a level similar to that of treatment 1 and
was significantly lower than that of treatment 3 (table 2). On adding 50%V sand,
the dry bulk density decreased while the total porosity increased as compared with
those in treatment 3 (table 2). As a result, plant growth in treatment 7 was of the
same order as in treatment | (table 2; plates 2, e&f, 3A & 4B). All seedlings established
in pure sand died after 4 months (plate 4C). It was thought that the particular grain
size of sand used might have given rise to a transient high water table after irrigation,
thereby damaging the root system. The cause of total seedling mortality in clay loam
amended with digested sludge (plate 4D) was tentatively attributed to a partial/total
reduced state in the rooting zone generated by the initial decomposition of the
sludge.
The value obtained from the oxygen microelectrode inserted to depths of 3 to 4cm
below the surface were all greater than 2.5 x 10°’ g cm’ min’. A value of
Water content °%> (w/w)
35 4O 45 50
10
Tension cm (HO)
U1 NRO
Oo On
~
on
100
b
Fig. | Water contents of a clay loam compacted by four methods and equilibrated at five moisture
tensions.
Soil compacted by: a, light finger pressure; b, rubber hammer; c, steel bar; d, mechanical press.
Note. Each point on the graphs is an average of 2 measurements.
Growth of Acacia pycnantha in compacted soil 87
1 x 10~’ gcm~ min“ is generally considered adequate for plant growth (Gradwell,
1972). The purely inorganic soil matrix supplemented with inorganic fertilizer had
the advantage of not inducing the type of reducing conditions as might be encountered
when digested sludge was incorporated in clay soils.
Figure 1 presents pF plots of the average water contents of cores from the
compacted clay loam soil after equilibration at five values of moisture tension.
Under the weak compaction exerted by finger pressure or rubber hammer, the soil
had about 10% more water available at tensions of 10 and 25 cm than in soils
compacted by steel bar or by machine compression. At high moisture tensions,
above 75 cm, the water contents of the cores were similar for all methods of
compaction. Mixing sand with the clay loam had marked influence on the water
content of the cores after equilibration at different moisture tensions. Figure 2
shows pF curves of water content against moisture tension for mixtures of clay loam
and sand, and clay loam and sludge, when all were compacted with the steel bar.
The water contents were all very similar at low moisture tensions. At high moisture
tensions, the water contents varied inversely according to the amount of sand that
had been added. In other words, addition of sand made water more easily available.
Water content °%o (w/w)
oe 30 35 40 45
10 - =
25 - =
Tension cm (HO)
g
25
100' ° A °
d € b
25+
| |
re x
@ a
Fig. 2 Water contents of a clay loam ameliorated by sand or sludge and compacted with a steel bar.
Composition: a, clay loam: b, 80% clay loam + 20% medium sand; c, 50% clay loam + 50%
medium sand; d, medium sand; e, 80% clay loam + 20% sludge.
Note. Each point on the graphs is an average of 2 measurements.
Plate 3. Development of plant top on clay loam under different levels of compaction.
A, above left, treatment 1; B, right, treatment 2; C, middle left, treatment 3; D, right, treatment
4; E, below, comparison of 4 rates of compaction on the development of plant top; /eft to right,
treatment | (control), treatments 2, 3 and 4.
88
Plate 4. Development of plant top on clay loam amended with 2 rates of sand, pure sand and clay
amended with sludge, all compacted with steel bar.
A, above left, treatment 6; B, right, treatment 7; C, middle left, treatment 5; D, right, treatment
8; E, below, comparison of the ameliorative effects of 2 rates of sand on the development of
plant top; /eft to right, treatment 1 (control), treatments 6, 7, and 5.
90 Gard. Bull. Sing. 37(1) (1984)
Conclusions
A. pycnantha seedlings were unable to establish roots on a scarified surface of a
pan material from the subsoil of Paremata clay or on the remoulded pan material
containing adequate mineral fertilizer.
It was evident that upon compaction, soil strength became the severely limiting
physical attribute that controlled the establishment and early growth of A. pycnan-
tha in the Paremata clay-loam subsoil. A direct relationship was found between this
property and the degree of compaction. Mechanical impedance, derived from machine
compaction, represented a limit to penetration by roots. Hence, the growth
of A. pycnantha, a very hardy plant, could be inimically affected by high degrees
of soil compaction.
Alleviation of the dramatic changes in soil strength due to compaction was
accomplished by adding a medium-size grade of sand on a 50%V basis prior to
compaction; this resulted in normal plant growth.
Acknowledgements
The work reported was carried out at the New Zealand Soil Bureau under a
New Zealand-Singapore Colombo Plan Co-sponsorship for the first author. He is
grateful to the Soil Bureau staff for their help and discussions during the course of
the practical attachment, and Mrs Foong Thai Wu for her assistance with the trial.
Literature Cited
Audus, J. W. (1934). Native trees of Australia, 84. Whitcomb & Tombs Limited,
Australia.
Baligar, V. C., V. E. Nash, F. D. Whisler & D. L. Myhre (1981). Sorghum and soy-
bean growth as influenced by synthethic pans. Communications. Jn Soil Sci.
Plant Anal. 12: 97-107.
Barley, K. P. & E. L. Greacen (1967). Mechanical resistance as a soil factor influenc-
ing the growth of roots and underground shoots. Advan. Agron. 19: 1-43.
Boone, F. R., J. Bouma, & L. A. H. de Smet (1978). A case study on the effect of
soil compaction on potato growth in a loamy sand soil. 1. Physical measurements
and rooting patterns. Neth. J. agric. Sci. 26: 405-420.
Cannell, R. Q. (1977). Soil aeration and compaction in relation to root growth and
soil management. Jn T. H. Coaker (ed.), Applied Biology Vol. II, 1-86. Academic
Press, London.
.(1982). Cereal root systems: Factors affecting their growth and function.
Monograph, British Plant Growth Regulator Group, No. 7, 118-129.
Drew. M. C. & M. J. Goss (1973). Effect of soil physical factors on root growth.
Chemistry & Industry 21: 679-684.
Growth of Acacia pycnantha in compacted soil 91
. (1974). Environmental stress and the growth of barley root systems: the
effect of nutrient ion concentration and mechanical impedance. Jn R. L. Bieleski,
A. R. Ferguson & M. M. Cresswell (eds.), Mechanisms of regulation of plant
growth, 111-119. The Royal Society of New Zealand, Wellington.
Eavis, B. W. (1972a). Soil physical conditions affecting seedling root growth I.
Mechanical impedance, aeration and moisture availability as influenced by bulk
density and moisture levels in a sandy loam soil. P/. Soil 36: 613-622.
.(1972b). Soil physical conditions affecting seedling root growth. I]. Com-
parisons between root growth in poorly aerated soil and at different oxygen par-
tial pressure. P/. Soil 37: 151-158.
Gradwell, M. W. (1972). Methods for physical analysis of soils. New Zealand Soil
Bureau Scientific Report 10c.
Greacen, E. L. & R. Sands (1980). Compaction of forest soils. Aust. J. Soil Res.
18: 163-189.
Heilman, P. (1981). Root penetration of Douglas-fir seedlings into compacted soil.
Forest Sci. 27: 660-666.
Lal, R. & D. J. Greenland (eds.) (1970). Soil physical properties and crop produc-
tion in the tropics. John Wiley & Sons, New York.
Mitchell, M. L., A. E. Hassan, C. B, Davey & J. D. Gregory (1981). Effect of soil
compaction on root development and seedling establishment. American Society
of Agricultural Engineers, Paper No. 81-1040.
New Zealand Soil Bureau (1968). Soils of New Zealand. Part 3. New Zealand Soil
Bureau Bulletin 26: 127.
Raghavan, G. S. V., F. Taylor, B. Vigier, L. Gauthier, & E. McKyes (1982). Effect
of compaction and root rot disease on development and yield of peas. Can.
Agric. Eng. 24: 31-34.
Ruark, G. A., D. L. Mader & T. A. Tattar (1982). The influence of soil compaction
and aeration on the root growth and vigour of trees — A literature review. Part
I. Arboriculture Journal 6: 251-265.
Russell, R. S. & M. J. Goss (1974). Physical aspects of soil fertility — The response
of roots to mechanical impedance. Neth. J. agric. Sci. 22: 305-318.
Sands, R. & G. D. Bowen (1978). Compaction of sandy soils in Radiata Pine forests.
Il. Effects of compaction on root configuration and growth of Radiata Pine
seedlings. Aust. For. Res. 8: 163-170.
Vigier, B., G. S. V. Raghavan & G. Drake (1983). Effect of soil compaction and
sterilization on root rot disease and yield of canning peas. Can. J. Plant Sci. 63:
327-331.
Zisa, R. P., H. G. Halverson & B. B. Stout (1980). Establishment and early growth
of conifers on compact soils in urban areas. Forest Service Research Paper
NE-451. USDA Forest Service Northeastern Forest Experiment Station.
The Embryology of Garcinia mangostana L. (Clusiaceae)
LIM AH LAN
Department of Botany, University of Malaya, Kuala Lumpur, Malaysia
Abstract
The anther is tetrasporangiate. The anther wall is four-layered and the secretory tapetum is binucleate.
Most of the pollen mother cells degenerate just before meiosis and a few undergo meiosis. Cytokinesis
is simultaneous and tetrads are tetrahedral.
The ovule is anatropous, bitegmic, tenuinucellate and the micropyle is formed by the outer integument
only. The embryo sac development is of the Polygonum type.
The endosperm is of the Nuclear type. Cell-wall formation proceeds from the periphery towards the
centre. The development of the adventive integumentary embryos is irregular and sometimes within the
same embryo sac, several embryos may be found. The mature embryo is straight. The seed coat develops
from the outer integument. The seed is non-endospermous and shows hypogeal germination and 10%
of the germinated seeds give multiple seedlings.
Introduction
The mangosteen (Garcinia mangostana L.) commonly known in Malaysia as the
*‘Queen of fruits’’ is indigenous to the Malay Archipelago (Burkill, 1935) but the
present distribution stretches from southern India through the Malesian region to
as far as the Philippines. The plant is dioecious, of medium height (5-15 m) and
produces a characteristic opaque, yellow latex. It is slow-growing and requires
approximately 15 years to bear fruits. In Peninsular Malaya, the mangosteen
produces fruits twice a year; between June and August, and October and December.
This seasonality can easily be upset by any change in the pattern of the wet and dry
seasons. Most individual trees, however produce fruits only once a year, some
during the first season and the others during the second. The fruits are usually eaten
fresh or they can be made into a pleasant preserve called ‘‘halwa manggis’’.
As early as 1919, Sprecher described the fruits, seeds and the embryogeny of
G. mangostana. Treub (1911), Puri (1939), Horn (1940) and much later Ha (1978)
contributed some information on the flower, fruit and seed development in Garcinia.
Nevertheless, our knowledge of the embryology on this genus remains fragmentary.
Materials and Methods
Buds, flowers and fruits of G. mangostana were collected at regular weekly
intervals from the fruit-tree nursery, Ministry of Agriculture, Serdang, West Malaysia.
A voucher specimen KLU 1/837 was deposited in the Herbarium, Botany Department
of the University of Malaya. The buds and flowers were fixed in formalin-propionic-
alcohol (5 c.c. formalin, 5 c.c propionic acid and 90 c.c 50% ethanol). For fruits,
the fixing solution used was Craf III (30 c.c. 1% chromic acid, 20 c.c. 10% acetic
acid, 10 c.c. 37% formaldehyde and 40 c.c. distilled water) at O°C, as 50% F.A.A.
93
94 Gard. Bull. Sing. 37(1) (1984)
hardened them excessively. Customary methods of dehydration and embedding
were followed. Much difficulty was encountered in sectioning the fruit tissues
because of the bulk and presence of sclerenchyma tissue. To overcome this, the wax
blocks, with the tissues exposed, were immersed for a month in a softening solution
of Molifex (B.D.H., U.K. Co.) or Bakers solution (36 c.c distilled water, 54 c.c.
95% ethanol, 10 c.c. glycerol). Microtome sections were cut at 12 and stained in
1% alcoholic (in 50% alcohol) safranin and 0.5% alcoholic (in 95% alcohol) fast
green FCF.
Observations and Results
FLORAL AND FRUIT MORPHOLOGY
The flowers about 4 cm wide, borne singly or in pairs, are terminal in position.
The four curved sepals are persistent on the fruit and the four fleshy petals are
tinged red (fig. 1!A-C). The superior ovary is round, capped by the sessile 5—8-lobed
stigma. The base of the ovary is surrounded by 14-16 staminodes. Anthesis is in the
evening between 4 and 6 pm and after 24 hours the petals drop off whereas the
sepals persist on the ripe fruit.
The fruits are approximately 8 cm in diameter and when ripe, have a purplish rind
0.5 cm thick. Within the rind, there are 5-8 fleshy or pulpy segments which, in some,
contain a light, brown seed that adheres to the flesh. The texture of the pulp is soft,
delicately flavoured and slightly acidic in taste.
The mature seeds are non-endospermous, flattened and of various sizes. They are
usually 1.0-1.5 cm long and 0.3-0.5 cm thick.
MICROSPORANGIUM, MICROSPOROGENESIS AND MICROGAMETOPHYTE
The anther is tetrasporangiate with four distinct wall layers within the epidermis
viz., the endothecium, two middle layers and the tapetum (fig. 2A-D). The secretory
tapetum is initially uninucleate but later becomes binucleate (fig. 2C). It degenerates
soon after the formation of microspore tetrads.
In young anthers, the microspore mother cells are well formed and prominent
(fig. 2B, 2C). However, as they enter into the meiotic division, their nuclei
disintegrate first and then their cytoplasm, causing most of them to degenerate
(fig. 2K). A few do undergo reduction division and simultaneous cytokinesis
(fig. 2F-J) but the chromosomes stain very faintly and show signs of disintegration.
At various stages of meiosis, the microspore mother cells continue to degenerate in
large numbers. A few microspore tetrads and single microspores appear normal but
the latter were non-viable as shown by their failure to stain in 1% 2, 3, 5-triphenyl
tetrazolium chloride.
Simultaneous with the degeneration of the tapetal cells, the epidermis accumulates
tanniniferous materials while fibrous thickenings are formed in the endothecium
and the middle layers (fig. 2E) and all these persist in the mature flower.
Fig. 1. Flower, fruit and seedling morphology. A, flower bud; B, flower; C, fruit; D-/, seed germination;
J, polyembryonic seedling; K, ‘‘three-leafed’’ seedling; L, twin seedlings partially fused along the
stem.
95
96 Gard. Bull. Sing. 37(1) (1984)
MEGASPORANGIUM, MEGASPOROGENESIS AND MEGAGAMETOPHYTE
The ovular primordium develops from the placenta. The differentiation of the
inner integument is followed by that of the outer (fig. 3A-C). Simultaneously, one
of the hypodermal, nucellar cells enlarges to form the archesporial cell (fig. 3A, 3B).
The ovule is anatropous, bitegmic, tenuinucellate and the micropyle is formed by
the outer integument only (fig. 3K). A short, thick funiculus is present. Only one
Ovule is present within each loculus.
The archesporial cell enlarges to function directly as the megaspore mother cell
(fig. 3A, 3B). At this stage, the outer integument gradually outgrows the inner,
enclosing the nucellus to form the micropyle (fig. 3C). Usually only one archesporial
cell is produced but sometimes two or three are formed. Following the first meiotic
division in the megaspore mother cell, a pair of unequal dyad cells is developed and
the pair divides further to form a linear tetrad. At the completion of meiosis, only
the chalazal megaspore enlarges further to form the female gametopyte while the
other three megaspores degenerate (fig. 3F). The nucleus of the functional megaspore
undergoes three successive divisions to form an 8-nucleate embryo sac which is of
the Polygonum type (fig. 3H-J). The antipodals usually degenerate soon after
formation but rarely do they persist even up to the stage when the egg apparatus
degenerates (fig. 4A). Approximately 50% of the ovules examined showed degenera-
tion of its embryonic content (fig. 3K). This was observed during various stages of
the embryo-sac formation. Often, the multiple archesporial cells differentiate into
the megaspore mother cells and further develop into multiple embryo sacs. The
development of multiple embryo sacs is not synchronous and frequently, all but one
degenerate (fig. 3D-G).
POLLINATION AND FERTILISATION
In the female flower, viable pollen grains are never found and those formed are
empty and non-viable. Viable pollen grains are probably produced in the male
flower but there is no report of a male G. mangostana tree in the Malay Peninsula.
Parthenocarpy and apomixis in the species had been suggested by Corner (1952).
In order to confirm this inference, bagging experiments were done. A set of 100
flower buds (diameter 1.5 cm) at similar stages of development was selected and
tagged. While 50 were bagged with the anthers removed, the rest were bagged with
the anthers intact. Fruit set in the samples bagged with anthers removed versus
anthers intact were 20% and 53% respectively. This suggests that cross pollination
and fertilisation are not necessary for fruit formation. The removal of anthers from
the buds before anthesis also excludes the possibility of self pollination. The percentage
of fruit set in flowers with intact stamens is much higher than that with the anthers
removed, possibly because of the unavoidable injury while emasculating.
ENDOSPERM
Before the degeneration of the synergids, the two polar nuclei fuse to form a
secondary nucleus. The division of this is not followed by wall formation and hence
Fig. 2. Microsporangium and microsporogenesis. (dp, degenerated microspore mother cell; ed, en-
dothecium; ep, epidermis; mic, microspore mother cell; m/, middle layer; osp, outer secondary
parietal layer; fa, tapetum). A, tetrasporangiate anther; B-E, stages of anther wall development;
F-J, meiosis in microspore mother cell; K, degenerated microspore mother cell.
97
98 Gard. Bull. Sing. 37(1) (1984)
endosperm formation is of the ab initio Nuclear type. The embryo sac enlarges at
the expense of the nucellus and the inner integumeni. Free nuclei increase in number
and remain scattered within the enlarging embryo sac (fig. 4B-D). Proceeding from
the periphery to the centre, they become cellular when the embryo is in its advanced
stage of development. This endosperm is progressively consumed by the developing
embryo and in a fully mature seed, only one or two layers of it remain.
EMBRYO
The degeneration of the egg apparatus follows that of the antipodals and the
fusion of the polar nuclei. However, one instance of an egg (probably developing
into an embryo) surrounded by endosperm nuclei was observed (fig. 4F). An
exceptional case of a persistent antipodal, was also observed simulating a developing
egg (fig. 4A). As the embryo sac enlarges, the nucellus disintegrates, leaving only
one or two layers of cells in the chalazal region. The degeneration of the nucellar
tissue is followed by that of the inner integument, leaving only one or two layers
of cells at the chalazal region of the embryo sac. When the embryo sac is about
0.4 cm long, the cells of the inner integument will have completely degenerated and
the innermost layer of the outer integument becomes meristematic and divides rapid-
ly. Undulations appear along the inner wall of the embryo sac. Cells of the outer
integument divide periclinally and anticlinally, producing buds which project into
the embryo sac. The divisions in these buds are irregular and buds of different
shapes result vis., button-like, linear etc. (fig. 4G-L). In the early stages of develop-
ment, numerous integumentary buds are formed, but as development proceeds only
a few of these attain maturity. Both the linear and button-shaped ‘proembryos’
eventually form broad-based, irregularly shaped embryos (fig. 4P). The developing
embryos apparently obtain nutrients mainly from the integument and the nuclear
endosperm.
The two cotyledons of mature embryos differ in size and shape. The epidermal
cells of the cotyledons are small and their nuclei prominent while all the other layers
have large and vacuolated cells with their nuclei peripheral. Polyembryony occurs
when more than one bud matures within an embryo sac. In 50% of the ovules
examined, the content of the embryo sac degenerates but the development of seed
coat and ovary wall proceeds normally (fig. 3K). This produces seedless fleshy
segments.
SEED COAT
In a mature ovule, the inner and outer integuments are respectively two layers and
five to six layers thick (fig. 4M). At the time wheu the outer integument becomes
meristematic, the outer epidermis of the outer integument accumulates tannin
(fig. 4N). As development proceeds, more tannin is deposited in the cells of the outer
integument. At the mature embryo stage, the enlarged embryo-sac cavity is
lined by one or two layers of parenchyma cells while the rest of the cells of the outer
integument are interspersed with tanniniferous cells (fig. 40).
Fig. 3. Megasporangium, megasporogenesis and megagametophyte (ac, archesporium; dm, degenerating
megaspore; eg, egg; i/, inner integument, m, megaspore; mmc megaspore mother cell; o/, outer
integument; sn, secondary endosperm nucleus; sy, synergid). A-C, development of integuments;
D-G, development of twin embryo sacs within an ovule; H, a developing embryo sac; J, egg
apparatus with secondary endosperm nucleus; J, f.s. embryo sac showing the egg, one synergid
and secondary endosperm nucleus; K, degeneration of embryonic content within an ovule.
99
100 Gard. Bull. Sing. 37(1) (1984)
FRUITS AND SEEDS
When the fruit fully mature, the diameter varies from 5 to 7 cm. The number of
segments (5-8) within it corresponds to the number of stigmatic lobes of the flower.
The fleshy segment may enclose a seed or it may be seedless. A random count of
fifty fruits showed that the ratio of seeded to seedless segments was 1:2.
The seeds are of various sizes ranging from 0.5 to 2.0 cm in width. They are
brown, flattened, have no endosperm and traversing its surface is a network of
vascular bundles.
SEED GERMINATION
Seed germination is hypogeal and the incubation period is one week. The viability
of fresh seed is very high, showing an average of 91% germination. During germina-
tion, the testa splits, one small root emerges from one end of the seed a few days
before the appearance of the plumule at the opposite end (fig. 1D-G). The erect
shoot system grows to about 4.0 cm before unfolding its first pair of leaves (fig. 1H-I).
Normally, only one seedling germinates from each seed but in approximately 10%
of the seeds germinated, polyembryony occurs and commonly, two or three, as
many as even four seedlings per seed have been observed. All these seedlings arise
from different positions of the seed and possess their own separate root systems
(fig. 1J). Occasionally, the two to three shoots may arise from the same point of
the seed and then they share one root system. This occurs as a result of the develop-
ment of lateral buds in the shoot as can be seen from the longitudinal section of the
multiple shoots which shows that the vascular bundles of the shoots are branches
of the main vascular bundle of the hypocotyl (fig. 4Q).
Fasciation is another common phenomenon among the seedlings. Often two
seedlings are partially fused along the stem or the two shoot systems are completely
fused except at the apices (fig. 1L). Sometimes, a seedling may have three leaves
at a node owing to incomplete leaf separation (fig. 1K).
Discussion
In G. mangostana, so far, in Malaya only trees with female flowers are known
in cultivation. According to Corner (1952), ‘“‘the male trees have never been found
in Malaya though they are said to occur rarely in Indochina’’. In addition, fertile
anthers are not found in the female flowers. Chevalier (Bull. Agric. Instit. Sci.
Saigon, I, 1919 pg. 101) in Burkill (1935) stated that the female plant produced
flowers which had male organs and that one could suppress all the male plants in
a garden without affecting fruit set.
Within the family Clusiaceae, the development of pollen grains has not been in-
vestigated (Davis, 1966). This present study shows that the anther of G. mangostana
is tetrasporangiate and degeneration in female flowers has been observed at various
stages of microsporogenesis. In addition, the few microspores produced were tested
and found non-viable. This observation strengthens the earlier report of Corner
Fig. 4.
Embryogeny and development of the seed coat. (ap, antipodal; ce, cellular endosperm; em,
embryo; en, endosperm nucleus; /b, integumentary buds; i/, inner integument; o/, outer integu-
ment; vs, vascular system). A, persistent antipodal developing into embryo; B-E, l.s. ovule
showing seed coat development; F, egg with endosperm nuclei, G-L, development of integumen-
tary embryo; M-O, portions of seed coat enlarged from C-E; P, an irregularly shaped embryo;
Q, germinated seed with multiple shoots.
101
102 Gard. Bull. Sing. 37(1) (1984)
(1952) and suggests apomixis as the mode of reproduction. Within the genus Garcinia
pollen-grain degeneration has also been reported in the staminodes of the female
flowers of G. livingstonii (Puri, 1939) whereas in G. forbesii, the microspore mother
cells do undergo simultaneous cytokinesis to produce binucleate, mature pollen
grains (Ha, 1978).
Preliminary bagging experiments confirm that the seeds are formed without
fertilization. However, the fruits formed from bagged flowers with the anthers
removed are slightly smaller (diameter 4.0 cm) than those with the stamens intact
(diameter 4.8 cm). This is probably due to physical damage during the process of
removing the anthers. Similarly, Maguire (1976) found that some Clusia species
occurring in the semi-deciduous forests in Guayana also produced fruits from unfer-
tilized flowers . But he did not establish the type of asexual reproduction.
The present study reaffirms the early observation of adventive embryony in
G. mangostana by Sprecher (1919). Horn (1940) further explained that since all the
seedlings were derived from essentially the same type of cells i.e., the integument,
they were of exactly the same genotype. That explains that hence throughout
Malaya, the fruits of G. mangostana taste alike and there is only one variety present.
A race with a thicker rind and more acid flesh is said to occur in the Sulu Islands
(Burkill, 1935).
Endosperm formation in the Clusiaceae is Nuclear in type and embryogeny
conforms to the Onagrad type (Davis, 1966). In G. mangostana, the actual type of
embryogeny could not be determined because of the irregular sequence of cell
division in the integumentary buds. The various species of Garcinia in particular and
the members within the Clusiaceae in general should be investigated extensively in
order to establish the embryological pattern of development in the family.
Acknowledgements
I express my sincere thanks to the Vice-Chancellor, University of Malaya for
research facilities and grant which made this study possible. I am very grateful to
Dr. N. Prakash, University of New England, Australia for advice and comments.
References
Allen, B. M. (1967). Malayan Fruits. Donald Moore Press Ltd. Singapore.
Burkill, I. H. (1935). A Dictionary of the Economic Products of the Malay Penin-
sula. Vol. Il. Crown Agents, London.
Corner, E. J. H. (1952). Wayside Trees of Malaya. Vol. 1. Government Printing
Office, Singapore.
Davis, G. L. (1966). Systematic Embryology of the Angiosperms. John Wiley and
Sons Inc. New York. London.
Fairchild, D. (1915). The Mangosteen. Jour. Hered. 6: 339-347.
Embryology of Garcinia mangostana 103
Horn, C. L. (1940). Existence of only one variety of cultivated mangosteen explained
by asexually formed ‘‘seed’’. Science 92: 237-238.
Ha, C. O. (1978). Embryological and cytological aspects of the reproductive biology
of some understorey rainforest trees. (Ph.D. thesis, University of Malaya,
unpublished).
Maguire, B. (1976). Apomixis in the genus C/usia (Clusiaceae) - A preliminary
report. Taxon 25: 241-244.
Puri, V. (1939). Studies in the order Parietales, 2. A contribution to the morphology
of Garcinia livingstonii. Proc. Indian Acad. Sci. B 9: 74-86.
Sprecher, M. A. (1919). Etude, sur la semence et la Germination du Garcinia
mangostana L. Rev. Gen. Bot. 31: 513-531.
Treub, M. (1911). Le sac embryonnaire et l’embryon dans les Angiospermes. Nouv.
Série des recherches. Ann. Jard. Bot. Buitenzorg 24: 1-17.
Vestal, P. A. (1937). The significance of comparative anatomy in establishing the
relationship of the Hypericaceae to the Guttiferae and their allies. Philipp. Journ.
Sci. 64: 199-256.
Multiple Branching at Nodes of
Symingtonia populnea (Hamamelidaceae)
FRANCIS E. PUTZ and N. MICHELE HOLBROOK
Department of Botany, University of Florida, Gainesville, Florida, U.S.A
Abstract
Vertically oriented sunlit shoots on Symingtonia populnea trees in the mountains of Malaysia produce
multiple branches at nodes. The multiple axillary branches originate as branches of the vascular trace to
the primary lateral bud. Plasticity in the number of branches produced per leaf axil allows S. populnea
considerable architectural flexibility in response to environmental heterogeneity.
Introduction
Symingtonia populnea (R. Br. ex Griff.) Steen. (Syn. Bucklandia populnea
R. Br. ex Griff. and Exbucklandia populnea R. W. Brown) is an early successional
tree species of upper montane ericaceous forests (100-300 m) in tropical SE. Asia.
On the mountains Gunong Ulu Kali and Gunong Brinchang in Peninsular Malaysia,
it is prominent along roadsides but within the forest it is restricted to large treefall
gaps and landslides. We were attracted to S. populnea by a branching pattern that
is rare among dicotyledons and possibly unique among dicotyledonous trees.
Symingtonia populnea trees continuously produce leaves (Ng, 1979) and sylleptic
branches, that is, branches that develop simultaneously with subtending leaves and
main shoots (Tomlinson and Gill, 1973). All axes are orthotropic with distichous
leaf arrangement and lateral inflorescences. Although there is no initial morphological
differentiation among branch axes, some grow more slowly than others and these
become displaced further from a vertical orientation than leader shoots. Using the
system proposed by Hallé and Oldeman (1970) for classifying plant architectural
types, this combination of characters conforms with Attim’s model. However,
S. populnea trees diverge from this familiar growth pattern when a second and
sometimes a third branch emerges from a leaf axil, especially on vigorously growing
leader shoots (plate 1).
The occurrence of multiple branching at nodes appears to depend on the particular
shoot’s vigour and position in the crown as well as on the position of the tree relative
to its neighbours. Shaded trees and shaded branches on well illuminated trees rarely
produce more than one branch per node. Symingtonia populnea trees planted in full
sun in the lowlands, well below their natural altitudinal range, manage to survive
but are unhealthy; the frequency of multiple branching at nodes on these trees is
extremely low.
To establish the frequency with which nodes on well illuminated leader shoots and
nodes on shaded subordinate branches give rise to more than one lateral branch, we
counted branches at nodes on five small (3-6 m tall) trees growing in full sun on
Gunong Ulu Kali. To allow branches time to develop and to avoid the potential
105
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id > SX f } es
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Plate 1. Pencil drawing of a S. populnea shoot showing development of multiple axillary branches along
the main axis. (About 1/5 life-size).
problem of branch loss, neither the youngest nodes nor nodes on branches more
than 3 cm diameter were included in the census. Branch departure from vertical
orientation was used as a measure of vigour; overtopped branches became more
horizontal with age.
On vertically oriented shoots (less than 10° from vertical), 177 of the 335 nodes
examined had two branches, while on branches diverging from vertical by more than
10° only 51 of the 872 nodes examined had two branches. This difference is highly
significant (G = 441, P < 0.01).
Buds on S. populnea trees are enveloped by pairs of large, tightly appressed,
obovate, to oblong stipules (plate 2). When the stipules unfold, the enclosed main
axis and at least one lateral branch are 1-3 cm long. The unique arrangement of
twice-folded petioles (Lubbock, 1899) allows for this extended development within
the stipules. In most cases in which a second branch develops, it does so slowly
during which time one or two new leaves and branches are produced by the main
axis. Sometimes the delay before emergence of the second branch is more prolonged
but we have never observed newly expanded branches from axes more than 3 cm
in diameter. The first branch produced at each node was smaller in diameter, had
shorter internodes, and usually had smaller leaves than the main axis. Second
branches at nodes are smaller than first branches. Where there was no branch loss,
these size differences were further accentuated by differential growth.
106
Plate 2. S. populnea shoot apex with one stipule removed. (Approximately life-size).
Developmental studies of S. popu/nea shoot apices revealed that multiple axillary
buds originate as branches of the vascular trace to the primary lateral bud. This
branching results in a proliferation of successively higher order axillary lateral
meristems (a secondary bud complex; Hallé et a/., 1978) forming what might be
called an embryonic short shoot. In S. popul/nea, each successive axillary branch bud
is oriented at approximately 90° to the previous bud, forming a spiral arrangement
(fig. 1). Both left-handed and right-handed spirals were observed. Up to four orders
of branching were observed within a single apex, and although such ‘‘precocious’’
branching was not limited to apices taken from vertically oriented shoots, it was
more vigorously expressed there. Prophyllary buds are known to occur in a variety
of species, for example, in many bamboos (McClure, 1976), Leptocarpus simplex
(Tomlinson, 1973), Liquidambar styraciflua (Kormanik and Brown, 1967), and
Gossypium spp. (Mauney and Ball, 1959; Cook, 1911). S. populnea shoot apices
failed to reveal any prophylls, stipules, or other organs subtending the higher order
branch buds. Vascular connections, however, show that each successive branch bud
(both in terms of initiation and maturation) is an offshoot of the preceding bud
(fig. 2).
Plasticity in branching pattern allows S. populnea trees to display many leaves
where conditions are favourable by responding architecturally to within-crown and
within-habitat environmental heterogeneity. A similar response was observed in
Acer saccharum in which leader shoots produce a significantly greater number of
107
as / © © )
stutes .
g AN
subtending leaf
Fig. 1. Diagrammatic representation of a S. populnea leader shoot apex showing the terminal (main)
and four axillary buds.
Fig. 2. Drawings of successive transverse sections of a S. populnea shoot apex (basipetal sequence,
A=uppermost; solid area represents vascular tissue).
108
Multiple branching in Symingtonia populnea 109
first order branches per second order branch than lower limbs (Steingraeber, 1982).
However, S. populnea is distinctive in altering both the frequency and the pattern
of branching.
Acknowledgements
We wish to thank F. S. P Ng, P. F. Stevens, B. C. Stone, P. B. Tomlinson, and
J. R. Vincent for their assistance and advice.
Literature Cited
Cook, O. F. (1911). Dimorphic branches in tropical crop plants. U. S. Dept. Agr.,
Bur. Plant Ind. Bull. No. 198.
Hallé, F., R. A. A. Oldeman, and P. B. Tomlinson (1978). Tropical trees and
forests, an architectural analysis. Springer-Verlag, Berlin.
Kormanik, P. P. and C. L. Brown (1967). Root buds and the development of root
suckers in the sweetgum. Forest Science 13: 338-345.
Lubbock, J. (1899). On buds and stipules. Kegan Paul, Trench, Trubner and
Company, London.
Mauney, J. R. and E. Ball (1959). The axillary buds of Gossypium. Bull. Torrey
Bot. Club 86: 236-244.
McClure, F. A. (1976). The bamboos, a fresh perpective. Harvard Univ. Press,
Cambridge.
Ng, F. S. P. (1979). Growth rhythms in tropical juvenile trees. Bull. Soc. Bot. Fr.
126, Actual. Bot. 3: 139-149.
Steingraeber, D. A. (1982). Phenotypic plasticity of branching pattern in sugar
maple (Acer saccharum). Amer. J. Bot. 69: 638-640.
Tomlinson, P. B. (1973). The monocotyledons; their evolution and comparative
biology. VII. Branching in the monocotyledons. Quart. Rev. Biol. 48: 458-461.
Tomlinson, P. B. and A. M. Gill (1973). Growth habits of tropical trees: some
guiding principles. Jn B. J. Meggers, E. S. Ayensu, and W. D. Duckworth (eds.).
Tropical forest ecosystems in Africa and South America: A comparative review.
Smithsonian Institution Press, Washington, D. C., 129-143.
Trichomanes proliferum Bl.
A new record for the fern flora of Singapore
RoE LJ VIANE
Botanical Institute, Rijksuniversiteit, Gent, Belgium
Summary
Trichomanes proliferum Bl. is reported for Singapore for the first time. Some morphological characters
are discussed.
Introduction
With the kind assistance of Dr. Chang of the Botanic Gardens I was able to collect
some ferns in the Nee Soon swamp forest in July 1982. New records could hardly
be expected as the fern flora of Singapore is rather well known. However, one of
the plants collected was identified as Trichomanes proliferum Bl., which was not
recorded for Singapore by either Holttum (1966) or by Johnson (1977), nor is
there a specimen from Singapore in the herbarium of the Botanic Gardens (SING)
(Dr. Chang, pers. comm.). It seems therefore that the following collection, deposited
in the Herbarium of the State University of Ghent (GENT), is the first from the
Island.
R. Viane & Noé 2369; Singapore, Nee Soon swamp-forest: on tree trunk along
rivulet, just above the water level; rare; together with Trichomanes motleyi v.d.
Bosch; 29 July 1982. Photo (plate 1) in SING.
T. proliferum is a filmy fern (Hymenophyllaceae) of the Gonocormus group that
is characterized by its proliferous frond axes. Copeland (1938: 56) stated about
Gonocormus: ‘‘... a well defined genus of exceedingly ill-defined species ...”’
and distinguished six ‘‘species’’ of which 7. proliferum was said to be the most
characteristic. Holttum (1966: 95-97) believed Gonocormus to be related to the
Vandenboschia group; he accepted two ‘‘not clearly distinct’’ species: 7. minutum
Bl. and T. proliferum, both recorded from Malaya but not from Singapore.
Distribution
T. proliferum was first described from Java by Blume (1828); according to
Copeland (1933: 150) ‘‘it is probably common throughout the Malay region”’ (incl.
Philippines). Holttum (1966) gave ‘‘Malaysia’’ (= area of Flora Malesiana) as the
overall range and stated that the plant is ‘‘found in both lowlands and mountains’.
The single population seen by the present author was growing among mosses on a
very sheltered wet tree trunk in the Nee Soon swamp-forest. More populations of
this small and easily overlooked fern probably exist in the area. Nee Soon is one of
the few remaining lowland swamp-forests that once covered large areas of Singapore
and Johore (Corner, 1978).
111
Plate 1. Trichomanes proliferum Bl. (R. Viane & Noé 2369). F, fertile plant; S, sterile plant. b, pro-
liferous bud with its frond; rp, primary rhizome; rs, secondary rhizome from a bud; s, sorus.
Morphological Notes
A brief description and figure are given by Holttum (1966). Bierhorst (1973, 1974)
studied the branching pattern in Gonocormus in relation to: (a) the concept of non-
appendicular fronds, and (b) the rhizome morphology of other Hymenophyllaceous
“‘genera’’. Yoroi and Iwatsuki (1977) discussed the variability of the frond form and
the proliferation of the Trichomanes minutum-group along with cytological and
anatomical data.
2
Trichomanes proliferum 113
The plants from Nee Soon (plate 1) have fronds up to 4 cm long (including the
stipe), well spaced on a delicate, wiry ‘‘rhizome’’. This ‘‘rhizome’’ has no roots but
is covered by many multicellular uniseriate brown hairs. Few fronds were fertile
(plate 1F). The stipes of (almost) all mature fronds have one to three dormant buds,
visible as slight swellings of the stipe tissue covered with a tuft of brown hairs.
Above these dormant stipe buds there often is a proliferating (= functional) bud
(plate 1: b, arrows) c. 1 mm above the cuneately narrowed lamina base; it forms the
“*secondary”’ frond. This ‘‘secondary’’ frond is similar to the ‘‘primary’’ frond; it
has dormant stipe buds too and the functional bud at its lamina base may produce
another frond, and so on. An additional dormant bud is sometimes present on the
vascular tissue of the lamina (c. 4 mm above its base). The functional buds occa-
sionally produce ‘“‘secondary rhizomes”’ in the abaxial axil of the secondary or
higher order leaves (plate 1F, rs, arrow). The secondary ‘“‘rhizomes’’ are mor-
phologically similar to the primary and develop from the bud only after it has
produced a frond. Another fern with buds forming small but conspicuous (elongate)
rhizomes is the West African ‘‘Ctenitis’’? jenseniae (C. Chr.) Tard.
Nomenclature
If 7. proliferum and T. minutum cannot be recognized as two separate species,
as suggested by Copeland (1958) as well as by Yoroi and Iwatsuki (1977), then
the name 7. minutum BI. sensu lato (Copeland, 1958) should be applied to the
Singapore plants.
Acknowledgements
I thank Dr. Chang for her help in organizing the visit to Nee Soon and for
checking the herbarium material in SING. Prof. K. U. Kramer kindly corrected the
manuscript.
Literature Cited
Bierhorst, D. W. (1973). Non-appendicular fronds in the Filicales. Jn Jermy, Crabbe
& Thomas (eds.), The phylogeny and classification of the ferns. Bot. Journ. Linn.
Soc. 67, Suppl. 1.
(1974). Variable expression of the appendicular status of the megaphyll in
extant ferns with particular reference to the Hymenophyllaceae. Ann. Mo. Bot.
Gard. 61 (2): 408-426
Blume, C. L. (1828) Enumeratio plantarum Javae. Vol. 2. J. W. van Leeuwen,
Leiden.
Copeland, E. B. (1933) Trichomanes. Philipp. Journ. Sci. 51(2): 119-280.
(1938). Genera Hymenophyllacearum. Philipp. Journ. Sci. 67(1): 1-110.
(1958). Fern Flora of the Philippines. Vol. 1. Bureau of Printing, Manila.
114 Gard. Bull. Sing. 37(1) (1984)
Corner, E. J. H. (1978). The freshwater swamp-forest of South Johore and
Singapore. Gardens’ Bull. Suppl. no. 1.
Holttum, R. E. (1966). A revised flora of Malaya. Vol. Il. Ferns of Malaya, 2nd
ed. Government Printing office, Singapore.
Johnson, A. (1977). The ferns of Singapore Island, 2nd ed. Singapore University
Press.
Yoroi, R. and K. Iwatsuki (1977). An observation on the variation of Trichomanes
minutum and allied species. Acta Phytotax. Geobot. 28: 152-159.
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ee ee a OS he
THE GARDENS’ BULLETIN
SINGAPORE
VOL. 37 (Part 2) 1st December 1984
CONTENTS
PAGES
WILDE, W. J. J. O. DE:
A New Account of the Genus Horsfieldia (Myristicaceae), Pt 1 .............2.ceeceeeeeeeeeeeeee 115-179
3 n\
LIM, AH LAN: ()
The Reproductive Biology of Rambutan (Nephelium lappaceum Linn., Sapindaceae) .... 181-192 Ni
WONG, KHOON MENG: \
A Revision of Rennellia (Rubiaceae) in the Malay Peninsula .................0..0c0ecseceeeeeees 193-198 5
I, FARAHD.: \\
ys to the Cultivars of Keladi (Colocasia esculenta — Araceae) in Peninsular @
To rere Es wean back ce eves eGeelee desea sicadsscaeesavecdevendéasendee 199-208 x
KIEW, R.:
a otes on the Systematy of Malayan Species of Chionanthus (Oleaceae)............06.e00e000 209-212
K RTHOF, H. M. & J. F. VELDKAMP: NY
A Revision of Aniselytron with Some New Combinations in Deyeuxia in SE. Asia @
tec Sai oe Soe cain covcassbecocnesccsaoseecasecvecocaassadesduacseates 213-223
(( \\
KIEW, R.: (
Preliminary Pollen Study of the Oleaceae in Malesia ...................cscseessescscessceccseeees 225-230
I gE encore - coc =m desose.ccceccsesvaesesseass neste cevecsenseescesneesem 231-238
Oe OE gE KE DKK.
Published by the Botanic Gardens
Parks and Recreation Department
Ministry of National Development
Cluny Road, Singapore 1025
Printed by Jin Jin Industry Pte. Ltd.
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GARDENS’ BULLETIN
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THE GARDENS’ BULLETIN
SINGAPORE
VOL. 37 (Part 2) lst December 1984
CONTENTS
PAGES
WILDE, W. J. J. O. DE:
A New Account of the Genus Horsfieldia (Myristicaceae), Pt 1 ................cceeeeceeceeceees 115-179
LIM, AH LAN:
The Reproductive Biology of Rambutan (Vephelium lappaceum Linn., Sapindaceae) .... 181-192
WONG, KHOON MENG:
A Revision of Rennellia (Rubiaceae) in the Malay Peninsula .......................c0eeeceee eee 193-198
GHANI, FARAH D.:
Keys to the Cultivars of Keladi (Colocasia esculenta — Araceae) in Peninsular
ee AN AINA REE yc Selec ca D8 alse eid gaatelv os aia. seiia'svielneVSicieaisiowasinasdemancciedeaestasessimetse 199-208
KIEW, R.:
Notes on the Systematy of Malayan Species of Chionanthus (Oleaceae)...............+200000 209-212
KORTHOF, H. M. & J. F. VELDKAMP:
A Revision of Aniselytron with Some New Combinations in Deyeuxia in SE. Asia
SaEe LESAN ISRICAGS Pepe eR eo nn 5 oc scald awa naeindcana is ses se ee/eaeer Hie ou Welowslards Teens 213-223
KIEW, R.:
Broumidacy Pollen Study of the Oleaceae in Malesia .........2..--002..0cssesesesesecssscnsaceoncs 225-230
SN SEMEN) UPR Re nee oe Ng oe g se Orton SS aisienjesaeee va salves see sas laveNemestaneossedae ss 231-238
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A New Account of the Genus Horsfieldia (Myristicaceae), Pt ]
We J O"de WILDE
Rijksherbarium, Leiden, The Netherlands
EFFECTIVE-PUBLICATION DATE: 7 MAR 1985
Contents
Page
Se a ee Pee NRL x ee Bins coin. coe doe see's snc ustsins sees cus Se emae ow cee asia d oot odaceseeneosesesee 115
Sara INCU IETS pene eee tee nee re Ce See esate on ses stabs ceolaene Seedavderchacecenecascss sessnentersceceeassseeses 116
MERE TE PCEATINIELT OLS ITCI OI Cpe sent ttn Ree See ae ore ae hoc ee aaa sean abe es ce 116
ESPMLE Cl APAU CIC UAL ACICLSE 6 cae tame nance aue wae Sols nk sasha 8 oxeae bes ees Sedtewel swede ob ocuesd cheeses des seeuee 117
ee RMSE TICE ASIU CIC MALACIELS omer ne ais ac ass <5 TOSSA Fos ne'sie Soe cisiece Goons Cove dc Gaisgioah ovis oases seas ees oacmeseeneeSs 119
Description and redefinition of the genus Horsfieldia .............cccccccccccceencncneueecececececeeeuceeneneeees 123
SEAM OMIG ISL OYSSICI IG jNtO Three SECLIONS |. f5.).0. 52.522 necednnebe sa scwcleetardicacsecdesssscaceceecausceensteanee 125
aE CR MeATISEAT CARSHEVEV. Ofpa LDC) SLOU PS irs nz: ose aes socom wcelelon scree eis cmeda bocca aeeamnied Socetecemnemeeasmesaaeet 126
aaa ree MAEM OMCLIS EEN LSTIETOM Pree tne eerste eee cece coche cce stokes <ccrercek ose tone ac eee ease eee oO OTE 136
Soe UE UNE SEES ES 6 heohddaebociSka aan ce Bae Annee AP Ee aAE PE nn ne a eo eee ee eee eee ee 138
(@)iGeneral key; based/onimale flowering specimens ..:..2.2.05.200.ccccc.csscescouecsecsesccaceccceeccsssoveess 138
(2-7) Regional keys to female flowering or fruiting speciMeNs ................2c0ceceececeecececeeeeeeeeeeeees 152
BaimMeEAnauralic GeScriptomOf the SPeCleS 1-5) «<2. ......c0acorcsccsoosceceseccenenaceecsesnssdcoscdaaterssecesee 167
Abstract
The genus Horsfieldia, extending from Ceylon to the Solomon Isls. is redefined to exclude the
species formerly assigned to H. macrocoma. The remaining 100 species are subdivided into 3 sections,
viz. (1) the monotypic section Horsfieldia with H. iryaghedhi from Ceylon, (2) section /rya (40 species)
and (3) section Pyrrhosa (59 species). Section /rya contains chiefly species with a 2-valved perianth and is
mainly distributed in East Malesia; the species of section Pyrrhosa have mainly a 3- or 4- valved perianth
and occur in SE. continental Asia and W. Malesia, west of Wallace’s Line. A further subdivision of the
sections into groups of species of closer affinity is proposed and discussed in the introduction. Warburg
(1897) recognized 52 species, Sinclair (1974, 1975) accepted 38 and a number of varieties. The most
important characters for the distinction of species are in the male flowers, especially the androecium. Of
the presently accepted 100 species, 41 are newly described, and in addition, 14 are new subspecies and
varieties. There are 6 new combinations. Most species have rather restricted but well-defined areas of
distribution; H. irya has a very large range, covering almost that of the genus. The centres of species
development are New Guinea and Borneo, and to a lesser extent Malaya-Sumatra.
iS
116 Gard. Bull. Sing 37(2)(1984)
A survey and discussion of all characters regarded as important in Horsfieldia is given. Two kinds of
keys are presented: a general key to all species, mainly based on male flowering specimens, and 6 regional
keys covering the whole area of the genus, concentrating on vegetative characters, for the identification
of female-flowering and fruiting specimens. All 100 species are fully treated and full synonymy, typifica-
tion, description, notes, and a brief enumeration of distribution and examined specimens are provided.
An index of all scientific names pertaining to Horsfieldia will be given in Part 3.
Introduction
With the demise of James Sinclair in 1968*, only the general treatment of
Horsfieldia — the Flora Malesiana precursor of the fourth genus of Malesian
Myristicaceae — remained unpublished, but an alphabetical account of the species,
which were already all worked out by him, was posthumously printed in Gard. Bull.
Sing. 27, 1 (1974) 133-141 and 28, 1 (1975) 1-181. Unfortunately a classification
system and most keys are lacking in these publications and before the treatment of
the family for Flora Malesiana could be started, these had to be provided for.
Moreover, much recently collected material needed first to be identified and
described.
In Sinclair’s earlier revision of the family Myristicaceae for Malaya and
Singapore (Gard. Bull. Sing. 16, 1958) the genus Horsfieldia was thoroughly dealt
with and keys were supplied. However, in order to grasp the whole genus, which ex-
tends over a large area from Ceylon and S. China east to the Solomon Islands and
North Australia, I had to scrutinize all the species myself, and this experience led to
different ideas about the circumscription of the species as a matter of course.
Because Sinclair had examined an enormous amount of specimens from all
major herbaria, I needed only to restrict my study to the Leiden collection com-
plemented by additional, personal visits to BM, K, and P, from where also rather
extensive selected material including type-specimens has been put at my disposal as
loans to Leiden.
Following a somewhat extended list of criteria, important for species delimita-
tion — among which are the highly diagnostic value provided by the presence of
small non-traumatic cork warts on the lower leaf surface, and a more detailed
evaluation of the androecium — I was forced to recognize considerably more species
than did Sinclair. Apart from new ones, most of the species accepted by Sinclair
have been retained in the present study, while others have been redefined, or
renamed, or taken out of synonymy, and some taxa accepted by Sinclair
subspecifically have been raised in rank. An example of the few abrogated taxa is H.
polyantha Warb., also accepted by Sinclair but presently reduced to H. laevigata, a
name which was reduced by Sinclair to H. parviflora.
a
In Sinclair’s enumeration of all species in his 1974/1975 publications, species
descriptions of the wide-spread H. irya as well as that of the variable and deviating
H. macrocoma are lacking. The latter possibly has some indicative value as, during
the present study, it has appeared that some species to be recognized within the H.
macrocoma-complex could be better segregated into a new genus, Endocomia.
Where my circumcriptions of species differ from Sinclair’s, brief reasons are
given in the notes.
The Characters in Horsfieldia
When Sinclair (1958) dealt with Horsfieldia for Malaya, he provided a rather
limited discussion of useful species characters, and in his posthumous publication on
the genus as a whole (1974, 1975), an explanation of the underlying considerations
* Obituary in Gard. Bull. Sing. 23 (1968), pp. i-xxiv.
New account of Horsfieldia 17
for the delimitation of the species is only given in the notes to the alphabetically
arranged descriptions of the species. In those treatments of the Malayan species, two
keys to all species were presented, one for fertile (male flowering) specimens, and
one for sterile material and fruit. From these keys it can be learned that he regarded
the following characters as important for the distinction of the species: (1) perianth
either 2- or 3-valved and (2) shape of the androecium (longer than broad or the
reverse, with or without a depression at the top); apparently of lesser importance to
him were (3) lenticels on twigs, distinct or not, (4) size of the male perianth and
length of the pedicel, (5) some properties of the leaves such as size and texture of the
blade, whether glabrous or pubescent beneath, and above, nerves either raised or
sunk; and, a further number of specific characters used for the distinction of some
individual species.
Although Warburg (1897), Sinclair and I use essentially the same characters for
the generic division, I agree with Sinclair (1958, p. 370) in discarding Warburg’s sub-
division into three sections; Sinclair had advocated a more natural division into two
sections i.e., on the basis of how most of the perianths in a specimen split at an-
thesis, in 2 or 3 valves. I also have the feeling that this criterion is one of the most im-
portant characters for a generic division, one that has phylogenetic significance.
However, in the case of H. iryaghedhi from Ceylon, I have set that apart because it
deviates from all the other Horsfieldias in various ways.
Characters, which in my opinion, are important for distinguishing groups of
species are given further on. They are also used in the list of species with a provi-
sional indication of grouping closer allied species, as presented and discussed in the
next chapter on the subdivision of the genus. Special attention is given to the
vegetative characters because the genus is dioecious and hence in much of the
material the most important taxonomical characters of the male flowers are lacking.
Measurements given in the descriptions of the species and used in the keys based
on vegetative characters and fruits are all taken from dried material; those of
flowers have all been measured after softening by boiling.
Vegetative characters
(1) Habit
The size of the tree sometimes seems specific, but most species are of medium
size. H. sylvestris with a recorded stature of up to 60 m high ranges among the tallest
whereas H. crux-melitensis is a shrub or shrubby treelet of 2-4 m. The growth of the
main stem is essentially orthotropic with the phyllotaxis dispersed, while the
plagiotropic side-branches are generally + horizontal or drooping, with the
phyllotaxis distichous. Characters of bark, buttresses, stilt roots etc., are mainly
observations of Sinclair, as he was acquainted with the species in the field.
(2) Indumentum
As in Knema, the indumentum of the leaf bud, leaves and twigs, and of the in-
florescences and flowers has proved to be important for the delimitation of the
species (de Wilde, 1979, p. 324). Except for the flowers of some species, all the
above-mentioned parts of the plant are initially covered byatomentum, but this
may be shed at a very early stage. Although there may be minor
differences, generally within the same specimen, the nature and size of the
tomentum-hairs on the various parts are largely similar, and to a large degree
characteristic of the taxa. For practical reasons, the tomentum is described only in a
general way, the lengths of the hairs (which themselves may be very complex)
appearing to be of considerable importance for the recognition of the species. Much
on the tomentum can be found in the study on the comparative leaf anatomy of the
Asiatic Myristicaceae by Koster and Baas (1981).
118 Gard. Bull. Sing 37(2)(1984)
(3) Twigs
The indumentum of the twigs is generally shed very early but sometimes may
remain on the young innovations; it is generally similar to that of the leaf bud and
can be simultaneously judged if required for the determination with the keys. The
internodes in the leaf-bearing portion may be terete or faintly lined. Sometimes these
lines, which run from both sides of the insertion of one petiole to the next, are very
pronounced and may render the twig angular or faintly winged, which is
characteristic for several species e.g., H. brachiata. The bark of older twigs i.e.,
behind the leaves and often where the infructescences are inserted, may be striate to
various degrees; only in rare cases does older bark flake more or less characteristical-
ly or crack longitudinally, as in H. sabulosa, H. xanthina or H. disticha. The bark of
young twigs usually dries brown or dark brown, contrasting little with the dark
colour of the dried petioles, but in some species (e.g., H. spicata, H. oligocarpa,
H. pallidicaula), the twigs dry to a conspicuously pale, greyish or straw-colour, con-
trasting well with the petioles, and the discrepancy is then used in the keys. Slightly
older bark nearly always bears lenticels, rather characteristically small or large,
often paler and contrasting well with the twig but at other times merging in colour
and then inconspicuous. In passing, it should be noted that the presence of lenticels
in Malesian taxa of the family seems to be largely or wholly restricted to the genus
Horsfieldia.
In the descriptions, the diameter of twigs was measured not far below the apex
in the leaf-bearing portion while the size cited in parentheses was taken lower down
where old inflorescences or infructescences may be attached. Excepting H. sabulosa,
Horsfieldias never flower on the older wood.
According to Armstrong and Wilson (1980) the wood of Horsfieldia is very
homogeneous and of little taxonomic value at a specific level. At the generic level, it
can be distinguished from the other three Asiatic genera of Myristicaceae. The wood
of one specimen of H. macrocoma (placed in a new genus Endocomia de Wilde,
1984) is apparently not significantly distinct from wood of other Horsfieldias.
(4) Phyllotaxis
In the majority of the species, leaves of the plagiotropic twigs i.e., generally
the fertile ones, are distichous. In some species, phyllotaxis is characteristically
dispersed, mostly with leaves in 3-5 rows (e.g., H. kingii, H. thorelii, H. sucosa, H.
sabulosa); only in a few species are the leaves either distichous or dispersed and
sometimes phyllotaxis is mixed as in one collection. The phyllotaxis is usually very
obvious and easy to see in older twig-portions, hence it is used in the keys. Those
species which have a variable phyllotaxis have been entered in the keys twice.
(5) Leaves
As a matter of course, many features of the leaves, as shape, size, texture, dry-
ing colour, number of lateral nerves and length of the petiole are of importance for
the recognition of the species, but only some characters of special interest will be
briefly discussed here (a-e).
(a) Pale markings on the upper surface
Often present in many species are very small markings, regularly spaced
and usually slightly raised. Their origin is unknown to me. In a few species,
much larger, pale-coloured, often nearly white blotches of irregular shape and
unpredictable location may prevail, sometimes only in a part of the leaves or
only locally present on some of the leaves. They are likely to be an artifact of
drying, of unknown origin and may be found ine.g., H. smithii, yet they are
very characteristic of almost all H. irya collections.
(b) Blackish or brown markings on the lower leaf-surface
A fine, pale, or more often a dark punctation in the form of small blackish
dots, possibly tannic conglomerations, is often present in many species, also in
New account of Horsfieldia 119
the fleshy tissue of perianths, androphores, or pericarps. Furthermore, there is
in certain species a very specific coarser punctation, regularly spaced dots of c.
0.1 mm diam., being a sort of non-traumatic cork warts of unknown origin,
most likely originating from hair-bases or hair scars. The presence or absence
of these dots should always be checked when determining a specimen, their
presence being very distinctive for several species e.g, H. punctatifolia,
H. glabra.
(c) Nerves
In most species, whether midrib and nerves above are flattish, sunken, or
distinctly raised, is of diagnostic significance. For the identification of a few
“‘difficult’’ species as e.g., the distinction between H. whitmorei and H.
laevigata, it is important to check on the submarginal nervei.e., whether its
arches are particularly marked and regularly shaped as is the case with the
former species against the less distinctive and less regular in the latter.
(d) The terminal leaf bud
The terminal leaf bud is usually present in every collection. It shows up as
a long subterete or + flattened cone, usually at least five times longer than
broad; in some species however, mainly in those with the leaves dispersed, it is
shorter and broader. The tomentum of the leaf bud, especially the length of
the hairs, has proved to be a good diagnostic character, as also indicated under
item 2.
(e) Petiole
The petiole is usually well-developed. H. sessilifolia and much materia! of
H. sylvestris have almost sessile leaves; the petioles of H. sabulosa are the
longest in the genus, up to 5 cm long.
Generative characters
(6) Inflorescences
The inflorescences are generally situated on the younger twigs, in-between the
older leaves or not far behind. Only in H. sabulosa does flowering occur on older
wood. They are almost always panicle-like (spike-like in most of H. spicata), (2 or) 3
or 4 times ramified, the mode of branching being essentially racemose. Flowers are
borne on the ultimate branches, either solitary or grouped in loose clusters of up to
10 each in the male inflorescence; those of H. sylvestris, especially in young in-
florescences, may be clustered in rather dense heads, but in the rather deviating
species H. iryaghedhi, the male flowers are grouped in true dense heads each with up
to nearly one hundred flowers.
In general, all flowers of an inflorescence of both sexes are of about the same
age or stage of development, only in H. amygdalina is there a tendency of some to
develop ahead of the others. This latter situation is prevalent in the segregated genus
Endocomia which contains the species EF. macrocoma formerly placed in
Horsfieldia.
Branching is subtended by bracts except the one bearing the ultimate flowers,
the bracts falling off early in all cases. Bracteoles are absent in Horsfieldia, the
segregated genus Endocomia, and Gymnacranthera but present in Knema and
Mpristica.
At the base of the inflorescence there are always a few cataphylls, short and
triangular in outline and clothed with a tomentum similar to that on the in-
florescences, leaf buds and twig-apices.
The largest inflorescences, to c. 30 cm long, are found in H. ampla and H.
ampliformis from New Guinea. In almost all species the male inflorescences are
larger and more elaborate as compared with the female ones; cnly in H. parviflora
are male and female inflorescences often equal in these respects.
120 Gard. Bull. Sing 37(2)(1984)
Following flowering, at fruiting stage, whole inflorescences are discarded as
in Endocomia, Gymnacranthera and in some Myristica, hence unlike the situation in
Knema and the rest of Myristica where flowering occurs on woody brachyoblasts, or
short shoots which are essentially of unlimited growth.
(7) Flowers
The flowers are unisexual, dioecious. The perianth of the male is usually
smaller than that of female flowers (considerably so ine.g., H. thorelii), or they are
equal, or even considerably larger in size. Their size is fairly constant for each
species. For instance, in H. parviflora which was described from a female specimen,
the perianth of the female flower is considerably smaller than the male perianth
which is one of the largest found in Horsfie/dia, up to 4 mm wide.
According to the species, the perianth may be cleft into predominantly 2 or 3
(or 4) valves, and it is on this character that the division of the genus into sections is
based. It is of interest to mention here that this manner of sectional division coin-
cides well with the various distributional areas as is pointed out in the next chapter,
on distribution.
The extent of the split of the perianth at anthesis varies with the species and
cleaving outright or almost to the base (in male flowers) is limited to only a few
species. At anthesis, the perianth opens only a little, the. lobes neither horizontally
spread nor curved nor reflexed as is often the case in Knema, Myristica, Gymnacran-
thera and always in Endocomia.
The flower colour ranges from yellowish green to a bright yellow or orange-
yellow.
The consistency of the perianths may be diagnostic for the species, especially
of male specimens, and sometimes it is useful to note whether the perianth collapses
on drying or not. Flowers may be glabrous, or very early glabrescent; or they are
pubescent, sometimes only so toward the perianth base or the pedicel. The inner
surface of the perianth is always glabrous, in contrast with species of Endocomia,
formerly included in Horsfieldia as H. macrocoma s.1.; it is only in H. subtilis var.
rostrata (formerly H. rostrata) that the inner side of the female perianth was
originally described as pilose, but I have not seen that specimen.
(8) Staminate flowers
Staminate flowers provide for the taxonomically most important characters,
at various levels.
(a) The shape of the male perianth shows much variation. In the 2-valved
species, the perianth is generally laterally compressed, in outline either
circular, or pear-shaped, or reniform, but in H. irya which is the type of the
2-valved section /rya, itis subglobose and not or but little compressed. The
perianth of the 3- or 4-valved species, prevalent in sect. Pyrrhosa, is usually
globose or pear-shaped and not laterally compressed but sometimes apically
depressed, and of various sizes. H. crassifolia has a 2-valved but globose
perianth, and also because of its non-compressed androecium, joins taxono-
mically the 3-valved species in sect. Pyrrhosa.
Peculiarly long-conical 2-valved male flowers prevail in H. crux-
melitensis, H. clavata, and H. squamulosa; they have club-shaped androecia,
aberrant for the genus, as discussed below.
(b) The depth towhich the perianth is cleft at anthesis is a diagnostic
character for several species. Sometimes the perianth opens only at the very
top e.g., in H. triandra, H. crux-melitensis, H. sterilis and H. pulverulenta; in
others it is split to the base or nearly so e.g., in H. basifissa, H. angularis,
H. obscura and several others, but in most species it is split to about halfway.
New account of Horsfieldia 121
(c) The pedicels are usually cylindrical or subterete, sometimes subtriangular
(in species of the group of H. polyspherula), slender or thickish, and it may
be of taxonomical importance whether they are longer or shorter than the
perianth. In some species with pear-shaped or club-shaped flowers e.g., H.
tuberculata (partly so), or H. crux-melitensis, H. lancifolia, H. decalvata and
a Sia the base of the perianth gradually narrows into a tapering
pedicel.
In many species the pedicel is typically articulated at the base, at least in
dried specimens; however, in a few species including H. glabra this character
does not seem to be very constant. One should be aware that the bracteal scars
of the uppermost inflorescence may be mistaken for the articulation in the
pedicel and therefore several pedicels should be checked. Bracteoles are lack-
ing in Horsfieldia, as already noted.
(d) The androecium of the male flowers shows much diversity in shape and
construction and provides the most important characters in the delimitation of
species. Again, at the generic level, the androecium also provides major
characters.
In Horsfieldia, as in the other Asiatic Myristicaceae, the stamens are
always connate through presumably the fusion of filaments into an an-
drophore, which usually bears a so-called central column. The androphore is
generally rather short and may be absent. The degree of coalescence of the
anther-bearing part or synandrium varies according to the species; in many,
the anthers are completely connate or almost so, at their back, forming a
broad central column usually of a rather spongy texture and showing much
specific variation in shape. In many other species the coalescence of the
anthers is only partial and the anthers may then have free apices of various
lengths, depending on the species. Opening of the thecae is extrorse to /atror-
sus (directed sideways).
A well-developed central column, if present, may be solid, but in the
majority of the species it is hollowed out in the apical portion in very diverse
ways and to various depths, according to the species. The anthers may be erect
or suberect, or curved and following the straight or rounded shape of the
central column. The free portions of the anthers, if prevalent, may stand erect
or curve inwards into the central-apical cavity of the central column to
various depths according to the species. The androecium may be elliptic in
transverse section, as in most species with a laterally compressed 2-valved
perianth (sect. /rya); or circular or subcircular as in most species with 3 or
4-valved perianths (sections /ryaghedhi and Pyrrhosa). In the group of
H. polyspherula (sect. Pyrrhosa) the anthers are typically erect or suberect,
mutually free for the upper half while the androecium is typically triangular in
transverse section.
Each anther consists essentially of a pair of bisporangiate lobes or thecae,
comparable to those of the presumably more primitive condition found in
Compsoneura and Dialyanthera (S. America), or Brochoneura (Africa) where
the anthers are monadelphous with their filaments fused only towards the
base. Armstrong and Wilson (1978, fig. 16) postulate that there is within
Horsfieldia an evolutionary trend in the androecia from a condition of a
relatively poorly developed central column, or a small degree of fusion of the
connective columns and hence with more protrusion of the anthers, towards
one with a greater fusion in the connective column (i.e.,a broad and well-
developed central column) and a reduction in the protrusion and pairing of the
anthers. The least specialized conditionsi.e., with relatively free anthers,
would be those as found in species like H. irya, H. parviflora, or H. ardisii-
folia, all with a predominantly 2-valved perianthi.e., in sect. /rya.
Schematic longitudinal and transverse sections of the typical androecium
forms of almost all species are represented in figure 1A-D. I have refrained
122 Gard. Bull. Sing 37(2)(1984)
from hypothesising as regards which forms may be more primitive or more
advanced, though in general, I agree with Armstrong and Wilson.
The number of anthers is generally a good species character, although in
some cases, it may vary considerably. When the anthers are tightly set, they
may be difficult to count, especially because they are much elongated and con-
sist of two equal thecae.
The size of the androecium in relation to the perianth is sometimes signi-
ficant; usually it almost completely fills the perianth, but ine.g., H. amplifor-
mis, the perianth is largely hollow because the androecium occupies only a
fraction of the space.
It is peculiar that in juvenile stages the thecae of the anthers may be
typically septate, the septa being of non-tapetal origin; this phenomenon has
been studied by Armstrong and Wilson (1978). I am not sure whether it occurs
in all species of Horsfieldia. Such a septation is unknown in Mpyristica and
Knema.
(9) Pistillate flowers
The female perianth is usually subglobose, ellipsoid or ovoid, and the shape
varies somewhat with the species. In most species the female flowers are larger than
the male for example, distinctly so in H. thorelii, but many exceptions exist. Within
the same species the number of perianth-valves is always similar in male and female
flowers, but the extent of the split in female flowers is usually considerably less.
The globose, ellipsoid or ovoid ovary may be glabrous or hairy, according to
the species. There is some variation in the size and shape of the stigma, which is
essentially 2-lipped and the style is usually absent or very short. In general, the small
differences in the stigma are difficult to describe and this has not been exhaustively
studied by me. H. squamulosa, for instance, has a relatively long, erect, slender style
and stigma, which show up as a minute appendage on the young fruit; in H. sepiken-
sis the stigma is sessile and conspicuously broad-lipped; in H. iryaghedhi the sessile
stigma tends to be few-lobulate and not 2-lobed.
(10) Fruits
The fruit agrees in general appearance with other myristicaceous fruit, but is
in Horsfieldia (and in the genus Knema) characterized by the aril being either com-
plete or split into short laciniae in the apical portion only. The pericarp is glabrous
or pubescent; in the latter case the hairs may remain only at the very base of the
fruit, near the insertion of its stalk. Fruit size is largely diagnostic for the species.
The fruit of H. grandis, c. 1 cm long, is among the smallest, that of H. punctatifolia,
up to 8 cm long, is among the largest in the genus. In variable species like H. poly-
spherula and H. pallidicaula, | have accepted several varieties mainly on the basis of
fruit size. The shape of the fruits is usually short-ellipsoid, and of the seed, ellipsoid;
only in a few species is the fruit globose or nearly soe.g., in H. subtilis var. subtilis
and H. sinclairii, but their seeds are short-ellipsoid; in H. irya fruit and seed are
globose.
The pericarp varies in thickness according to the species; it is hard-succulent
in the fresh state, shrinking considerably on drying. For example, the pericarp of
large fruits of H. punctatifolia would usually shrink from a 4-cm thickness when
fresh to 1 cm. The surface of the dry pericarp is in many species provided with con-
spicuous pale warts or dots, apparently becoming more pronounced with drying,
showing up as lenticel-like tubercles. A striking example is the fruit of H. /aevigata.
Finally, it may be remarked here that in several species the female perianth
remains persistent under the fruit. This phenomenon occurs in species which are, if
considered on other grounds, not regarded as closely related. However, it provides
an easily observed character for species like H. kingii, H. wallichii, H. splendida, H.
pallidicaula and others.
New account of Horsfieldia 123
Description and Redefinition of the Genus Horsfieldia
Horsfieldia Willd.
Horsfieldia Willd., Sp. Pl. 4 (1806) 872 [non Bl. = Harmsiopanax Warb. (Aral.)]; Pers., Symb. 2 (1807)
635; Warb., Mon. Myrist. (1897) 130, 262; Sinclair, Gard. Bull. Sing. 16 (1958) 368; 27 (1974)
133-141; 28 (1975) 1-181. — Pyrrhosa Endl., Gen. Pl. (1839) 830 (see note). — Type: Horsfieldia
odorata Willd. =H. iryaghedhi (Gaertn.) Warb.
Myristica sect. Pyrrhosa BI., Rumphia | (1837) 190-192, Tab. 62-64. — With subsequent authors the
genus Horsfieldia as presently defined was treated partly under Myristica sect. Pyrrhosa as well as
under several other sections of Myristica viz., sects. Caloneura p.p., Eumyristica p.p., Horsfieldia,
Irya); see Sinclair (1958), p. 368 and under the presently accepted sections. — Lectotype of Myristica
sect. Pyrrhosa: Myristica glabra B\. = Horsfieldia glabra (Bl.) Warb.
Shrubs or usually trees, 2-40 (-60) m, dioecious. Twigs usually early glabrescent,
terete or sometimes angular or with two raised lines or ridges from petiole to petiole,
bark usually striate, always lenticellate but sometimes inconspicuously so. Leaves
distichous or in some species dispersed, usually petioled, blades up to 45 cm long,
membranous to coriaceous, often brittle when dry, pubescent or glabrescent, nerves
prominent or not above, reticulations above usually lax, never forming a dense,
close network as in Knema, lower surface not glaucous, papillose (alveolar tissue)
only in H. iryaghedhi; in some species non-traumatic cork warts present. In-
florescences axillary, situated in between and behind the leaves, rarely on the older
wood (H. sabulosa), paniculate, usually branched several times, pubescent or
glabrescent, at base of common peduncle with a few minute cataphylls; female in-
florescences usually smaller than the male. Flowers usually pedicelled, glabrous or
pubescent, solitary or in loose, sometimes sub-umbellate clusters or fascicles,
generally all of about the same age in the same inflorescence, in H. iryaghedhi the
male with flowers sessile in dense flower-heads; bracts broad-triangular to elliptic or
boat-shaped, caducous. Perianth thin-leathery to succulent, either 2 (or 3) or (2 or)
3- or 4-lobed, inside glabrous, greenish to yellowish, never red, the lobes splitting the
perianth to various depths, never spreading; bracteoles absent. Male perianth usual-
ly small, either globose, depressed-globose, transversely ellipsoid, ellipsoid,
reniform, pear-shaped, or clavate, laterally compressed or not. Androecium very
diverse in shape, either cupshaped, or globose to ellipsoid, cylindrical, or trigonous,
laterally compressed or not, sessile or with short androphore; the anthers usually
entirely or largely mutually connate and adnate at their back into a narrow or broad
central column various in shape, the latter usually with a depression or excavation to
various depths at apex. Anthers 2-c. 25, either straight and + erect, or curved or the
apical parts incurved or inflexed into the cavity of the column to various depths;
thecae often septate in the juvenile stage, opening extrorsely. Female perianth
usually larger than the male, subglobose to ovoid-ellipsoid; ovary globose or ovoid,
glabrous or pubescent, style absent, stigma 2-lobed or 2-lipped, usually small, more-
lobed only in H. iryaghedhi. Infructescences of moderate size, smaller than male in-
florescences. Fruits globose or usually ellipsoid, pericarp usually somewhat fleshy,
drying brown or blackish, often with lenticel-like tubercles, glabrous or pubescent,
or subglabrescent, perianth sometimes persistent under the fruit; seed ellipsoid, rare-
ly globose, testa not variegated; aril completely covering the seed, entire or at apex
shallowly lobed or convoluted; albumen ruminate, with fatty oil but no starch;
cotyledons connate at base.
NOTES
Fieldnotes. Trees of primary rain forest, often persisting in secondary growth;
also in marshy forest, and stilt roots present in some species.
According to Sinclair (1958) the bark of Malayan species is usually reddish-
brown, smooth or more often striate or rough with circular or irregular dents,
sometimes flaking but mostly not. The flowers are mostly waxy yellow, and often
sweet scented; those of H. iryaghedhi have a particularly strong smell.
124 Gard. Bull. Sing 37(2)(1984)
Anatomy. Koster and Baas (1981) published a paper on the comparative leaf
anatomy of the Asiatic Myristicaceae and found the genera anatomically quite
distinct, especially in the hairs and the vascular pattern of the midrib and petiole;
most species examined can be distinguished by their epidermal features.
Taxonomy. After having examined all species of Horsfieldia, including
H. macrocoma, {| came to the conclusion that within H. macrocoma s.\. four species
could be distinguished. These rather deviate from all other Horsfieldias and I have
segregated them into a new genus Endocomia (1984), which differs from Horsfieldia
by the following characters: flowers monoecious; flowers at the ultimate ramifica-
tions of the inflorescences developing in rather different stages; perianth inside
hairy or with appendages, at anthesis cleft to the base into (3 or) 4 (or 5) valves and
these spreading or recurved and not erect; androecium long-stalked, of different
construction; aril generally laciniated to about halfway the seed or more; testa of the
seed variegated.
Synonymy. Pyrrhosa Endl. is illegitimate because Horsfieldia Willd., as genus,
is cited in its synonymy; according to art. 7, 11 of the Code, Pyrrhosa Endl. should
be typified by Horsfieldia odorata Willd., being the type of the name which should
have been adopted. Pyrrhosa BI. as section is presently lectotypified by Myristica
glabra, a practice initiated by Hook. f. and Thomson (1855) with the division of
Mpristica sect. Pyrrhosa Bl. into three sections of smaller circumscription viz., sect.
Eumyristica, sect. Irya, and sect. Pyrrhosa; these three coincide largely with the
three presently accepted sections in Horsfieldia.
Table I
Table I enumerates all species in Horsfie/ldia and indicates by horizontal lines
demarcations between accepted sections and informal groups of species; broken
lines indicate marked discrepancies within a group as Is discussed in the text.
The distribution of three marked characters which are used at species level, but
not primarily used to distinguish sections or groups, is indicated with symbols before
the species names. A deviating perianth valve-number within a section is indicated
between the brackets; [] = pedicel articulated at base, + = perianth persistent
under the fruit, © = lower leaf surface with coarse dark-coloured dots from cork
warts.
17. H. olens (valves 3)
18. H. sepikensis (valves 3)
. Iryaghedhi
. kingii (valves 4)
p)
3. H. longiflora (valves 2) sylvestris
4 . thorelti (valves 2 or 3) :
5
. amygdalina (valves 2 or 3) . australiana
. crux-melitensis
22. H. clavata
. spicata
: . squamulosa
. H. inflexa
. H. moluccana . ampla
10. H. parviflora ae _ ampliformis Pella lee
11. H. obscurinervia H. angularis (valves 2-4)
12. H. ardistifolia . H. triana
13. H. talaudensis __H. aruana oe
14. H. samarensis MED tilis
bey ASE _ H. schlechteri
H. palauensis.\)\ ©) 9 99701 t5im = al 2 le ee
. H. basifissa
New account of Horsfieldia
(QO)
32. H. sinclairii
33. H. psilantha
34. H. whitmorei
35. H. laevigata
36. H. pilifera
37. H. lancifolia
38. H. decalvata
H. tuberculata
H. corrugata
41. H. pachycarpa
H.
. pulverulenta
. leptantha
. hellwigii
. ralunensis
. sabulosa
. atjehensis
- SUCOSA
. Sparsa
. triandra
. tristis
. fulva
. superba
H.
H.
H.
H.
H.
H.
. H. pallidicaula
H.
H.
H.
H.
H.
H. sessilifolia
H.
. grandis
motleyi
. tomentosa
. gracilis
. paucinervis
. splendida
an
LS) ;
By sae Sy
. rufo-lanata
ap ey Wea) fea) ea]
a
(i) a) a
O
125
. affinis
. reticulata
. crassifolia (valves 2)
. carnosa
. Sterilis (valves 2)
. hirtiflora
. brachiata
. pachyrachis
ridleyana
obtusa
disticha
. tenuifolia
macilenta
. laticostata
. nervosa
. polyspherula
. oligocarpa
~
m|[rzaarzzrzzrzzzzlsl(zlzla2
ee A de
. valida
borneensis
. fragillima
. androphora
TuTtIrt.z
. amplomontana
. H. montana
90. H. punctata
. subalpina
. obscura
. coriacea
. penangiana
. punctatifolia
. macrothyrsa
os Se | ae
. glabra
Subdivision of Horsfieldia into Three Sections
As can be seen in the preceding survey of characters to be used in the taxonomy
of Horsfieldia as well as from the species descriptions, there is a large diversity in the
genus. This led Warburg (1897) to subdivide the genus into 3 sections, with 5 sub-
sections and 2 series, with 11 additional accepted species not placed in any section
because their male flowers were unknown. Warburg’s subdivision appears to me
very artificial, at least for the greater part. For the Malayan species only, Sinclair
(1958) advocated a more simple subdivision into ‘‘Bivalves’’ and ‘‘Trivalves’’, and
some unnamed subsections, but a definite decision, pending on the study of the
whole genus, was frustrated by his early death.
126 Gard. Bull. Sing 37(2)(1984)
I have chosen to subdivide the 100 presently accepted species into three sections
although I realize that these are of considerably unequal taxonomic weight. The
three are: (1) sect. Horsfieldia, containing one single species, being the type species
of the genus, but rather deviating from all others, (2) sect. Jrya, containing most
species with a predominantly 2-valved perianth, and (3) sect. Pyrrhosa with most
species having predominantly a 3- or 4-valved perianth. Although the two last-
named sections, which are by no means sharply segregated but which have their own
range of distribution, coincide with Sinclair’s subdivision, they should be named dif-
ferently on nomenclatural grounds. Each contains a few species with the perianth
predominantly of a deviating valve-number, but it will be argued in the discussion
on the groups that these species are tentatively best regarded as casual deviations still
to be placed in that section.
As regards a further formal subdivision, I have refrained from making any. In-
stead, I have listed in Table I all the species and indicated with horizontal lines
(either continuous, or broken, according to the supposed strength of the demarca-
tions) which groups or clusters can be distinguished. These groups, without formal
status, contain species which are presumably closer to each other on grounds to be
discussed in the next chapter. At first sight several of these groups of more or less
coherent species seem to represent formal entities e.g., subsections or series, but in
view of the diversity in the variation of all species concerned, this certainly would
lead to a complicated system of entities differing much in taxonomic weight and
with blurred circumscriptions.
It is obvious that the species in general have reticulate relationships, and the
groups are only consistent on the basis of a single or a few putatively important
characters. Other characters, which could be regarded as important as well will often
be found distributed in various other groups, without necessarily expressing that
such characters, respective to their groups, are less valuable or the groups unnatural.
The erratic distribution of such characters not used to assemble species into groups,
as the presence of non-traumatic cork warts on the lower leaf surface, the presence
of an articulation at the base of the pedicel, or the calyx being persistent under the
fruit, is indicated in the list of species in Table I, according to the symbols explained
in the legend. I have refrained from arguing about any phylogenetic value of these
groups as I feel that it is arbitrary and difficult to judge what characters should rank
phylogenetically behind others, and often, one can only guess which character-states
should be ranked as more primitive or more advanced.
The sections and groups will be discussed briefly below, with references to links
with other groups or individual species. The majority of the characters binding a
group can be extracted from those used subsequently in the general key to the
species.
The Sections and a Survey of the Groups
1. Sect. Horsfieldia
Mpristica sect. Horsfieldia A. DC., Prod. 14 (1856) 200; Miq., Fl. Ned. Ind. 1, 2 (1859) 63. — Myristica
sect. Jrya auct. non Hook. f. & Th.: Benth. & Hook.f., Gen. Pl. 3 (1880) 137, for Horsfieldia Willd.
only. — Myristica sect. Eumyristica subsect. Horsfieldia (A. DC.) King, Ann. Roy. Bot. Gard. Cale.
3 (1891) 282. — Type: Horsfieldia odorata Willd.
Mpyristica sect. Pyrrhosa Bl|., Rumphia 1 (1835) 190, p.p., for M. horsfieldii only, not the lectotype. —
Mpristica sect. Eumyristica Hook.f. & Thomson, FI. Ind. (1855) 162, p.p., for M. horsfieldii only.
Horsfieldia sect. Orthanthera Warb., Mon. Myrist. (1897) 268, p.p., for the lectotype only. —
Horsfieldia sect. Triavalves subsect. Orthanthera (Warb.) Sincl., Gard. Bull. Sing. 16 (1958) 371,
p.p., vom. inval., provisional name only. — lectotype: Horsfieldia iryaghedhi (Gaertn.) Warb.
Phyllotaxis of plagiotropic shoots distichous. Lower leaf-surface with alveolar
tissue, epidermis papillose, stomatal complex sunken; without larger dark coloured
New account of Horsfieldia 127
dots (cork warts). Flowers in male sessile, at base not articulate, arranged in many-
flowered subglobose dense heads; perianths elongate-obconical, before anthesis +
angled, 3- or 4-(in 2 :2- or 4-) valved, splitting the bud to c. 1/2-3/4. Androecium
elongate, sub-cylindrical, androphore distinct; central column narrow, narrowly
hollowed out for over 1/2-way; anthers 3-5, erect. Stigma sessile, many-lobulate.
Ceylon. — Figure 1, I; 21; species 1.
This section is monotypic with H. iryaghedhi', a species deviating from all other
Horsfieldias by the following characters: — some anatomical characters of the leaf,
the male flowers being: sessile and arranged in dense heads which have a thick recep-
tacle, the anthers being mostly connate but in most instances not back to back so
that a narrowly hollowed central column is thereby formed, the angular perianths,
and the many-lobed (not 2-lobed) stigma in the female flowers.
2. Sect. Irya (Hook.f. & Th.) Warb.
Myristica sect. Pyrrhosa B\., Rumphia 1 (1837) 190, p.p., for M. javanica and a few other species only,
excl. lectotype M. glabra (sect. Pyrrhosa) and M. horsfieldii (= H. irvaghedhi, sect. Horsfieldia);
A.DC., Prod. 14 (1856) 202, p.p.; Miq., Fl. Ned. Ind. 1, 2 (1859) 64, p.p., excl. M. glabra (= sect.
Pyrrhosa).
Myristica sect. Irya Hook.f. & Thomson, Fl. Ind. (1855) 159; A.DC., Prod. 14 (1856) 202; Miq., Fl. Ned.
Ind. 1, 2 (1859) 64; Benth. & Hook.f., Gen. Pl. 3 (1880) 137, p.p., excl. sect. Horsfieldia; King, Ann.
Roy. Bot. Gard. Calc. 3 (1891) 284, p.p., for the smaller part only. — Horsfieldia sect. Irva (Hook.f.
& Th.) Warb. subsect. Euirva Warb., Mon. Myrist. (1897) 123, 267, p.p., for the type only. — Type:
Mpristica irya Gaertn. = Horsfieldia irya (Gaertn.) Warb.
Horsfieldia sect. Pyrrhosa subsect. Bivalves Warb., Mon. Myrist. (1897) 262 (incl. series Smithii and
sezies Globularia). — Horsfieldia sect. Biva/ves Sinclair, Gard. Bull. Sing. 16 (1958) 370, 371, comb.
inval., provisional name only. — Type not indicated, several species listed.
Phyllotaxis it: plagiotropic shoots distichous. Lower leaf surface without
alveolar tissue, epidermis not papillose, stomatal complex not sunken; rarely
(H. inflexa) with larger dark coloured dots. Flowers pedicelled, base not articulate,
solitary or in loose clusters; perianth rarely globose or obovoid, usually laterally
compressed, in outline either circular or elliptic or pear-shaped, before anthesis not
angular or in young stages of H. sy/vestris faintly so, usually all or predominantly
2-valved, rarely 3-valved, cleft at anthesis to various depths even nearly to the base.
Androecium various, nearly always more or less zygomorphic, either usually lateral-
ly compressed or in some species with 3- (or 4-) valved flowers, the androecium
broad-ellipsoid with the anthers at apex bi-laterally incurved, rarely obconical and
actinomorphic; androphore distinct or not, central column broad or narrow, little to
much hollowed; anthers few to many, erect or sub-erect, or some or all incurved
over or to various depths into the cavity of the central column. Stigma minutely
2-lobed. Mainly E. Malesia (incl. Philippines), only H. irya distributed over almost
the whole area of the genus. — Figure 1, 6-45; 2 II; species 6-45.
Almost all of the 40 species of this section have exclusively or predominantly
2-valved perianths in the same inflorescence, and usually have a more or less
zygomorphic androecium, either because it is laterally compressed or because the
anthers curve in apically from two opposite sides only.
Aberrant are H. olens!’, H. sepikensis'® with 3- or 4-valved perianths, but with
the androecium distinctly tending to be zygomorphic, the anthers from two
opposite sides incurved in a manner reminiscent of the condition as found in species
like H. parviflora, H. moluccana; H. angularis* has a 2-4 valved perianth.
Also aberrant are the species of the group of H. clavata ”?, with a 2-lobed
perianth but a club-shaped non-zygomorphic androecium.
A few mutually related species from continental SE. Asia, presently placed in
the next section, (3) Pyrrhosa, viz. H. longiflora, H. thorelii and H. amygdalina
128 Gard. Bull. Sing 37(2)(1984)
have all or partly predominantly a 2-valved perianth, and partly a zygomorphic
androecium, especially H. /ongiflora. This latter species blurs the distinction bet-
ween sect. /rya and sect. Pyrrhosa, because, if judged from its morphology of the
perianth and androecium, it seems to agree with sect. Jrya. However, it clearly
belongs to the group of the other above-named species, which obviously link up with
sect. Pyrrhosa. Moreover, since sect. /rya is mainly of East Malesian distribution,
H. longiflora would be occupying an aberrant locality as it is restricted to Indo-
China. Compared with other species in sect. Jrya, H. irya extends far beyond their
range of distribution; besides, its relationship with H. /ongiflora is rather remote.
The phylogenetic backgrounds of this strayed species within sect. Pyrrhosa, known
only from a few Vietnam collections, pose a problem. Within sect. Jrya, 8 groups
(1—8) can be distinguished.
SURVEY OF GROUPS WITHIN SECT. IRYA
(1) H. irya-group
H. irya-group consists of a single species, H. irya®, which occupies a rather
isolated position. It is distinct in habit, with thin leaves often provided with irregular
pale blotches, and unique by having globose fruits and seeds, the globose 2-valved
male perianth and a typical androecium which is only slightly laterally compressed,
and with the anthers almost free, and attached to the rim of the cup-shaped an-
droecium. It is linked with the next group by H. palauensis'® which occupies a rather
intermediate position between H. irya and H. smithii’>.
(2) H. parviflora-group, species 7-16
Group 2 is chiefly characterized by the distinctly laterally compressed, male
perianth, which is either + pear-shaped or circular or more or less transversely ellip-
tic or reniform in lateral view. The laterally compressed androecium follows the
shape of the male perianth and consists of anthers connate to various degrees and
deeply curved inward into the central cavity at one or both sides of the androecium.
This is partly Warburg’s sect. Pyrrhosa: subsect. Bivalvis: series Globularia and
series Smithii.
(3) H. olens — H. sepikensis-group
The two species of group 3, 17 and 18, are aberrant in the section because of
their 3-valved perianths. The androecium is rather broadly obovoid and only slightly
zygomorphic, but the apical parts of the anthers curve inwards + bilaterally, in a
manner reminiscent of that in the foregoing group.
(4) H. sylvestris-group
H. sylvestris-group contains a sole species (16), occupying an isolated position,
and is readily recognized by a number of characters. Warburg (1897) placed it with
H. iryaghedhi and H. ralunensis in a separate section called Orthanthera because the
male flowers are elongate and densely clustered in flower heads, but this latter
feature only holds for H. iryaghedhi, because in H. sylvestris similar flower heads
are only present in juvenile inflorescences, which later on expand into separately
attached flowers. The flowers of H. sylvestris have an elongate perianth and
androecium, not or but little compressed, and the anthers are mutually completely
connate forming a narrow, completely solid, central column.
(5) H. australiana-group
A rather isolated species, H. australiana”’, is the sole member. Sinclair had
assigned plants from New Guinea to this species but here I accept them as belonging
to a distinct species: H. sinclairii. In H. australiana the androecium is rather broad-
ovoid, overarching over a broad and + shallow, apical hollow, into which the
central column protrudes conspicuously. Based on the flower-features, H. sinclairii
is assigned to the group of H. /aevigata*’.
Editorial Note; superior figures following species names are numbers of the author’s accepted species.
|
g
WY
CG
S) :
ALAS
Figs. 1A-D Semi-schematic drawings of the androecium of most species of Horsfieldia, all depicted ex-
cept species 24, 55, 62, 76; lateral view (left), longitudinal section (right), apical view (top);
white: anthers, black: sterile tissue (i.e. androphore and central column).
19
Fig. 1A. Species 1-23: /, H. iryaghedhi (x 10, Jayasurika & Bandaranaika 1869); 2, H. kingii (x 10,
Haines 842); 3, H. longiflora (x 10, Eberhardt 3050); 4, thorelii (x 10, Poilane 19887); 5, H.
amygdalina var. lanata (x 10, Kerr 8556); 6, H. irya (x 10, Muller 1020, from Indo-China); 7,
H. spicata (x 5, Beguin 1407); 8, H. inflexa (x 5, LAE 52866); 9, H. moluccana var. moluccana
(x 5, Kostermans 673a); 10, H. parviflora (x 5, Ding Hou 134); Il, H. obscurinervia (x 5, For.
Bur. 26503); 12, H. ardisiifolia (x 5, Cuming 1702); 13, H. talaudensis (x 5, Lam 2628); 14, H.
samarensis (x 5, PNH 117374); 15, H. smithii (x 5, Rutten 1776); 16, H. palauensis (x 5,
Takahide Hosakawa 6756); 17, H. olens (x 10, NGF 31966); 18, H. sepikensis (x 10, Hoogland
& Craven 10255); 19, H. sylvestris (x 10, Craven & Schodde 739); 20, H. australiana (x 5,
Dunlop. 3585); 21, H. crux-melitensis (x 5, Schlechter 19246); 22, H. clavata (x 5, Hoogland
3663); 23, H. squamulosa (x 5, Pullen 8287).
130 Gard. Bull. Sing 37(2)(1984)
(6) H. clavata-group, species 21-23
The three species in the H. clavata-group are from New Guinea. They form a
distinct group based on various characters dominated by the peculiar club-shaped,
not zygomorphic androecium with small sessile anthers at the apex. A thickened
sterile androecium with small, apically attached anthers is also found in H. pulveru-
lenta, H. sterilis and H. triandra but in a somewhat different manner. The species of
the H. clavata-group are shrubs or low trees.
(7) H. laevigata-group, species 24-4]
The group has mainly species from New Guinea. Several sub-groups can be
distinguished, but all are characterized by a laterally compressed perianth and
androecium. The anthers are erect and the rather narrow central column is narrowly
cleft to various depths.
The two species H. ampla’* and H. ampliformis?> are characterized by their
large male inflorescence and the androphore in H. ampliformis is conspicuous.
H. angularis, H. iriana and H. aruana, species 26-28, are distinct by their ridged
or angled twigs; H. angularis*° has the male perianth variably 2- to 4-lobed.
H. subtilis?? and H. schlechteri?° are small trees; both have the androecium
rather narrowly stalked with the central column somewhat thickened towards the
base, and hence the androecium is often slightly sagged.
Species 31-41 are rather heterogeneous and some have special features:—
H. basifissa*! and H. sinclairii?? both have rather small globose or subglobose fruits,
but in H. basifissa the perianth is deeply cleft; H psilantha*} comes close to H. /aevi-
gata*’ but its flowers are (almost) glabrous, the leaves are larger, etc., H. lancifolia’’
and H. decalvata*® from Celebes and the Moluccas respectively, stand apart within
the present group by their pear-shaped perianth with a tapering pedicel, but, the
wide-spread and variable H. tuberculata’’ usually has somewhat pear-shaped
flowers also; H. corrugata® and H. pachycarpa*' are mountainous species with very
large fruits as compared with the other species of the group.
(8) H. hellwigii-group, species 42-45
H. hellwigii-group is restricted to New Guinea and the Bismarck Archipelago,
and is generally easily recognized by the rather stout habit ((omewhat resembling the
wide-spread H. sylvestris) and the coarse tomentum on the leaf bud and twig apex;
H. pulverulenta®? is very distinct within this group by the peculiar androecium,
which is reduced to a largely sterile cone with 2 minute anthers at the top, and the
hard-fleshed perianth which opens apically only by a small slit just above the
anthers.
3. Sect. Pyrrhosa (Bl.) Warb.
Myristica sect. Pyrrhosa Bl., Rumphia 1 (1837) 190-192, Tab. 62-64, p.p., for the smallest part incl. the
lectotype Tab. 64, fig. 1A, B; Hook.f. & Thomson, Fl. Ind. (1855) 160; A.DC., Prod. 14 (1856) 202,
p.p.; Miq., Fl. Ned. Ind. 1, 2 (1859) 64, p.p.; Benth. & Hook.f., Gen. Pl. 3 (1880) 136; King, Ann.
Roy. Bot. Gard. Calc. 3 (1891) 282. — Horsfieldia sect. Pyrrhosa (Bl.) Warb. subsect. Eupyrrhosa
Warb., Mon. Myrist. (1897) 265 (excl. H. macrocoma = Endocomia gen. nov. — Lectotype:
Myristica glabra B\. = H. glabra (B\.) Warb.
Mpristica sect. Eumyristica Hook.f. & Thomson, FI. Ind. (1855) 162, p.p., for M. superba = Horsfieldia
superba (Hook.f. & Th.) Warb. only. — Myristica sect. Caloneura A.DC., Prod. 14 (1856) 192, p.p.,
for M. superba Hook.f. & Th. = Horsfieldia superba (Hook.f. & Th.) Warb. only.
Myristica sect. Irya auct. non Hook.f. & Th: King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 284, p.p.
Horsfieldia sect. Irya (Hook.f. & Th.) Warb. subsect. Euirya Warb., Mon. Myrist. (1897) 267, p.p;,
excl. H. irya (type).
‘Ni |e
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Fig. 1B. Semi-schematic drawings of Horsfieldia cont.
Species 24-54: (24, H. ampla, male flowers not extant); 25, H. ampliformis (x 10, Hoogland &
Craven 11085); 26, H. angularis (x 5, lwanggin BW 5828); 27, H. iriana (x 5, Zipelius 139d);
28, H. aruana (x 5, Zipelelius s.n.); 29, H. subtilis var. subtills (x 5, Versteegh & Vink BW
8377); 30, H. schlechteri (x 5, NGF 13293); 31, H. basifissa (x 5, Womersley 3821); 32, H.
sinclairii (x 5, NGF 28886, type); 33, H. psilantha (x 10, Sands et al. 2047); 34, H. whitmorei (x
5, BSIP 7565); 35, H. laevigata var. laevigata (x 5, Craven & Schodde 874); 36, H. pilifera (x 5,
Clemens 1710); 37, H. lancifolia (x 5, bb. Cel./I1. 464); 38, H. decalvata (x 5, Idjan & Mochtar
181); 39, H. tuberculata (x 5, Waterhouse 820B); 40, H. corrugata (x 5, Carr 14123); 41,
H. pachycarpa (x 5, Manner & Street 307); 42, H. pulverulenta (x 5, LAE 43567); 43, H. lep-
tantha (x 5, Vink BW 12194); 44, H. hellwigii var. hellwigii (x 5, NGF 26253); 45, H. ralunen-
sis) x 5, NGF 44388); 46, H. sabulosa (x 10, SAN 15146, immature); 47, H. atjehensis (x 10,
Bangham 882); 48, H. sucosa subsp. sucosa (x 10, SNF 40629); 49, H. pallidicaula var.
pallidicaula (x 10, Hose 29); 50 H. sparsa (x 10, FRI 7982); 51, H. triandra (x 5, Forbes 2465);
52 H. tristis (x 5, S 37470); 53, H. fulva (x 5, KEP 99334); 54, H. superba (x 5, FRI 4571).
131
132 Gard. Bull. Sing 37(2)(1984)
Horsfieldia sect. Irva (Hook.f. & Th.) Warb. subsect. Trivalves Warb., Mon. Myrist. (1897) 267. —
Horsfieldia sect. Trivalves subsect. Trivalves Sinclair, Gard. Bull. Sing. 16 (1958) 370, 371, comb.
inval., provisional name only. — Type not indicated.
Horsfieldia sect. Orthanthera Warb., Mon. Myrist. (1897) 268, p.p., for H. ralunensis and H. sylvestris
only, excl. the lectotype H. irvaghedhi.
Phyllotaxis of plagiotropic shoots dispersed or distichous. Lower leaf surface
without alveolar tissue, epidermis not papillose, stomatal complex not sunken; with
or without larger dark coloured dots or cork warts. Flowers with a pedicel (short in
H. wallichii), at base with or without articulation, solitary or in loose clusters;
perianth (depressed) globose, or obovoid or ellipsoid, not laterally compressed, not
or but faintly angular, generally perianths all or predominantly 3- (or 4-) valved,
rarely 2-valved (H. longiflora, H. sterilis), cleft at anthesis to c. 2/3 or less. An-
droecium actinomorphic or more or less triquetrous in transverse section, never
laterally compressed, depressed-globose or ellipsoid or obovoid, usually with a
broad central column with an apical hollow of various shape and depth; androphore
short or rather long, usually narrow at base; anthers various in number, + straight
or curved, either almost entirely connate or + straight-erect and mutually free for
about the upper half (H. polyspherula-group). Stigma minutely 2-lobed. Continen-
tal SE. Asia, West Malesia (incl. Philippines). — Figures 1, 2-5, 46-100; 2 III;
species 2-5, 46-100.
This section of 59 species contains mainly those with exclusively or
predominantly a 3- (or 4-) valved perianth, the rest are 3 species with a 2-valved
perianth, viz., H. longiflora from Vietnam, H. crassifolia (Malaya, Sumatra,
Borneo) and H. sterilis from Sabah. I believe that these exceptions should be placed
among the species of the present section Pyrrhosa as is briefly discussed in the notes
to the preceding section for H. /ongiflora, and argued for H. crassifolia and
H. sterilis below.
Within sect. Pyrrhosa some 15 groups (9-23) can be distinguished, some of
them are heterogeneous, others have a clear circumscription particularly the
H. polyspherula-group or contain only a single aberrant species.
SURVEY OF GROUPS WITHIN SECT. PYRRHOSA
(9) H.amygdalina-group, species 2-5
Group 9 clearly links up with the H. glabra-group (sp. 97-100) because of the
largely identical construction of the male flowers, and the phyllotaxis which is either
distichous or dispersed, but differs in the absence of cork warts on the lower leaf
surface. The group is keyed out at the beginning: — because it has a distinct
distributional area in continental SE Asia, and because of the variable number of
perianth-valves (2-4) and the variable phyllotaxis, and this situation rather interferes
with the keying out of most of the West Malesian species, which are clearly distinc-
tive. H. kingii is readily distinguished by its pubescent, 4-valved male perianth.
(10) H. sabulosa-group
Group 10 consists only of H. sabulosa®, which, from the study of the rather im-
mature male flowers, links up with the group of e.g. H. borneensis*’ or
H. punctata®. It is however readily distinguished by its ramiflorous flowering,
dispersed phyllotaxis, long-petioled leaves, dark-coloured dots or stripes on the
lower leaf surface, etc. The leaves have an iso-bilateral anatomy as was discovered
by Koster and Baas (1981).
——
New account of Horsfieldia 133
(11) H. pallidicaula-group, species 47-50
The species in group 11 are usually easily distinguished by the pale, grey-brown
or straw-colour of the dried twigs, contrasting with the blackish colour of the dried
leaves. Usually the flowers, leaves and fruits also dry to a relatively black colour.
The phyllotaxis is mostly dispersed. In H. sucosa*® and H. pallidicaula*p.p. the
calyx remains persistent under the fruit; H. atjehensis*’ is distinct by the presence of
blackish brown dots or cork warts on the lower leaf surface. The architecture of the
androecium of this group links up with that of many other species, e.g., the
H. glabra-group. In H. sucosa subsp. bifissa**® the perianth is predominantly
2-valved, which is aberrant in the present section.
(12) H. triandra-group
Group 12 consists of a single species, H. triandra*', from C. and S. Sumatra, and
is distinct by flowers in that the perianth is thick-leathery and opens by 3 valves split
to only c. 1/5, and in the turbinate androecium, with 3 reduced anthers at the top.
Similar androecia with ‘reduced’ anthers occur ine.g., H. sterilis, H. pulverulenta,
and the H. clavata-group of sect. Jrya, but I doubt that these species are otherwise
related.
(13) H. fulva-group, species 52-54
These three species occupy a separate position because of the leaves drying
to a dull colour above. The male perianth and androecium are of an elongated
shape. In H. fulva and H. superba the perianth remains persistent under the fruit.
(14) H. sessilifolia-group
Group 14 consists of a possibly isolated species (55), characterized by the
almost-sessile leaves; its male flowers are not known.
(15) A. grandis — H. motleyi-group, species 56-60
Group 15 is named here merely for convenience, as its five constituents, H.
grandis®*, H. wallichii’’, H. pulcherrima®®, H. flocculosa’’, H. motleyi® all differ
considerably from each other in general habit as well as the shape of the male
perianth and especially the androecium. Only H. grandis and H. pulcherrima seem
more closely related according to the male flowers, but for the rest, all five species
could represent separate groups as well. H. flocculosa*’ has a somewhat elongated
male perianth and an elongate androecium, and therefore suggests connections with
the H. fulva-group.
(16) H. affinis-group, species 61-67
Group 16 is characterized by a globose or depressed globose androecium with a
moderate apical hollow, and a short and slender androphore; in these it seems much
related to the group of H. endertii — H. montana (83-89). Our present group,
however, is segregated by a generally persistent tomentum on the lower leaf-surface.
H. affinis© and H. reticulata’ have very distinctly articulated pedicels.
(17) H. crassifolia-group
Group 17 consists of only one species, H. crassifolia®’, which deviates in many
respects viz., a 2-valved perianth, an actinomorphic subglobose androecium with
largely free anthers (fig. 1C), a persistent perianth under the fruits and a dotted
lower leaf surface. The pedicels at the base are not articulated. Because of the large-
ly free anthers and the 2-valved perianth, this species may have affinity to the wide-
spread H. irya® from section /rya.
(18) A. carnosa-group
Group 18 possibly should not have been mentioned as a group, because its
single species (69) links up with species from the group of H. borneensis** by its male
134 Gard. Bull. Sing 37(2)(1984)
flowers. Vegetatively and ecologically (heath forest, peat swamp forest) ‘it is,
however, quite distinct.
(19) H. sterilis-group
The position of group 19, with its single species H. sterilis” from SE. Sabah
is problematic within section Pyrrhosa. Its perianths are 2-valved, opening at
anthesis only at the very apex; the androecium is, however, actinomorphic,i.e.,
faintly blunt-triangular, and consists of a large sterile conical part with 6 anthers
apically attached (or possibly 3 anthers each with 2 thecae). It has twigs drying
rather pale and with dark leaves and is hence vegetatively reminiscent of the
H. pallidicaula-group with which it has no affinity if judged from the androecium.
The female perianth persists under the young fruit.
(20) H. polyspherula-group, species 71-82
Group 20 is remarkably coherent on account of the following characters: pedicels
articulated at the base, androecium (generally) triquetrous in transverse section,
anthers suberect and mutually free for about the upper half; furthermore, most
species (except H. ridleyana”) have the lateral nerves distinctly raised above. The
group can be divided in 2 subgroups based on whether the internodes are distinctly
ridged (or short-winged) or angular as in species H. hirtiflora’' and H. brachiata”,
respectively. In H. hirtiflora, a rather deviating species with its pubescent 4-valved
perianth, and leaves partly dispersed, the androecium is rather roundish, not
triquetrous in section, but the erect and largely free anthers still point to a relation-
ship within the H. polyspherula-group. This group correlates with Warburg’s sect.
Irya subsect. Trivalves.
(21) The group of H. endertii®, H. valida**, H. borneensis**, etc. to H. obscura”
(spec. 83-93) is rather heterogeneous, but most of its species are clearly interrelated
as they have in common a depressed globose narrowly-stalked androecium with a
moderately deep, apical hollow. In H. fragillima® this hollow is exceedingly broad,
rendering the androecium saucer-shaped; in H. androphora*’ the androphore is con-
spicuously elongated; H. borneensis** has distinctly articulated pedicels and on the
lower leaf surface blackish marks, presumably cork warts. Through H. punctata®,
with a dotted lower leaf surface, there is a strong connection of the present group
with the group of H. glabra'’™. Species 83-89 all have in common a coarse tomentum
on the leaf bud. Through its coarse pubescence H. endertii*’ leads the present group
though it deviates by its elongate male perianth and androecium, which it shares
with the species of the next group.
(22) H. xanthina-group, species 94-96
Species of group 22 and H. endertii®’? of the foregoing group have in common
an elongate male perianth and androecium. Whether these species are in reality more
Facing Page
Fig. 1C. Semi-schematic drawings of Horsfieldia cont.
Species 55-89: (55, H. sessilifolia, male flowers not known); 56; H. grandis (x 10, Chew Wee-
lek 261); 57, H. wallichii (x 5, FRI 12135); 58, H. pulcherrima (x 10, FRI 8008); 59, H. floc-
culosa (x 5, KEP 110225); 60, H. motleyi (x 10, Kostermans 6859); 61, H. tomentosa (x 10, FRI
15957); (62, H. gracilis, male flowers not known); 63, H. paucinervis (x 10, Haviland 3075);
64, H. splendida (x 10, S 33723); 65, H. rufo-lanata (x 10, Richards 1667); 66, H. affinis (x 10,
S 24718); 67, H reticulata (x 10, Hans Winkler 388); 68, H. crassifolia (x 10, SEN 40898); 69,
H. carnosa (x 10, van Niel 4519); 70, H. sterilis (x 10, SAN 30597); 71, H. hirtiflora (x 10,
Rahmat si Boeea 9257); 72, H. brachiata (x 10, King’s Coll. 4704); 73, H. pachyrachis (x 10,
bb. 28128); 74, H. ridleyana (x 10, Scortechini 12); 75, H. obtusa (x 10, Native Coll. BS 821);
(76, H. disticha, male flowers not known); 77, H. tenuifolia (x 10, S 24945); 78, H. macilenta
(x 10, B.N.B. 4204); 79, H. laticostata (x 10, S. 17252); 80, H. nervosa (x 10, S 16652); 81,
H. polyspherula var. polyspherula (x 10, KEP 38/29); 82, H. oligocarpa (x 10, Ashton &
Whitmore BRUN 398); 83, H. endertii (x 5, Endert 3996); 84, H. valida (x 5, Lorzing 5896); 85,
H. borneensis (x 10, S 14610); 86, H. fragillima (x 10, Bakar S 4361); 87, H. androphora (x 10,
S 35443); 88, H. amplomontana (x 10, SAN 18843); 89, H. montana (x 10, Sinclair 8987).
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closely inter-allied remains uncertain. The androecium of H. majuscula® is
triquetrous in section and the pedicel is articulated at the base, and therefore this
species might be close to the H. polyspherula-group (species 71-82).
(23) H. glabra-group, species 97-100
As already mentioned under the H. amygdalina-group (spec. 2-5), and e.g., also
in the comments on H. punctata®, H. glabra-group has several connections with
Others, especially in its largely similar, subglobose or ellipsoid androecium with a
135
136 Gard. Bull. Sing 37(2)(1984)
smallish apical cavity. The group is readily distinguished by the very short tomentum
of the leaf bud and of the inflorescence, and by the presence of dark-coloured dots,
i.e. cork warts, on the lower leaf surface.
Geographical Distribution
Horsfieldia, with 100 species, ranges from Ceylon through NE. India to S.
China (Kwangsi, Hainan) and throigh Malesia and the Caroline Isls. east to the
Solomon Isls. and N. Australia. It is absent from the Lesser Sunda Isls. Apart from
a few widely distributed species, e.g., H. irya, H. glabra, H. amygdalina,
H. laevigata, or H. tuberculata, most species are of limited distribution. Distinct
centres of species-development are New Guinea and Borneo, and to a lesser extent
Malaya-Sumatra. Here recognized are three sections, which occupy largely mutually
exclusive areas. Section Horsfieldia, with only H. iryaghedhi, is confined to
Ceylon. Section Jrya (with 40 species) is, except for the wide-spread H. irya (see fig.
2), confined to E. Malesia and the Solomon Isls. and N. Australia, as indicated in
figure 2. The section Pyrrhosa occurs west of Wallace’s Line as also indicated in that
figure by a broken line.In distribution, Sections /rya and Pyrrhosa overlap for a
relatively narrow area in the Philippines and Celebes. They are segregated mainly
because of a different valve-number of the perianths viz., predominantly 2 in
the east-occurring sect. Jrya and predominantly 3 in the west-occurring sect.
Pyrrhosa. Some species with the perianth-valve number deviating from that in the
section but still regarded as belonging to it are for sect. Jrya: H. olens!’, H. sepiken-
sis'’, and H. angularis’’. In the west-occurring section Pyrrhosa the deviating valve
number 2 is found in: H. longiflora?, H. thorelii# (and H. amygdalina’, for a
minor part), H. crassifolia® (see fig. 2) and H. sterilis”. Why these species are better
listed in this section is briefly explained in the foregoing chapter in the discussion on
sections and groups. Apart from these few species with a deviating valve-number,
the division into the sections Pyrrhosa and Irya based onthe character of the
perianth being 3- and 2-valved respectively, correlated with their respective dif-
ferent geographical distribution is peculiar because in most individual specimens the
number of perianth-valves is not at all constant: perianths with a deviating valve-
number are usually found in low percentages. In general, however, though not
strictly constant, even in the specimens, the character of the predominant valve-
number has apparently some phylogenetic significance connected with Wallace’s
Line.
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Fig. 1D. Semi-schematic drawings of Horsfieldia cont.
Species 90-100b: 90, H. punctata (x 10, FRI 90/4, slightly immature); 9/, H. costulata (x 10,
PNH 2685); 92, H. subalpina subsp. subalpina (x 10, Whitmore FRI 3884); 93, H. obscura (x
10, Kostermans 13773); 94, H. xanthina subsp. xanthina (x 10, Richards 1927); 95.
H. majuscula (x 10, Ramat si Boeea 8772); 96, H. coriacea (x 10, bb. Cel. III/-27); 97, H.
penangiana (x 10, Curtis 2406 in BM); 98, H. punctatifolia (x 10, S 36580); 99, H. macrothyrsa
(x 10, Lorzing 17195); 100, H. glabra var. glabra (x 10, Nengah Wirawan 95); 100 b, H. glabra
var. javanica (x 10, Koorders 21635 B ).
= = T T eS a
° 0 430 449 130 190
Horsfieldia
T sect. Horsfieldia
TI sect. Irya
II sect. Pyrrhosa
I
2 ete,
* J
ROR ke KE
woe irya
» xxxxxH.crassifolia
| Fig.2. Distribution of Horsfieldia
. L sect, Horsfieldia (1 species). — II. sect. Irya, 40 species, distributed east of Wallace’s Line as
drawn by broken line; distribution of H. irya as stippled line — III. sect. Pyrrhosa, 59 species,
distributed west of Wallace’s Line. Crosses indicate the approximate locatlities or areas of
species with valve-number of perianth straying from the sectons. For further explanation see
the text.
137
138
Gard. Bull. Sing 37(2)(1984)
Keys to the Species
Seven keys are presented, a general key (1) to all species of Horsfieldia, based
on male flowering specimens, and six regional keys (2-7) for female flowering and
fruiting specimens, based partly on vegetative characters and partly on distribution.
la.
2a.
3a.
4a.
(1) GENERAL KEY TO THE SPECIES
based on male flowering specimens
Male flowers sessile, packed into dense subglobose capitula; perianths +
obconical, in bud mutually appressed and angular. Male perianth 3-valved,
at anthesis cleft to c. 1/5-2/3. Androecium stalked, anthers 3-5. Leaves
papillose beneath. Ceylon, elsewhere cultivated................. H. iryaghedhi’
Male flowers subsessile or usually pedicelled; perianths various, in bud not
or only somewhat angular, mutually free or at least not densely clustered
(perianth densely clustered before anthesis in H. sylvestris from E.
Malesia) Leaves not papillose beneath. 7.2.:....2.25-2.0e- ese eee eee 2
Phyllotaxis of plagiotropic (fertile) twigs distichous. Leaves membranous,
usually with largish whitish marks of irregular shape and size. Male
perianth 2-valved, globose, c. 1-1.5 (-2.0) mm diam. Androecium not or
hardly laterally compressed; androphore distinct, tapering, broad above
and hollowed to a saucer-shape, shallowly to deeply; anthers 6-10, their
apices free for only c. 0.3 mm and + in-curved. Ceylon to Solomon Isls.,
generally not too far from the COGSL .....skacsas- -ueees eee. cack eee eee H. irya®
Phyllotaxis of plagiotropic twigs either distichous (alternate), or dispersed
(spirally), or mixed in the same specimen. Leaf consistency various,
usually, without whitish marks. Male perianth 2-4 valved, various in shape
and size. Androecium various, but if the hollow deeply cup- or saucer-
shaped then the anther-apices either mutually free for at least 1/2-way, or
the anthers strongly inflexed into the cup (anthers connate or mutually free,
numberof anthers few to mamny)'::.....2c..-.cseseadee tose cas ceese sees ete eater ane 3
Perianths in any one inflorescence either 2- and 3-valved evenly mixed, or
all the flowers with perianths either 2-, or 3-, or 4-valved. Phyllotaxis of
plagiotropic (fertile) twigs at least partly dispersed (1/3-2/5 spiral), except in
H. longiflora (Annam) with leaves distichous (always?). Leaves without
brown or blackish dots of non-traumatic origin (lens!) beneath. Dried twigs
dark brown, not pale and not much contrasting with the colour of the
petioles. Continental SE. Asia, Andaman Isls., not S. of the Isthmus of Kra
Peete ee eee eee eee eH HEHEHE EE EEE EEE HEHEHE EEE EEE EEE EEE HH EEEEEEE EEE EEEEH HEHEHE EE EE EE EERE
Perianths either 2-, or 3-, or 4-valved, the valve number in flowers of any
One specimen constant or almost so; sometimes a few flowers excepted.
Phyllotaxis distichous or dispersed. Leaves with or without dark dots
beneath. Dried twigs straw or grey (pale) to dark brown, much contrasting
with the dark colour of the dried petioles or not. Peninsular Thailand
through Malesia to Solomon Isls. and N. Australia ..........0ccccecceecneeeenes 7
Perianths (2- or) 3- or 4-valved, pubescent, in male c. 3.0-4.0 mm diam.
H. kingii’
Perianths either 2-, or 3-valved, or 2 and 3 valves mixed, outside glabrous,
in male‘c., 170-3 O0' mm 1ONe...--ssccccomecceeee tee ceossere tree accent eas 5
New account of Horsfieldia 139
Sa.
6a.
Ta.
8a.
9a.
10a.
ia.
Phyllotaxis of plagiotropic shoots (fertile) distichous (always?). Male
perianth + elongate-ellipsoid, 2.2-3.0 mm long. Androecium ellipsoid, c.
PEMAMIMITNOMEDT OA TIENIIN eS ecco. as hae sence ck choca haps. dened. H. longiflora’
Phyllotaxis, usually at least partly, dispersed. Male perianth shorter, c.
1.0-2.3 mm long. Androecium depressed-globose to obovoid, rarely ellip-
soid, c. 0.6-2.0 mm long
Male perianth subglobose or depressed-globose, 1.0-1.5 (-1.7) mm long.
Anthers 7-10. Thailand, Laos, Cambodia, Vietnam: Annam, Cochin-China
5009 JOLIE CES SOR CAEOAGSORS CREE Ses OORRON SEER BCC ECOEE enc SE ESS nee ene He aa tne H. thorelii’
Male perianth short-ellipsoid, or subglobose, or obovoid, c. 1.5-2.3 mm
long. Anthers 8-15. S. China, Indo-China (excl. Annam, Cochin-China)
Ni a1OvAssam, Andaman Isls. (2 vars.) <.cccc.-.ccccscosces+ee H. amygdalina’
Perianths predominantly 2-valved, or sometimes a few flowers in an in-
florescence 3- or 4-valved. Species mainly from E. of Wallace’s line, or the
following from W. Malesia: H. crassifolia (Malaya, Singapore, Sumatra,
Borneo), H. penangiana, p.p. (Sumatra), H. sterilis (Borneo), H. sucosa
REPS COM HOLMEI ye sma. 383 RT EB Ce A, wecsoubecdcawee 8
Perianths predominantly 3-valved, or predominantly 4-valved, or both 3
and 4 valves present; sometimes the odd flower in an inflorescence has
a 2-valved perianth. Species from W. Malesia and penins. Thailand, and
the following from New Guinea: H. angularis, H. olens, H. sepikensis
Nt oa cla ieeinss Vo uolae tsa saeinauidoansoene nae eceaets seaamea 48
Androecium not laterally compressed, in transverse section + circular;
Mover aistinctly longer thami wide. W. MGleSIG <......ccc.sss+-cccecoecccce+cceecs: 9
Androecium laterally compressed or not; if not or only slightly then the an-
droecium including androphore longer than wide; androecium not or little
laterally compressed in occasional 3- or 4-valved flowers. E. Malesia:
MMnCEHS TO SOLOMON ISIS., IN. AUSUGIIG .ouccsocvscsacesecessinasseneenedicean sad 72
Leaves coriaceous, with inconspicuous, subpersistent, dense tomentum of
stellate-scaly hairs with scattered emergents beneath (in specimens of
Borneo the hairs often deciduous, leaving distinct scars); leaves with scat-
tered brown or blackish dots or streaks (lens!) beneath. Colour of dried
twigs and of petioles not much contrasting. Peat swamp forest, swampy
I REAOTUSIVIELYISONU 345, 505088 chiess ocunebenceeessonttwoesoemansoie: H. crassifolia®’
Leaves + membranous, glabrous or early glabrescent beneath. Not from
FOTIA LTT Ia) ONES lam ercran cca nacaeuins teas ssi ale os (anise ciscin cn os eistee atiare sales wamptuacmaene 10
Twigs slender, towards apex c. 2 mm diam.; bark not drying to a pale
colour. Leaves small, c. 7-12 cm long, with regularly scattered dark dots
beneath; lateral nerves flat and inconspicuous above ........ H. penangiana”’
Twigs towards apex c. 2-5 (-10) mm diam.; bark usually drying to a pale
colour and contrasting with the blackish dried petioles or not. Leaves 14 cm
MBC MOLE MOL COLLEGE, DENGAN) 0)..540 .sisiesactecsscrdsssecveesseceaccSdoodausiet 11
Androecium depressed-globose, largely consisting of anthers; central-apical
Cavilvatlateanarshallowseseececccccnece cee see aeteaceeace as H. sucosa subsp. bifissa*
Androecium broadly obovoid, consisting of a large sterile base, and an
upper half bearing 3 (or 6?) inconspicuous anthers ................. H. sterilis”
140
13a.
14a.
1Sa.
16a.
liar
18a.
19a.
Gard. Bull. Sing 37(2)(1984)
Male inflorescence spike-like, i.e., either not so or but slightly so and short-
ramified, lateral branches up to 5 mm long. Inflorescences, flowers and
petioles drying blackish, usually contrasting with the paler grey-brown dried
twigs. Anthers inflexed inwards. MOIUCCGS ...........0cecececeseceees H. spicata’
Male inflorescence usually ramified, the side branches at least 5 mm long.
Inflorescences, flowers, and petioles drying brownish or blackish, usually
not contrasting withithe colour of the/dnied (wigs .:.......-csse.sse-ee eee 13
Androecium + cup-, or bowl-, or saucer-shaped, moderately laterally com-
pressed; the anthers at one or both sides of the androecium distally distinct-
ly incurved or inflexed into -the cup-shaped central cavity. E. Malesia:
Philippines, Celebes, Moluccas, NW. New Guinea, Palau Isls............... 14
Androecium laterally flattened or not, the central column either (1) solid, or
at apex, (2) broadly but shallowly hollowed only up toc. 1/3, or (3) narrow-
ly channelled like a slit to various depths; the anthers straight or only slight-
ly in curved, never inflexed into the cup-shaped central cavity. E. Malesia:
Moluccas to:Solomon Ists:; N= AUSUGNIG 5. c.000c<csessncs nace eee eee 22
Leaves below with regularly spaced reddish-brown to brown-black dots
presumed to be non-traumatic cork warts originating from hair-scars
(lens!), not to be confused with a much finer punctation if also present.
Twigs angular or ridged especially in the upper portion ............ H. inflexa®
Leaves without dots. Twigs either terete, or angular, or winged ............ 15
Male perianth together with the pedicel + pear-shaped, the pedicel taper-
ing. Petiole relatively long, c. 1.0-2.6 cm, leaf blade (6-) 8-22 (-25) cm.
Twigs at apex generally terete, not angular .................... H. moluccana’
Perianth (in laterally view, on the broad side) either short-cuneate, or
rounded, or subtruncate at base; the pedicel not tapered, + abruptly pass-
ing into the perianth. Petioles generally shorter. Twigs terete or angular 16
Anthers entirely connate; androphore short or absent. Male perianth at an-
thesis cleft'to'c. I/2-way <2. 4...cccssocsosenenewion sa zectcen cbr sen dee tee et tee aaa 17
Anthers or thecae mutually free at least in the incurved or inflexed portion.
Male perianth at.anthesis clefitoic.:2/3:0r Over asus: .ce-- ceveceee sieeeeneeeeeee 18
Anthers 18-25, usually inflexed at both sides of the laterally compressed an-
droecium into the thin-walled androecium-cup. Male perianth 2.5-4mm
wide. Leaves membranous, drying matt. Moluccas ............ H. parviflora’
Anthers 11-12, usually inflexed at only one side of the androecium-cup; the
cup thick- and firm-walled. Male perianth c. 2-2.2 mm wide. Leaves char-
taceous, + glossy above. Philippines: Luzon .............++ H. obscurinervia’’
Anthers mutually free only in the inflexed distal portions, the basal non-
inflexed portions connate intoacup-shaped androecium. Androphore ©
minute, only c: 1/10 of the androecium-lengtlt, . cs. -cs-e-ce- eee eee eee emer 19) 9
Anthers free for at least 2/3. Androphore relatively large, its length c. 1/3 |
of the androecium-lempthyicey tae. tad een eas aes as Cale ee 21
Twigs angular or winged. Male perianth c. 4 mm wide, glabrous. Philip-
pines 2 Foe ee © Ue ee ere Cee eee eee eee H. ardisiifolia’?
|
|
Twigs terete, neither ridged nor winged. Male perianth 2.5-3 mm wide ... 20 I
|
New account of Horsfieldia 141
20a.
Bla.
22a.
23a.
24a.
25a.
em
26a.
27a.
Pedicel of male flowers pubescent, shorter than the perianth. In-
florescences rather densely finely pubescent. Anthers inflexed at both sides
of the laterally compressed androecium. Talaud Isls ......... H. talaudensis”’
Pedicels of male flowers glabrous, longer than the perianth. Inflorescences
sparingly pubescent with hairs less than 0.1 mm long. Anthers inflexed only
at one side into the androecium-cup. Philippines: Samar Isls. .............000++
Tha BE ce totes apd Sele ce sewstonasnaeliveseeusindscheencts H. samarensis’*
Twigs generally angular or ridged. Male perianth c. 3-4 mm wide; pedicel
glabrous. Anthers all inflexed into the centre of the androecium. Moluccas
rE eee ee me at ALES Uk poor re H. smithii’®
Twigs terete, usually lined. Male perianth c. 2-2.5(-3) mm wide; pedicel
pubescent or glabrescent. Anthers deeply inflexed at one side of the
androecium, always partly covered over by those of the other side. Palau
I Ee I Ee cncce tonsa conccreencecais H. palauensis ’°
Male perianths angular before anthesis, arranged into dense semi-globose
clusters. Androecium much longer than wide, not or but slightly laterally
compressed. Leaves lanceolate to lanceolate-linear, usually + parallel-
Ee ne 2 Rises so upasimepeNcwenede H. sylvestris’?
Flowers not densely clustered, not angular in mature buds .................. 23
Mature male perianth either + lengthwise ellipsoid, or subglobose, c. 2-3
mm long, very thinly pubescent; at anthesis cleft to c. 1/2-way. Androecium
only slightly laterally compressed, the central column broad, broadly
hollow in the apical portion, but the central column protruding centrally in-
aN SICAMALY HAMUSITONAN S250 eS RE H. australiana”’
Male perianth various. Central column of androecium not protruding. E.
7 EET ES Bee ss Soe OM ae Sales i a ARS OO EAE Parr tek 2 a eee os 24
Twigs sharply angular, or markedly ridged, or winged from petiole to
petiole at the apex of the twig as well as lower down in between the older
leaves and in older wood; i.e., twigs not merely more or less distinctly lined
Nr eter es ih ce Phe Mech Bins co 5 ccc s gdicapldca'awesesscntadasapnceacce 25
Twigs not winged, i.e., terete or only somewhat angular at apex, lower
down terete or merely provided with lines from petiole to petiole (the bark
TORRES OF GOD) boas see ae te Rae 8 Scenes eos aos Sees 27
Male perianth either 2-, or 3-, or 4-valved, subspherical, hardly or not
laterally compressed, drying + glossy, not collapsing on drying.
New Guinea: Vogelkop Penns. ........0.sccccccceeceseceessceeeoes H. angularis”’
Male perianth predominantly 2-valved, little or much laterally compressed,
drying matt, slightly or strongly collapsing on drying .............-.2:+ss00+ 26
Leaves thinly coriaceous. Pedicels of male flowers about as long as or
longer than the perianth. Perianth at anthesis cleft almost to the base. Hairs
of inflorescence and pedicels c. 0.2-0.3 mm. Anthers 10-14; the central
column of androecium at apex hollow for c. 1/4 ...........eeeeeee ee H. iriana’’
Leaves membranous. Pedicel shorter than the perianth. Perianth at anthesis
cleft to c. 2/3-3/4. Inflorescences and pedicels almost glabrous, with hairs of
c. 0.1 mm or less. Anthers (12-) 14-18; the central column of androecium
PRAMEIATTARAIUIS SI. a= 52 ccna cnc tenn nn-vcsescscssotasansessnccesxeeaens H. aruana”’
Male inflorescence large, 25-35 cm long. Male perianth + pear-shaped.
Androecium longer than broad; androphore c. 0.5 mm long or more, about
142
28a.
29a.
30a.
Biliak
32a.
33a.
34a.
Gard. Bull. Sing 37(2)(1984)
half astong’as the anthers orlongery.. Bt ctsc cna. deeneete se cee eet en ean eaeeeee 28
Male inflorescences c. 20cm long or less. Male perianth of various shapes.
Androecium longer or shorter than broad; androphore short or long
Assi elds RE QA sh eee et EAS eae ected: ee 29
Male perianth glabrous (?), c. 4 x 2mm. Anthers 10, androphore nearly as
long as the anthers. Inflorescence glabrescent, to 25 cm long .... H. ampla”4
Male perianth pubescent, c. 3 X 3 mm. Anthers 7, androphore about half
as long as the anthers. Inflorescence pubescent, c. 25-35 cm long ..............
BOT ee oe BEG aarinnade doannoncn db ocosnetbcnaantedsoaAbboobosonon H. ampliformis”
Male inflorescence delicate, 2-5(-8) cm long, 1 or 2 (or 3) times ramified.
NewiGuineaninel VArivlsis.c.. 22ers eee Eee 30
Male inflorescences generally stouter, c.5-20cm long, not or 1-4 times
ramified (inflorescences of H. sinclairii and H. basifissa from New Guinea
sometimes small). Whole of E. Malesia, incl. Celebes, Pacific; not in the
PRU DP iM eS.vx catiriae dzeeieian hem seach eee eb ace nos bee case a eee 34
Male perianth pubescent or late glabrescent, distinctly or only somewhat
longer than broad. Anther-bearing part of the androecium much shorter
than the elongate club-shapediandrophoreisx. 7y.cce te eeee cee 31
Male perianth glabrous or early glabrescent, about as long as or shorter
than broad. Androphore much shorter than the anthers ...................... 33
Leaves broadly obovate or elliptic to oblong, c. 12-17 x 5-11 cm. Male
perianth gradually passing into the thick and tapered pedicel, both together
c. 10-12 mm long. Androphore glabrous ................... H. crux-melitensis7’
Leaves elliptic to lanceolate, 4.5-20 « 0.7-6 cm. Male perianth with pedicel
A-6(=9). MM csdscnasateeeaads Lack Oe DW aleee ae. bee dees en ae eee 32
Leaves drying olivaceous. Male perianth with pedicel together club-shaped,
65-5256 2-2-2)mm Androphoreiglabrousmeese setae teeta eee H. clavata”’
Leaves drying dark brown. Male perianth broadly ellipsoid, c. 2.0-2.4 x
1.8-2.2 (-2.4) mm, at base narrowed into a slender pedicel 2-3.5(-6) mm.
The surface of the androphore in the upper part wrinkled-bullate, towards
the base either densely, minutely scaly-hairy, or striate but glabrous ..........
gis e-o.ainin define layoTe ae falesic tea ols IAS UIE rater ote eee H. squamulosa”’
Male perianth + laterally compressed, generally + obtriangular in lateral
view, 1.8-3 mm wide, usually + collapsing on drying. Androphore slender,
c. 0.2.-0.5 mm long, usually much shorter than the anthers ..... H. subtilis?’
Male perianth generally subglobose, 1-2 mm wide, not or but slightly com-
pressed, the surface wrinkled on drying but not collapsing. Androphore
0.4.-0.5 mm long, about half the length of anthers ............ H. schlechteri*’
Male perianths together with the pedicels generally + pear-shaped; the
upper part of the perianth broadly rounded in lateral view, the lower (1/4-)
1/3-1/2 tapered and gradually passing into the + tapered pedicel (these
characters not always clear in certain specimens of H. tuberculata).
Perianth glabrous, pubescent or glabrescent ............-0.0.csesssecsessesanee 35
Male perianth in lateral view of various shapes, either circular, or ovate,
or obovate, or elliptic, or transversely elliptic, or reniform; at base either
New account of Horsfieldia 143
37a.
38a.
39a.
4la.
short-attenuate, or rounded, or truncate; not tapered, the pedicels generally
not thickened towards the perianth. Perianth either hairy, at least at the
base, or in H. psilantha, H. basifissa, H. sinclairii glabrous, subglabrous,
OP Sees RAN ess Rent eee ee 2 39
Perianth glabrous. A variable species ..............0ceceeeeeeees H. tuberculata*’
Perianth minutely pubescent, or early glabrescent in H. corrugata ......... 36
Leaves small, + lanceolate, 5-16 cm long. Male perianth at anthesis cleft to
CUD? STAG C2 27S Ee A ee H. lancifolia*’
Leaves elliptic to lanceolate, generally much larger, 12-30 cm long. Male
Perini atanthesis cleft (©. C40/22Way goss. <td vcke 1 dbeccostdedadoeRedendsalieces 37
Pedicel c. 1.5-2 mm long; perianth c. 2.3 mm long, thinly pubescent. An-
PETS C56) MOIUCCHS,. AE LOW QUILUGE c5...00 0500s c0ecsseseseiesdaermee H decalvata**
Pedicel generally longer; perianth c. 2.5-3.5 mm long. New Guinea, at c.
450-2000 m alt. Vegetatively and sometimes in fruit much resembling
Lat, WE ILGTIGE? oc cocndaane SOC ISECOE CBO O TRCN COON SE TOA CBRE A EE PARTE eE See Annee 38
Male pedicel stoutish, 2-4 mm long. Male perianth + membranous,
glabrescent, with or without a few scattered blackish warts. Anthers c. 8-12
sos ose tounegpadodoopbeBERpoasoSBOB SEO SH ORC an PCa ES na Hae a annaes Saran ax H. corrugata”’
Male pedicel stout or rather slender, 2-5 mm long. Male perianth + fleshy,
pubescent, without blackish warts. Anthers 5-10 ............. H. pachycarpa®’
Inflorescence moderately pubescent to nearly glabrous. Tomentum on leaf
buds and apical portions of twigs short, rusty or greyish, the hairs
0.1-0.4(-0.5) mm long. Leaf largely or entirely (early) glabrescent, or with
scattered stellate hairs beneath especially the younger ones .................. 40
Inflorescence generally thick-woolly tomentose. Tomentum on leaf bud
and apical portion of twig usually rust-coloured, conspicuous, its hairs
coarse and long, (0.3-) 0.5-1.5 mm. Leaves with (sub) persistent tomentum,
areas’ Onalid near the midrib beneath .....:-...00502c.0.seosccessoorceenmmensatesee 45
Male perianth largely glabrous, subglobose, 2-3 mm diam., at anthesis cleft
FOE DaSe NOL GCOMAPSING. ON ALYING oo. oa. se sjers sweeter opision saneeens H. basifissa*’
Male perianth glabrous or hairy, of various sizes and shapes, at anthesis
cleft to c. 1/2-way to near the base; perianth collapsing on drying or not 41
Male perianth wholly or almost wholly clothed with + persistent
tomentum, the hairs may be scattered and very minute. Leaves drying
olivaceous to brown, without a reddish tinge; marginal nerve not con-
spicuously regularly looped. Hairs of leaf bud c. 0.1-0.2 mm long, usually
greyish. Moluccas, New Guinea, New Britain; not in the Solomon Isls. 42
Male perianth either glabrous, or early or late glabrescent, at least the upper
ee INCH MINED TO WOIOIMON IS)... 20's oka. vnesatcenennesacoemees ae team ores veh 43
Mature male perianth small, c. 1.2-1.9 mm diam. A variable species ..........
EOE vee ten eet ee er de Ere! borat busts saute Pholooe epee qahie « pamecele H. pilifera”’
Mature male perianth 2.0-3.3 mm diam. A variable species ... H. laevigata”
Perianths and pedicels glabrous; male perianth c. (20-) 2.5-3.5 (-4.0) mm
diam. Leaves rather large, 20-40 cm long, drying olivaceous to brown, the
144
45a.
46a.
47a.
48a.
49a.
50a.
Gard. Bull. Sing 37(2)(1984)
marginal nerve rather indistinct and not very regularly looped. Bagabag
Isis: Longisl:, New Britain, New lreland ee H. psilantha’?
Perianths and pedicels glabrous or pubescent; male perianth (1.0-) 1.5-2.0
(2.5 mm diam., see note 5 under H. sinclairii). Leaves drying generally with
a reddish tinge, especially the midrib and nerves ................seeeeeeeeeeeees 44
Leaves usually large, 9-30 (-40) cm long, the marginal nerve very distinct
and conspicuously regularly looped. Hairs on leaf bud 0.1-0.4 mm, average
c. 0.3 mm long, usually reddish brown, sometimes greyish. Pedicel and base
of perianth minutely pubescent. Male perianth c. 1.5-2.0 mm diam., cleft at
anthesis: tore 9/10 SSOlornOnmUds | iresn its) tens esac ee eee H. whitmorei*’
Leaves smaller, 6-20 cm long, the marginal nerve indistinct and not very
regularly looped. Hairs of leaf bud to c. 0.1 mm long, greyish brown.
Pedicel and perianth glabrous (but see the notes). Male perianth c. 1.1-2.0
mm diam., at anthesis cleft to c. 1/2-way; perianth larger and pedicel pubes-
cent in deviating specimens as discussed in note 5. E. New Guinea .............
Soares waainsteare esa icteaclsoxas Sila tattle Sed oid elaine breil reset ae eet ee ee H. sinclairii*?
Perianth in the male largely pubescent, towards the base thick-walled and
coriaceous, the remainder collapsing on drying; both ¢ and @ opening at
apex by small pore-like slit less than 1 mm long. Androecium subellipsoid,
mainly consisting of the column with 2 minute anthers situated at the top,
just below the pore. Leaves usually coriaceous, + bullate; clothed with
harsh hairs which when shed leave rough thickened bases . H. pulverulenta”
Perianth glabrous or pubescent, membranous or chartaceous, not much
collapsing on drying, opening by a slit to at least c. 1/3. Androecium mainly
consisting of 10-16 sessile anthers. Hairs on lower leaf surface not harsh,
when shed not leaving rough thickened bases. ........5 00+ «s«» «cues seeeebesseneeiene 46
Perianth pubescent, in male at anthesis cleft to c. 3/4-5/6. Anthers 10-14.
Leaves chartaceousor membranousmc:sses-sesastetestres seer eaee H. leptantha®
Perianth glabrous, except at the very base; at anthesis cleft to c. 1/2-way or
less... Leaves generally, membranous ys te-pone ds ac ceceey en nas te 47
Male perianth subglobose. Anthers 12-16. Leaves generally oblong to
oblong-lanceolate, at apex not-caudate (always?) ................ H. hellwigii*
Male perianth obovoid or ellipsoid. Anthers c. 10(-12). Leaves generally
oblong-lanceolateatiapexicallGalieiee-sseeseeeenreser eee eeeeseEaeee H. ralunensis*
Phyllotaxis in plagiotropic shoots (i.e. usually the fertile) dispersed,
i.e., leaves in 3 or more rows along the twigs. Terminal leaf bud generally
rather short-and, DrOad) «tec cmctc ccs teehee weet mae eases cee eee 49
Leaves in plagiotropic shoots distichous (alternate), in rare cases, a few on
the same plant in 3 rows. Terminal leaf bud generally more slender ........ 57
Leaves bunched towards the end of the twigs. Leaf bud and inflorescence
pubescent with hairs c. 0.5-1.0 mm long. Bark of older twigs often blackish
and flaking. BOrneOs.. -\ cccassosancnccs cies teouncaecteer trees H. sabulosa”
Leaves bunched or not. Tomentum on leaf bud and inflorescence short,
the hairs up to c. 0.2 mm long, or, if plant from Continental Asia, then
hairs short or long, up toc. 1 mm. Bark of older twigs flaking slightly or not
dss bbdg asi. od Sluis ba vain lbh ONE voces SR ECR ERC Reon aOR NS eee UNG Case NeA CAE Cr eee ee aeentCr 50
(VEST) ici acsiers oiscsis ee sie’ ravardevc Svovs jars o alate lelelefelelsaie sevsva ser electsia state estes oticielete tate Teceeeieetonne 51
New account of Horsfieldia 145
b.
Sla.
524°
59a.
WowerleatsuriaceswithoOutidots) ius: He. ees OR eee 52
Bark of twigs generally drying brown or grey-brown, not contrasting with
the dark colour of the dried petioles; older bark not flaking. Leaves in 2 or 3
LOSE oP tids Sect 12 ee See ee ee ee re ae ene te H. glabra’”
Bark rather pale, grey-brown to pale yellowish-brown, rather contrasting
with the petioles; older bark + flaking. Leaves in 3-5 rows. Aceh
TEES AC NOU TI oo a2 2c d5 sai oncad waja au niceapasawaseasiese ses H. atjehensis*’
Bark of twigs drying brown, not contrasting with the colour of the dried
(OL OUBE Se scne op ce oct OER EE EE Dee AE Es re ae ee on eee es ee 53
Bark of twigs drying pale, greyish-white or straw, contrasting rather well
with the blackish brown colour of the dried petioles ....................0c0000s 55
Twigs distinctly ridged or short-winged. Sumatra ............... H. hirtiflora”’
Meter rerere, NeUNel TGDEd NOL WINGE 6c. ..6o5 5 vie dosieecumaaaesicncs sceneasceceds 54
Leaf pubescent beneath. Sumatra, Malaya ..............c0cceceeees H. superba”
Beam PlADLOUS DENEAU: BOICO. ........cc0c.c0vcescenccsosesecweosne H. fragillima®
Male pedicles at base articulated. Androecium much depressed, with the
apical cavity conspicuous, broad, either shallow or rather deep, reaching up
to nearly 1/2-way the androecium and bottom flattish; androphore narrow,
largely hidden by the anthers. The leaves in most specimens distichous,
IE PIMCS 4 -SLICHOUS)- VVIGIGYG, SUMAGUTG o... 002000000. c0ceedeanoesss secs scsvedsos
Male pedicels all or mostly not articulated. Androecium not or but slightly
depressed, the apical cavity usually narrow and inconspicuous. Leaves
GTA (TL SES) TOMS pea birt cose Soo c ROO RE CSE O une OUe na nedc anaes aes cceoaanen eee 56
Androphore indistinct or absent. Leaves generally drying blackish-brown.
PSR COM Pe EERIE EN, Ua tersichs bs agistes odeawle cos unten ncsedeuwes H. pallidicaula*’
Androphore relatively distinct, 0.3-0.4 mm long. Leaves generally drying
bright brown. Sumatra, Malaya, penins. Thailand ................. H. sparsa™’
Male perianth + obconical-obovoid, very leathery, at anthesis opening for
1/6-1/5 only. Androecium turbinate, anthers 3. Sumatra ....... H. triandra*’
Male perianth of various shapes, opening for c. 1/5 or more. Androecium
WAALS MATIUNELSY4 Ole MMOLE }. jase ntet oka bees Nee aacune cee ee oa ate ecb ana ee ote cue ee 58
Mature male perianth in bud ellipsoid or obovoid, 3-8 mm long, at anthesis
splitting c. 1/5-1/4 (-1/3). Leaves usually + parchment-like, on drying matt
above owing to the finely wrinkled surface (in H. sessilifolia male flowers
notknown and leaves not'so distinctly matt)... ....:....2.....c-scesesccscesenes 59
Mature male perianth in bud either (1) ellipsoid or short-pear-shaped, c. 3.5
mm long or less (though sometimes rather large in H. endertii, H. floc-
culosa, H. majuscula, H. wallichii), or (2) perianth somewhat globose or
depressed globose; perianth at anthesis cleft to (1/4-) 1/3-1/2, or more.
Texture of leaves various including coriaceous, but never parchment-like
ana leaves nObiypically, Matt.aAbOVe cad duas ates sascid tees Jemeslaceaecplee Bobs ececs 62
Plant stout: twigs towards apex 5-7 (-10) mm diam.; leaves 25-70 cm long,
tomentum: present. and persistent beneath 2552.2... cac cote tic ceect ok ewseats 61
146
6la.
62a.
66a.
Gard. Bull. Sing 37(2)(1984)
Twigs towards apex 3-5 mm diam. Leaves up to 30 cm long, (largely)
glabrous beneath, or with some hairs persisting on the midrib. Male
perianth 3.0-5.0.mm long... ain.s eh eee eee ee eee 60
Leaves drying dark olivaceous; up to 32 cm long. Bark of twigs bright
brown or yellowish-brown, coarsely striate and tending to crack
longitudinally. Inflorescence (almost) glabrous. Pedicel not articulated.
Anthers: 1'2220)33.oec ee ee eee Tee hs Pee HH. tristis‘?
Leaves drying fulvous-brown; up to 24 cm long. Bark of twigs brown, finely
striate, not cracking. Inflorescence pubescent. Pedicel articulated at base.
Anthers 110-12)... secusaspeseameusaasmmesanoenaneri acne Cee eere een eee Ree H. fulva®’
Male perianth 7-8 mm long, perianth and pedicel glabrous. Petiole 6-15 mm
long: Malaya :SUmaira serio cco ee eee ee ee H. superba’
Male flower not known; female perianth and pedicel pubescent. Petiole
almost:absent. Borneo (Sardwahe) v.25... eecneeae eee ene H. sessilifolia*’
Leaves with a persistent or subpersistent tomentum below, in H. gracilis
and H. wallichii the tomentum sometimes vestigial on and near midrib and
REEVES, cciss.c.s1e neediness oletnoree sister oesiciene eelsse sec ec meer ence EEC CR eee eRe Recetas 63
Leaves either glabrous or early glabrescent beneath, or in some species often
with a vestigial pubescence on the lower midrib ................cccccececeeeeees 74
Leaf venation reticulate or not, not scabrous above ..............0cceceeeeeees 64
Venation distinctly reticulate on both surfaces, older leaves scabrous above
ob b0.d alja'ese GeGieweiciaieis's oie s'sie SS wes sae See ae Croce OO REN S oe Icne eIc eee ee H. grandis”
Leaves with scattered dark brown or blackish dots and/or streaks beneath,
obscured by hairs Or mot-seeter erase es a a eh ee 65
Leaves without dots or streaks beneath) .:..-s2..0..ceaseasneeeeeeeeee eee 66
Mature male perianth in bud short-pear-shaped, c. 2-2.5 mm long; pedicel
indistinct, c. 03-1.0 mm long. Leaves rather thinly pubescent, often +
glabrescent beneathye 05 Ae eee H. wallichii*’
Mature male perianth subglobose, c. 1-1.5 mm diam., pedicel distinct, c.
1-1.5 mm long. Leaves with persistent, conspicuous tomentum beneath
SERRE CoC REO ennionondedobaadec osoaccandnscsboataososassonocecadoaaaouboc H. pulcherrima”’
Mature male perianth in bud broadly ellipsoid or obovoid, 2-3 mm long;
androecium longer than broad. Pedicel (1.5-) 3-4 mm long. Tomentum on
twigs and lower leaf surface with hairs c. 1.5-2 mm long..... H. flocculosa*’
Mature male perianth subglobose or depressed globose, c. 2.5 mm diam. or
less; androecium as broad as or broader than long. Pedicel 3 mm long or
less. Tomentum on twigs and lower leaf surface with hairs to c. 1.5 mm long
saves auheose se tineay aalcotbes ta taeeltetes ae tede somes toe tenes te etene natn ont ete ete aaa 67
Male perianth to c. 1 mm diam., with persistent tomentum or late glabres-
r=) 1 eee em ANE ERR FSET een odadecedooonaddbabaT[c008 H. motleyi®
Male perianth c. 1.0-2.5 mm diam., glabrous or early glabrescent ......... 68
Male pedicel at base not articulated (character not quite clear in H. rufo-
lanata; male flower not seen in H. QracillS) ...........ececeesenencenencenseeeese 69
Male pedicel with distinct articulation at base .............s:eseeeeseeeeee reese 13
New account of Horsfieldia 147
69a.
70a.
Tla.
72a.
73a.
74a.
75a.
76a.
via.
Upper leaf surface drying dark brown, with the venation (reticulation)
usually indistinct. Malaya, S. penins. Thailand ................. H. tomentosa®
Leaves drying olivaceous to brown, with the reticulation on upper surface
ETE Ge OEE ae Pe 70
Twigs stoutish, diam. towards the apex 3.5-7 (-13) mm. Leaves 10-45 cm
long. Tertiary venation usually distinct above. Lateral nerves c. 11 pairs or
more. Male perianth c. 1.5-2.3 mm diam.; anthers 8 or more. Plant grow-
ing generally on soils richer than kerangas Or SANA ...........cccecceceeceseeee 72
Twigs slender, diam. towards the apex up to c. 3 mm. Leaves small, c. 7-21
cm long. Tertiary venation usually indistinct above ....................eeceeee 71
Twigs 1.5-2.5 mm diam. Leaves thinly membranous, 12-21 cm long; lateral
nerves 14-17 pairs. Lower leaf surface with tomentum rather sparse, subper-
sistent, the hairs vestigial mainly on nerves and midrib. Male flower not
MSMEL TINIE Y TOWING JOTESL &o0csc0cc 2 co secc osces cde sees cece cessecveecc H. gracilis”
Twigs c. 2-3 mm diam. Leaves 7-15 cm long, + chartaceous; lateral nerves
5-9 pairs. Lower leaf-surface with denser tomentum. Male perianth c. 1.0
mm diam.; anthers 4 or 5. Kerangas forest, or forest on coastal white sand
EEN ARREST eee ode Soo: Nostay SedaStiisesacssscesbecestece H. paucinervis®
Leaves generally large, 18-45 cm long; lateral nerves 18-25 pairs, sunken in
upper surface. Male perianth c. 1.5-2.0 mm diam., anthers 8-10 ...............
Ee ne Seen canes cen ae coast usinesioabingtee ove Wewssipetasveeses pee soes H. splendida®
Leaves smaller, 10-23 cm long; lateral nerves 11-16 pairs, raised above.
Male perianth c. 2-2.3 mm diam.; anthers c. 15. A montane species at c.
COON 400 alts tt Sarawak; SQDQH 0. .2.....0ss000sse0ccseseoee H. rufo-lanata®
Leaves membranous, not bullate; veins in upper surface flat or sunken,
reticulation rather faint to distinct. Male perianth 1.5-2.2 mm diam.;
pedicel equal to or somewhat longer than the perianth ........... H. affinis”
Leaves chartaceous, bullate; veins sunken, reticulation very distinct. Male
perianth 2-2.5 mm diam.; pedicel as long as or shorter than the perianth
oc oo occ seccoe caGscnae seew loess eseaseemesQemeeaeee H. reticulata®’
Twigs in apical portion pale, grey-whitish or pale straw coloured, con-
trasting with the blackish or dark brown colour of the dried petiole ...... ib)
Twigs in apical portion drying brown to dark grey-brown, not contrasting
with the brown to blackish colour of the dried petiole ...............0.0..... 78
Androecium in transverse section triangular. Anthers mutually free for the
upper half of more. Borneo: heath forest on sand or peat soil .................
re ae wat ecvapes sent ce see aston seeds dsesdieuccens H. oligocarpa’’
Androecium in transverse section circular or subcircular. Anthers largely
SMR err ae -Eeee eee Pat te aasoe nett ede don tendescadsetows asecacsewsanidcdtasenaes 76
Leaves chartaceous, drying bright brown. Pedicel at base not articulate.
Androecium with small apical cavity. Borneo: mostly in heath forest or
Keripas JOrest OM SANGY SOUS ....2.00.0<20.0000s01esne0esseerenssonens H. carnosa®’
Leaves membranous, drying usually + brown or blackish brown. Plant
growing usually in forest on richer soils, incl. SANG. ...........000000cceeceeeee 77
Pedicel articulated at base. Androecium strongly-depressed globose, with
the apical cavity broad and the bottom flattish, cavity nearly 1/2-way deep.
148
78a.
8la.
82a.
83a.
84a.
Gard. Bull. Sing 37(2)(1984)
Malaya Sumatraicessh gece ee ee H. sucosa subsp. sucosa**
Pedicel not articulated at base. Androecium ellipsoid to slightly depressed
globose, the apical cavity small and narrow. Borneo ........ H. pallidicaula”’
Twigs distinctly ridged or nearly winged in between the petiole insertions,
also mthe ‘older wood 7.5. 200 sees ccto eet eh One cone ee ee 79
Twigs not ridged; sometimes twigs faintly ridged, or lined or angular in the
apical portion ONLY vsoseccasiantodensae- tect Sacto tw anl ae ee eas ee ee 82
Male perianth at anthesis cleft nearly to the base. New Guinea ............. 80
Male perianth at anthesis cleft to c. 1/2-2/3. West Malesia ..............0.0 81
Male perianth slightly broader than long, short-pubescent in the lower half.
Androecium slightly broader than long; anthers erect, not incurved .........
uw aebawoepGeecclweenlstlvceiscean nsbidsis/eh nts oh sts oaE RT ERee ACRE raat aaa aeeaeaee H. angularis”°
Male perianth subglobose to broadly ellipsoid, glabrous. Androecium
longer than broad, + obovoid, the anthers with apex free and incurved,
those of one side of the androecium clasping the others ............. H. olens’’
Mature male perianth in bud c. 2.5 mm diam., pubescent. N. Sumatra .......
Pee oe ese REE RRS S ST At Te Bocataceacdbs dadbancuooddaposudenabedoe H. hirtiflora”’
Mature male perianth in bud c. 1.0-1.5 mm diam., glabrous. S. penins.
Thailand, Malaya, Sumatra, BOrneo ..............0ccceeccceeceees H. brachiata”
Mature male perianth in bud short-pear-shaped, c. 2.0 (-2.5) mm long,
subsessile; the pedicel much shorter than the perianth, c. 0.3-1.0 mm long,
thickish. Leaves coriaceous; lateral nerves flat or usually sunken above,
pubescent or glabrescent beneath and with scattered blackish dots and
streaks (lens!). Twigs towards the apex usually conspicuously hollow
safe a's « aso/asiesne's n Ganaastaes se osicter Shane daca hice een Scars Oa See eee H. wallichii*’
Male perianth various in shape and size, the pedicel proportionally longer
and more slender; perianth obovoid with pedicel short in H. glabra var.
oviflora. Leaves various, nerves raised or sunken, dotted or not beneath.
Twigs solid or generally less strikingly hollow ..................sceecereeeeeeeees 83
Male inflorescence very stout, the rachis towards the base 5-8 mm diam.
Androecium about as broad as long, triquetrous, the anthers entirely
connate. WicBOrneo sashes ee ee eee H. pachyrachis”
Male inflorescence large or small, the rachis towards base c. 4.0 (-4.5) mm
thick or less. Androecium of various shape, in transverse section triquetrous
or circular. Whole:Of W.iMGIeSIG viet icssocckeoscasedan haan sc hcapeeh eee eee 84
Androecium in transverse section 3- or 4-angular. Anthers + erect, for a
large part mutually free, usually for c. 1/2-way or more. Lateral nerves
raised above, except in H. ridleyana (with leaves small, and male perianth
c. 1 mm diam.). Leaves not dotted beneath. Male perianth c. 1.5 (-2.0) mm
diam. or less. Pedicel articulated at base. Most of W. Malesia, not in
Celebes, rare.incthe Philippines tithisc. Jcodesa- sieges. Hae dh SR eee 85
Androecium in transverse section either circular or + ellipsoid, or sub-
triangular with rounded angles, neither triquetrous nor quadrangular.
Anthers + curved, almost entirely connate; free apices c. 1/3 or less. Leaf,
with the lateral nerves raised above, or level, or sunken; with or without
blackish dots (lens!) beneath, the dots not to be confused with a much finer
punctation present or not. Male perianth c. (1.3-) 1.5 mm diam. or more.
New account of Horsfieldia 149
85a.
86a.
87a.
88a.
89a.
epee Beas ae lem ateel OROL is . bee reheat bore ty, Jocad toe bs, 92
Leaves small, 5-16 cm long. Lateral nerves + level or sunken above; midrib
PeMrPECHeTAMSEH OF fidt— OF SUTKEN 2... <<... ssocc. feccce cocoon ce coonnecckoee 86
Leaves small or large, 5-28 cm long. Midrib and lateral nerves raised above
otis zieedgpceet 22k ps 2k Seg See ae ee oe oe Se 87
Anthers 4-6. Nerves above usually indistinct or invisible. Leaf apex acute or
acute-acuminate. Twigs at apex and inflorescence rather glabrescent.
III ARN is oe Eh A i et dixn! hoe mSateren deez H. ridleyana~”
Anthers 9 or 10. Nerves flat or but little raised above, well visible. Leaf apex
blunt. Twig apex rather late glabrescent, inflorescence with persistent
ET SEA AOL SRL eee ee a A A ee H. obtusa”
Flower not seen. Leaves early glabrescent, also on the midrib; leaf apex
long acute-acuminate. Bark of twigs rather smooth, lower down cracking
OD LLL LES D cal 2077 ae pO Sree H. disticha”
Leaves early glabrescent, the midrib beneath early or later glabrescent; leaf
apex acute-acuminate. Bark of twigs striate, lower down coarsely striate or
NCR wren ca. re Ret DE 8) NO Se RO IOP 88
Twigs slender, towards apex 1-2 (-4) mm diam. Leaves rather small, 7-18 cm
long, thinly membranous to subchartaceous; petiole slender, diam. 1-1.5
(-2.0) mm. Male inflorescence delicate, up to 9 cm long. Mature male
Semen n ET AI PISEAYEY fe “ER ININEY oe oo oes ce rte tows, ee Ek ON 89
Twigs towards apex 1-5 (-8) mm diam. Leaves of various size, chartaceous
to coriaceous; petiole 1.5-4 (-8) mm diam. Male inflorescence up to
15(-20) cm long. Male perianth 1.0-2.0 mm diam .......................c0e00ee 90
Leaf bud with tomentum composed of hairs c. (0.1-) 0.2 mm long; twigs at
apex and leaves glabrous; inflorescence with rather sparse tomentum of
stellate hairs c. 0.2 mm long, glabrescent. Leaves drying to a greyish
tinge. Mature male perianth in bud rather short-pear-shaped, shortly taper-
SRM METENCUN COM Se ee Loe et eaicn Sa oee Coeuue ees een en ceesek eet H. tenuifolia”
Leaf bud, apical portion of twig, petiole and inflorescence with woolly
tomentum of stellate-dendroid hairs c. (0.2-) 0.5 mm long; leaves with
rather persisting hairs on midrib beneath, drying olivaceous. Mature male
perianth in bud globose or depressed-globose ................... H. macilenta”
Plant robust, twigs stout, leaves coarse, the midrib broad above, at the tran-
sition to the petiole at least 3 mm wide. Male inflorescence 10-20 cm long.
Borneo: forests on poor soil, including sand and peat ........ H. laticostata”’
Plant generally less robust; leaves various, the midrib above towards the in-
sertion of the petiole less than 3 mm wide. Male inflorescence up to 15 cm
EROS OF OOP OF TICK SOW 3. «ona oconnasceeacesspsmstasacietecedswanancduesss 91
Leaves 16-28 cm long; nerves 16-19 pairs, very prominent above; leaf base
+ rounded or short-attenuate. Borneo: Sarawak .............604. H. nervosa‘?
Leaves 7-28 cm long, base short- to long-attenuate; nerves 6-16 pairs, raised
to various degree above. On drying, colour of leaves above and below
usually much contrasting, generally more so than in the related species. (A
variable species, especially the fruits.) Malaya, Sumatra, whole of Borneo,
REPAID Sao os seca ne tan ontnnhases qngeecss H. polyspherula®’
93a.
94a.
9Sa.
96a.
97a.
98a.
99a.
100a.
Gard. Bull. Sing 37(2)(1984)
Leaf bud, apical portion of twig, and inflorescence with tomentum of hairs
c. 0.2 mm long or more; hairs 0.1-0.4 mm long in H. punctata .............. 93
Leaf bud, apical portion of twig, and inflorescence with tomentum of hairs
less than 0.2 mm long; hairs usually c. 0.1 mm long or less ................. 100
Mature male perianth in bud ellipsoid, c. 2.5-3.5 mm long; androecium
longer than broad. Leaves generally drying thick and brittle, tip bluntish.
Lower leaf surface usually with conspicuous pale golden hair scars (lens!) ...
a aia wid'e ws eyoisialoaje%e isle eisiowein sleiotolo elatereraiolete oles ores seats soocemiot eisece eiemenecaeenee H. endertii*
0) (01-10 ERNE RREEe Cees ee ar | ane a aie AER Mon | ane 5 he nu dbdcankonsdostocosaoe 94
Mature male perianth in bud c. 2.5-3.0 mm diam. (or c. 1.5 mm in a
deviating specimen Hallier 624 from W. Borneo, see the notes), at anthesis
Cleft t0:¢5,4/S ASSUMES... Socssvietoctund cowadtaeucnaceees eee ee H. valida‘
Mature male perianth in bud c. (1.0-) 1.2-2.5 mm diam., at anthesis cleft to
1/3-273'(-3/4): ‘Malaya, Borneo: 5.2225. 2e seas haneee ke eee ee 95
Pedicel in male flowers not articulated at the base. Leaves without blackish
dots or streaks below (lens!). Lateral nerves largely raised above. On dry-
ing, colour of upper and lower leaf surface not much contrasting .......... 96
Pedicel at base articulated. Leaves with scattered blackish brown dots and
stripes beneath. Lateral nerves above largely flat or sunken, rarely faintly or
partially raised. Upper leaf surface drying greenish or blackish brown,
usually rather contrasting in colour with the cinnamon- or chocolate-
coloured lower sutfaceisuc?. easter. Ange ete eens eee H. borneensis®*
Male perianth c. 2.0-2.5 mm diam. Androecium sessile, + saucer-shaped
and broadly hollow. Leaves 20-45 cm long. Lowland forest ..............0.0000
Male perianth c. (1.0-) 1.4-2.2 mm diam. Androecium + globose or
depressed-globose, with the apical cavity small, concealed by the overhang-
ing anther tips. Leaves 4-35 cm long. Montane forest at 800-200 m ........ 97
Androecium borne by a distinct slender androphore c. 0.3-0.8 mm long, not
hidden by the anthers. Leaves membranous, 9-18 cm long, drying dark
brown, leaf tip acute-acuminate ...................00-e0ccsseee H. androphora *”
Androecium sessile, with the androphore absent or short, up to 0.5 mm,
largely or completely hidden by the anthers 2s.2..9.2..0<..-s cee te cee eee 98
Leaves chartaceous or membranous, to c. 35 cm long, drying olivaceous to
brown above; leaf tip acute-acuminate. Male inflorescence large, to c. 20
cm long: (Borneo. seces. See Hain Gosia eae H. amplomontana*®
Leaves coriaceous, 4-14 cm long, drying olivaceous brown or blackish
above; leaf tip obtuse to subacute. Male inflorescence 4-16cm long...... 99
Leaves drying dark brown to blackish, lower surface not punctate. Borneo
H. montana’?
Leaves drying olivaceous brown, lower surface densely brown-black
punctate (lens!) Malaya... ..saceueeatesnacconsneseee aoe eee H. punctata”
Leaves dotted or coarsely punctate beneath; dots brown to blackish, or
rarely pale brown, + regularly spaced and equal-sized, of non-traumatic
OTIQIN ..cevscecscenuncaacudnoncansacesceestacseeereeeneteaeet en aueeee ena ae eee ee ee eeee 107
New account of Horsfieldia iL Syl
b.
10la.
102a.
103a.
106a.
107a.
Leaves not dotted beneath (dotting should not be confused with generally
smaller, irregularly spaced, blackish mottles or points, or with dots of
AACS SEACS ONCE AMI ALC OMIPIN)) 5-2. .oc senseccareccesssdeccesedcedassncesactens 101
Male perianth + ellipsoid; androecium + obovoid, the anthers with their
apical portions deeply inflexed into a deep apical cavity. New Guinea ........
es eee ee ee Bers SSS si Siatein xa SH T6Sa6 abv ct sciswe sWievdde sees H. sepikensis’*
Male perianth and androecium of various shapes; the anthers + straight or
curved, at apex not inflexed. W. and E. Malesia, not in New Guinea .... 102
Male perianth coriaceous; valves thick, towards the base (0.3-) 0.4-1.0 mm
thick. Androecium ellipsoid-obovoid, longer than broad ................... 105
Male perianth thinner, valves at base c. 0.2-0.3 mm thick. Androecium
subglobose or broadly ellipsoid or broadly obovoid, not or but little longer
than broad. Androphore narrow, and only c. 0.2 mm long. Leaves mem-
NE TEPID REASEACEONG he ooo. oot Ses sc cctcs lacus ccedessecdceccesciossosesdteass 103
ETE CORE ee Be Re Be SL ich icictalo\s cig nasi oe sasiele tian Gueeeee da geeeees 104
Male pedicel shorter than the perianth, c. 0.5 mm long. Celebes, Philippines
Eee ot ae Tae ease oe siveie wad aeiaaw ens e veeacaecsarsonveveaess H. costulata?’
Male perianth at anthesis cleft to c. 1/2-way ...............eeeee H. subalpina”
Male perianth at anthesis cleft to c. 2/3-4/5 ...........ceceeeeeeeeees H. obscura”
Bark of twigs not flaking. Leaves membranous
Bark of twigs flaking or not. Leaves coriaceous. Male pedicel at base not
articulated. Anthers 5-8. Androphore rather broad, tapering, (0.1-) 0.2-0.3
mm long. Kerangas forest, mountain forest; 800-1200 m alt.; Sarawak,
NUE cet has nSs one maaan dsb <idsvela en scntalnecmeeass H. xanthina”
Pedicel of male flowers articulate at base. Anthers 7-9. Androphore rather
broad and tapering, 0.2-0.5 mm long. Forest, 0-1000 m; Malaya; Sumatra
I Mea c Si esicewaid on a nbicicinbie ss svise0%s susie spins H. majuscula®”
Pedicel at base not articulated. Anthers 5 or 6. Androphore narrow,
0.1-0.2 mm long, hidden by the anthers. Forest, c. 100-450m,; C. Celebes
aN Ce REE pep vale <8 fos cic Sloteiats ols /sle bisiaisia-siale alsietaisiane sia asians H. coriacea”
Twigs moderately stout, towards the apex c. 2.5-3 (-4) mm diam. Leaves (8-)
12 cm long or more. Male perianth globose or subglobose, diam. c.
Mase? INIT oe eee im eee sca eine ores 2 8 aie <(e so sles sins siooeisem(saieiaie nieleiseieiniselsise== 108
Twigs slender, diam. towards the apex 1.5-2.0 mm. Leaves c. 5-12 cm long.
Perianth + ellipsoid to globose, 1.2-1.8 mm long ............ H. penangiana’”’”
Male perianth at anthesis cleft to c. 3/4-4/5. Anthers 7-11 ...............5+5 109
Male perianth at anthesis cleft to c. 1/3-2/3. Anthers 9-20. Sumatra, Java
ees eng tek Si Sa5h58 2 Ancoc ssenteccss cosas sdcheectecsdscee-eStetesessecstcesecs's 110
Anthers 7-9. Dry fruits c. 4-5 cm long, pericarp 10-20 mm thick. N.
NETL IGA VAS BOTTICO\ 855-2 sas gece a odds soo sain snjnae= sain H. punctatifolia®”’
Anthers c. 11. Dry fruits c. 2 cm long, pericarp c. 1.5 mm thick. Malaya
RPE RISD RN Sd Hs AD tet coe cca acces senetenees tenes teasasneseten ase H. punctata”
Gard. Bull. Sing 37(2)(1984)
Male perianth 3.0-4.2 mm diam.; anthers 15-20............. H. macrothyrsa”’
Male perianth c. 1.5-2.5 mm diam.; anthers 9-15. A variable species with 3
VATICNOS ? 55 evi ebasddese ideasads seule hiacesaatTcae emanates erat ao tennaeeNe H. glabra’
(2-7) REGIONAL KEYS TO THE SPECIES FOR FEMALE FLOWERING
AND FRUITING SPECIMENS
based partly on vegetative characters and distribution
(2) CEYLON, CONTINENTAL SE. ASIA (INCL. PENINSULAR THAILAND, EXCL. MALAYSIA
la.
2a.
3a.
4a.
Sa.
6a.
Ta.
8a.
AND SINGAPORE), AND ANDAMAN SL.
Ovary pubescent. Fruit pubescent at least at the base, or in H. kingii
glabrescent dse<sct ansadd egeepent . elaewmenseael shee etree eee eee eee 2
Ovary glabrous Fruit glabrousiy.)5.cscdeins. ccheebecs ete ce tele eee 4
Female perianth pubescent at least at the base. Leaves glabrous or late-
glabrescent below, or with vestigial tomentum on the midrib. Fruit c. 2.5 cm
LONG OT MOTE: gies nases.s deweacignsao'acels uesakieeb alent eae Se ae la satee CRE RE ERE ER eae anna 3
Perianth glabrous. Leaves with persistent tomentum below. Fruit c. 1.5-2.0
cm long, glabrescent but with vestigial tomentum towards the base;
perianth not persisting. S. penins. Thailand ............00..0000+ H. tomentosa®’
Perianth pubescent at base. Fruit ellipsoid, c. 2.5-4 cm long, wholly pubes-
cent; perianth not persisting. Geylon):.\ienc-cssscccten ates teeet H. iryaghedhi’
Perianth wholly pubescent, or partly glabrescent. Fruit + ellipsoid-oblong,
(3-) 4-6 cm long, glabrescent, with the perianth persisting. NE. India to S.
0) (111) RRR REE EPEESE o ho enocnaan st tosecp arco cic rn ubeorcpiymacrgan ands H. kingii?
Twigs towards apex either subterete, or usually + flattened or angular, and
lined or ridged from petiole to petiole. Phyllotaxis distichous ................ >)
Twigs terete, not lined. Leaves distichous or in 3-5 rows along the twigs.
Fruit ellipsoid 3.2.4) cb ccb coche cecdesbine oteeees osapte Rhea eas te aie een 6
Twigs lined. Perianth 2-valved. Leaves often with irregularly shaped whitish
marks or blotches here and there. Fruit globose, c. 1.5-2.0 cm diam.,
pericarp 1-2 mm thick; seed globose. Plant usually growing not too far
SOM IRE COGSE .cstoories dei comuehdinboodtiausbaretiete tee cae enna H. irya’
Twigs more prominently lined or ridged, usually distinctly angular in
section. Leaves without whitish marks. Fruit ellipsoid, c. 2.0-3.0 cm long,
pericarp 1.5-4 mm thick; seed ellipsoid. Not coastal penins. Thailand
ae wcsisved aa Tenningse nanactanted oma onaece tae qataste tence teen as Se ee eee enna H. brachiata”
Bark of twigs drying pale, grey-brown, contrasting with the blackish colour
of the dried petiole . Fruit drying blackish, c. 3.0-5.5 cm long; pericarp c.
5-15 mm thick. Perianth 3- (or 4-) valved. Penins. Thailand ...............0000.
Twigs drying brown, not much contrasting with the petiole. Fruit usually
(dark) brown, c. 1.5-3.5 cm long; pericarp c. 1-2.5 mm thick. Perianth 2- or
3=Valved ssc vscasvessostaocoeamnsse eumostusene tees ueeaeectenaeeet Care eee mmr eerntee 7
Fruit c. 1.5 cm long. Leaves distichous.C. Vietnam (Annam) H. longiflora;
Fruit generally larger, c. 1.8-3.4 cm long. Leaves distichous or dispersed .. 8
Leaves in 3-5 rows along the twigs. Tomentum of leaf bud composed of
hairs c. 0.1-0.3 mm long. Male perianth c. 1.0-1.5 mm diam. /ndo-China
New account of Horsfieldia 153
2a.
3a.
4a.
Sa.
6a.
at
8a.
9a.
Re oe eee eh aroiete cee 5 aes aisle Said widen ale a'5 araTaions Suhelotets disc ee a alMeeide v Sewlddeebide H. thorelii’
Leaves either distichous or dispersed in 3 or 5 rows. Tomentum of leaf bud
composed of hairs either c. 0.1 mm long or c. 0.5-1.0 mm long. Male
perianth c. 1.5-2.3 mm diam. NE. India, S. China (Yunnan), Indo-China,
3 UTE TELU LOL et eee le ca H. amygdalina*
(3) MALAYA, SINGAPORE
Female perianth at base pubescent; ovary and fruit pubescent. Fruit ellip-
soid, c. 2.5-4 cm long. Flowers in dense clusters and strongly fragrant in
male specimens. Originating from Ceylon, cultivated in Penang, Singapore
I EEE at oe iota Sosia}a aes cle sroiaors'se ole os win cidlai's's easravecenaoaeewe x H. iryaghedhi’
2 LTE DLT CIBC UG ac eee ees So SRR SA lek Aaa 2
Pea VAIVER” OVATY PIADIOUS! <oicio0sc0006050000ce00ecesetecdecerssaniavecsion ne 3
Perianth 3- (or 4-) valved. Ovary glabrous or pubescent ..................00005 5
Twigs usually shallowly ridged or lined from petiole to petiole. Leaves
membranous, often with irregularly shaped whitish marks or blotches here
and there. Fruit globose, c. 1.5-2.0 cm diam., glabrous; pericarp 1-2 mm
thick; seed globose. Plant usually growing not too far from the coast
NNO RE re RE IS ve asaisiesclasomisaawid ais Se psstais tiololoiore slagaiorceroaaler H. irya‘
Twigs not lined. Leaves without pale blotches. Fruit and seed ellipsoid .... 4
Leaves membranous, glabrous beneath, not brown dotted. Gardens’
PE SEIOTIPTOIF EE Ph e290 28S. SUI USRa GON leh Sud aeend Joedetewendinn H. parviflora’?
Leaves coriaceous, pubescent below and with regularly spaced, small,
brown to blackish dots (lens!). Kerangas forest, peat swamp forest ...........
«sc ne DUA ODE AONE ROT CEEEIB CO HOS BOCE ERE EE RACES Ene enEMHnoE tere seeks H. crassifolia®*
Leaves on lower surface with + regularly spaced, dark brown or blackish
dots being cork warts originating from hair scars; dots not to be confused
with usually smaller dark punctation of different origin, present or not
TINY) EE Reece eens ccs eis ooesdsbpnisdesisiure ce eeienierseinaceuemeenccers 19
MERION COLCOIMENEAGN 2.08. ccc 5tnssuinuc-P- dex ds se sdnarnew ns cuplehosbeeruestlen +e. 6
Beaves with persistent tomentum beneath ............2.2.2.0. ees scnccteceteceeces 7
MEAVES PIADFOUS OF PlADTESCENT DENALI. &. oc... 2. caccccesnseeecsssccooucessesees 1]
Ovary pubescent. Fruit pubescent or at least with vestigial tomentum near
the base; perianth not persisting under the fruit ................ H. tomentosa®
Ovary glabrous or only with some incidental minute hairs. Fruit glabrous,
me periantn (at least at first) PeEISISLING /0./..66.2.60.000.0cceecencsoncasonssneseses 8
Hairs on lower (and upper) leaf surface harsh, with hardened hair-bases, in
older leaves rendering the surface scabrous. Fruit c. 1.0-1.4cm long..........
eS ee osc aia sinc seictasbroths eins sie Tejeide eau seo elaaes aero ane H. grandis‘
Twigs of moderate habit, towards the apex c. 3-5 mm diam. Tomentum of
leaf bud and the very twig apex composed of hairs c. 0.2-0.3 mm long.
Leaves drying dull above, 13-21 cm long. Fruit c. 2.2-2.4 (-3.0) cm long
I REE RE Corie 1 P1955. te dadnaaaadaweuicsmascceenaeaneeeotebacserts H. fulva™’
154
10a.
ilar
12a.
13a.
16a.
Gard. Bull. Sing 37(2)(1984)
Twigs stouter, diam. towards apex c. 5-10 mm. Tomentum with hairs c. 0.5
mm long or more. Leaves c. 20-40 (-70)icm long ..2-4..tesseestees cee eee 10
Tomentum of leaf bud, twig apex and lower leaf surface with hairs c.
0.5-1.0 mm long, rather stiff, rust-coloured. Fruit c. 3.8-5.5 cm long
Jareinielaacle ateleaee Suma aostcloe sts neonate einai cee eRe ere ene Raee eee ree eee H. superba’
Tomentum composed of woolly hairs c. 1.0-2.0 mm long, yellow-brown or
pale brown. Fruit:cy3icmilongeeacscoteee ee ee H. flocculosa®?’
Bark of twigs drying rather pale, pale brown or yellowish brown, or straw,
contrasting with the blackish colour of the dried petiole. Phyllotaxis either
distichous, or the leaves in 3-5 rows along the twigs ................cceeeeeeees 12
Twigs drying brown, the colour not considerably contrasting with the
petiole: ‘Leaves distichous::.....c:e.-csacasuieaseveccn eset eae ene eee 13
Leaves distichous or in 3 rows. Fruit c. 2.5-3.5 cm long, with the perianth
persisting. Pedicel articulated at base (this character best seen in male
FIOWEES) ives gs nisdis avian Teplonete Bence ceee eee ene H. sucosa subsp. sucosa**’
Leaves in 3-5 rows along the twigs. Fruit c. 3-5.5 cm long, the perianth not
persisting. Pedicel at base not articwlated2..--.0--s5h.4e cena eee H. sparsa~’
Leaves drying very dull above because of finely wrinkled surface; nerves flat
or sunken. Fruit usually with persistent perianth ..................... H. fulva®
Leaves above not particularly dull, not finely wrinkled; nerves flat or
raised. Perianth not persisting under the prulin....-.se--ecce se -eeene eet 14
ateral nerves flatonbuttamtlynaisediabovems.nssaeseeeeoeetsseteeeeeteeene 15
Lateral nerves on upper leaf surface distinctly raised. Pedicel at base
articulated (this character best seen in male flowers) .................:ee0ee0+ 16
Tomentum of leaf bud composed of hairs c. 0.1 mm long. Twigs towards
the apex c. 2.5-5 mm diam. Leaves 15-27 cm long. Pedicel at.base not
articulated! 8.50)... ccoscee ote icetintrereenene H. subalpina subsp. subalpina?”’
Tomentum of leaf bud with hairs c. 0.2-0.4 mm long. Twigs slender,
towards the apex c 1.5-3.5 mm diam. Leaves c. 5-15 cm long. Pedicel at
base articulated (:<..ccecsccceacsueaseeneee au eco eee eee eee H. ridleyana”
Tomentum of leaf bud composed of hairs c. 0.1 mm long. Midrib beneath
early glabrescent. Fruit c. 4.5-6.5 cm long, usually with thick pericarp
Re PRO Se eee eee eb oroe re ae ce ocekk cadens H. majuscula”’
Tomentum of leaf bud with hairs c. 0.1 mm long or usually much longer.
Midrib beneath often rather late glabrescent. Fruit c. 2-4 cm long, pericarp
©. -2-5:(-7) mam thiiekorz isn sssasatects sao ansacoe aieatenee soca te een eee eee eee 17
Twigs + angular by lines or low ridges from petiole to petiole. Fruit c. 2-3
(-4.0) eM TONG: oo. 2. abs ctinaes ree ecee nee Le heer naa ota a a see nee eee H. brachiata”
Twigs terete, neither lined nor ridged y.ic.cssccs-ce cee seeeacnsee eee ce eee 18
Twigs towards the apex c. 2-5 mm diam. Leaves usually chartaceous. Fruit
19-3 -SxCm ONG sc 2: ses cewastees come eee OR CREA CER H. polyspherula®’
Twigs more delicate, towards the apex c. 1-3 mm diam. Leaves mem-
branOuss EnuUitien2-5-244.clm ONG eeeae eee eeteeener eee reenter H. macilenta”
New account of Horsfieldia SS
Bila:
22a.
la.
2a.
3a.
4a.
Sa.
6a.
Leaves usually with persistent tomentum beneath
Leaves glabrous or glabrescent beneath
Lower leaf surface with small dots and dashes; pubescent, sometimes late
glabrescent. Ovary glabrous. Fruit c. 4-6 cm long, glabrous, with the
PermAasTISUAlly PEXSIStENE G2). 05.. ooa08 edokslae eh adonbe Bede eee H. wallichii*’
Lower leaf surface always pubescent, provided with only dots, not with
streaks. Ovary pubescent. Fruit c. 1.6-1.8 cm long, shaggy-hairy, the
Pai AU RIIIO PDCKSISLOMG ooo cio oars dsicie donickinicsjacie na daeawictcedeegeulets H. pulcherrima*’
Twigs delicate, towards the apex c. 1.5-2 mm diam. Fruit c. 1.1-2.0 cm long
See ects totale tata tecies eyeiaaa sis a newicis slew ote sh cierdlas seaman igasaaactice H. penangiana”’
Twigs somewhat stouter, diam. towards apex c. 2.5-5 mm. Fruit c. 2 cm
2 BER TOTS Pac ee ee ol a 22
Leaves coriaceous, tip blunt or subacute. Fruit c. 2-2.3 cm long, pericarp
thin. A mountain species of central Malaya ............0.20ceee0ee H. punctata”
Leaves membranous, tip acute-acuminate. Fruit 4.5-8 cm long, pericarp
10-20 mm thick. Forests up toc. 1100 m. alt. ..........0... H. punctatifolia®
(4) SUMATRA, JAVA
Female perianth at base pubescent; ovary pubescent. Fruit ellipsoid, c. 2.5-4
cm long, pubescent. Male specimens with the flowers in dense clusters,
strongly fragrant. Cultivated, originating from Ceylon ....... H. iryaghedhi’
Estat pe AD OUSWHA corte cee ene ie seala swe a bata ciclod oi elves a Sleeoeetee Suinsatee dase ceeen 2
Leaves membranous, often with irregularly shaped whitish blotches here
and there. Female perianth 2-valved; ovary glabrous. Fruit globose, c.
1.5-2.0 cm diam., glabrous; pericarp 1-2 mm thick; seed globose. Plant
usually growing not too far from the COGS ..........0cccccencenccnceenees H. iryaé
Leaves of various consistency, usually not white-blotched. Fruit ellipsoid;
RECRMEINESOIG. $1 G71 COASLA) ODNOL oc. cesacelsseese tse aldo Gelbade Shcaddlee Seseae wd wwe 3
Perianth 2-valved. Lower leaf surface with + regularly scattered brown to
blackish dots, originating from hair scars; dots not to be confused with
usually smaller punctation of different origin (lens!) ...................eeeeeeee 4
Perianth 3-(or 4-) valved. Lower leaf surface dotted or not .................... 5
Leaves coriaceous, 10-20 (-28) cm long, finely pubescent on the lower sur-
face. Twigs towards apex 2-6 mm diam. Fruit c. 1.5-2.2 cm long, with per-
MIBLEHE PELIANUM. EO? SWAIIP fOTESL 12.0.2. .00c.catecssnneneencns H. crassifolia
Leaves membranous or thinly chartaceous, 5-12 cm long, glabrous beneath.
Twigs c. 1.5-2 mm diam. Fruits 1-2 cm long; perianth not persisting. Mixed
CEE, cts ARES COEE CS COE LCC EC EE EEE ROR EECA Een he cone Sencerrece H. penangiana”’
Ovary and fruit (at least at base) pubescent. Lower leaf surface with persis-
LO MMAMTOMTIETIRUETAN Gee eo oss synod w seiaie/cS n'a clare sina oN aiero ni beewnsiva eee ono useSoeloe Evloce aac 6
Twigs towards the apex c. 2-5 mm diam. Leaves c. 9-27 cm long, lower
surface without dots. Fruit with hairs c. 0.5 mm long or less. Thailand,
156
Ta.
8a.
10a.
lla.
12a.
13a.
14a.
Gard. Bull. Sing 37(2)(1984)
Malaya; specimens from Sumatra nOt S€@N ........ccecceseeeeee H. tomentosa’
Twigs stouter, towards apex 5-8 mm diam. Leaves c. 20-36 cm long, lower
surface with regularly scattered brown to blackish dots (lens!). Fruit with
hairs, c. 2 MMplOny .oias2 och accoueeecoeac ees aaa eee eee H. pulcherrima”’
Lower leaf surface with more or less regularly spaced brown to blackish
dots (lens!). Lateral nerves in upper surface generally flat or sunken ...... 21
Lower leaf surface without dots. Nerves either raised, or flat to sunken ... 8
Lower leaf surface with persistent tomentum. Fruit with persistent perianth
BE re ire Leonie ier et At ea nOee SCE bon poanncbtnontbaneroegouss00 9
Lower leaf surface glabrous or glabrescent. Fruit with perianth persistent or
MOD Fo 5. nics vasa Sea tedenve code adccnecoceed obalnaes aaetidae chen etcnee tea ae eae ae ana eee 11
Hairs harsh, the hair scars making the lower surface of older leaves
scabrous? Fruit<l=1.4 ein long) eo ce tone ee meee cerns H. grandis
Olderleaves not'scabrous| beneathieeseeseese ete ee eee ene eee 10
Plant stout, twigs towards apex c. 5-8 mm diam.; leaves c. 20-40 (-70) cm
long. Tomentum of leaf bud with hairs c. 0.5-1.0 mm. Fruit 3.8-5.5 cm long
By Pn Re en a PPR San Sores bol Boseccdon GanbocsncodaBonbousunde H. superba’
Twigs towards the apex c. 3-5 mm diam.; leaves 13-21 cm long. Hairs on
leaf bud c. 0.2-0.3 mm long. Fruit c. 2-3 cm long ..................00. H. fulva®?
Lateral nerves in upper leaf surface flat or sunken or but faintly raised. On
drying, colour of lower leaf surface generally greyish brown, not con-
trasting with that of the upper surfaces ....ccc.asn0;scceeet oheoee aaa eee eee 12
Lateral nerves distinctly raised above. On drying, colour of the lower leaf
surface bright brown or chocolate, usually much contrasting with the upper
SUIPACE Tr. fic. PE Lass dacciod to taemee Somes sonman’ ately Seales neice Ceara Nees ne 16
Twigs delicate, towards the apex c. 1.5-3 mm diam.; leaf bud, the very twig
apex and young inflorescence woolly-pubescent with hairs c. 0.3-0.7 mm.
Leaves 5-9 cm long. Female flower and fruit not known ...... H. triandra’’
Twigs generally stouter, towards apex (2-) 3-10 mm diam.; hairs of leaf bud
and inflorescence c. 0.1-0.3 mm long. Leaves more than 10 cm long ..... 13
Leaves distichous, drying dull because of minutely wrinkled upper surface.
Bark of twigs straw-colour Or BrOWMs..220; oaccc. sees. cosets eee nee eee 14
Leaves distichous or arranged along the twigs in 3-5 rows; not particularly
dull from drying, upper surface not finely wrinkled. Bark of twigs pale,
grey-brown or straw, contrasting with the blackish colour of the dried
PELION |! SL FER SD. Hae hdres Mak iS o es. te See tee teehee ee 15
Leaves elliptic-oblong to oblong, drying olivaceous to brown above.
Stem grey-brown, not conspicuously contrasting with the colour of the
dried petiole. Fruit drying brown, c. 2.2-3 cm long, perianth persistent ......
s mu awinwiesarwpiaaewies vbldvw te slew qoute getadl cep oneal Ate acess acme vate eee ene H. fulva*’
Leaves elliptic-oblong to lanceolate, drying to a rather dark olivaceous
colour above. Stem pale + yellowish-brown, rather contrasting with the
petiole. Fruit drying blackish, c. 1.5 cm long, the perianth not persisting ....
He La wcind Sets SDs. isa TING ch Adon de Rade thebeats dathaatca teen cee E ee coe H. tristis*?
—— ne
——
New account of Horsfieldia ily
a:
l6a.
20a.
Zila.
22a.
Leaves distichous or in 3 rows. Fruit c. 2.5-3.5 cm long, with the perianth
persisting. Pedicel articulated at base (this character best seen in male
emerswere 4 eer So eyeres: Foi. Daeracelb. 325. H. sucosa subsp. sucosa*’
Leaves in 3-5 rows along the twigs. Fruit c. 3-5.5 cm long, the perianth not
Bostisune:.Pedicel-at base not articulated. . :.:.ssescicc3003sccscse0e see H. sparsa°’
Perianth 4-valved. Pedicel at base not articulate (this character best seen in
male flowers). (Female perianth known only from the remnants persisting
under young fruits). Fruit c. 8-9 cm long .................ccceeeeee eee H. valida‘
Perianth generally 3-valved. Pedicel articulate at base. Fruit up to 6.5 cm
op nesesdeitie EL ARS 2 Peet reid heed ae gee eee ee 17
Twigs angular or subterete, provided with lines or ridges from petiole to
ee IE CeCe cee dan ans Goce S vacates men nas voammee Cees coaek ee 18
Twigs terete or but faintly angular, neither lined nor ridged
Perianth pubescent (known only from male flowers). Fruit c. 5-6 cm long.
SUT AETEIT Brake cde aS Res CBE ee seperate op neers H. hirtiflora”’
Perianth glabrous.Fruit c. 2-4 cm long ..............00.c0ce0ce000. H. brachiata ”
Twigs delicate, towards the apex c. 1-3 mm diam. Leaves membranous,
midrib beneath late-glabrescent. Fruit c. 2.3-2.4cm long .... H. macilenta”
Twigs generally stouter, towards the apex c. 2-5 mm diam. Leaves usually
ene ODEN 9-6-5 Cia LONG. 6h oc. Sanes ocdecnearons od/endtecen tees teank 20
Tomentum of leaf bud and inflorescence composed of hairs c. 0.1-0.2 mm
long. Midrib beneath early glabrescent. Fruit c. 4.5-6.5 cm long ...............
SEE 2 tea? 1s ok ice Ubon eee hnkawds et uddem eee ee emet ue H. majuscula®”
Tomentum with hairs 0.1-0.6 mm long. Midrib often late-glabrescent. Fruit
es cunlone..A Vvaridble SPCCICS .. sci .innn0snosnennd-n0ees H. polyspherula®’
Lower leaf surface usually with persistent tomentum (sometimes glabres-
cent), and with small scattered dots and streaks (lens!). Twigs conspicuously
hollow. Fruit c. 4-6 cm long, the perianth generally persistent ..................
Leaves beneath glabrous or glabrescent; dotted, not streaked. Twigs not
PERIMCHOUSIYeMUNOW. PII VARIOUS sic-.c.sc0.cisececssencccaccnsen secon cdeeve sss 22
Bark of twigs pale, greyish to straw, much contrasting with the blackish
colour of dried petiole. Phyllotaxis dispersed, with the leaves in 3-5 rows.
Female flower and fruit not known. N. Aceh, atc. 1300m
RE obs Rice hate opicumede depp ease vepobwuaed «sudden S<seveaetaeetbs H. atjehensis*”
Bark of twigs brown, not contrasting in colour with the petiole. Leaves
PNCMGIS OL Mt SLOWS MP1. PIGDTE, DD. ose ons ss tence sasas scones ecsncdcsens 23
Fruit large, (4.5-) 5-8 cm long, with thick pericarp ......... H. punctatifolia”’
Hae. 1-2-5 cm long, pericarp much thinner (o..-- 2.0 5..<....cectecescoseeees 24
Twigs slender, diam. towards the apex c. 1.5-2 mm. Leaves 5-12 cm long.
AS ew U6 770 (6) a ni i H. penangiana”’
Twigs generally stouter, towards apex c. 2.5-4 (-6) mm diam. Leaves (8-) 12
ERMONEOF MIGre PINUC. £.8-2.5 CHT IONE: °°... .0.<--c.c-c.cs<cncnccsceceesn ass 25
la.
ue\-
3a.
4a.
8a.
9a.
Gard. Bull. Sing 37(2)(1984)
Leaves distichous. C. GaN Sumatran stot eee H. macrothyrsa”
Leaves distichous or dispersed in 3 rows. Java, S. Sumatra, Mentawai
Isis: north to) Simeulue Isls ts. eee ecase, cee eee aes H. glabra’”
(5) BORNEO
Leaves membranous, usually with irregularly shaped, whitish blotches here
and there. Female perianth 2-valved; ovary glabrous. Fruit globose, 1.5-2.0
cm diam., glabrous; pericarp 1-2 mm thick; seed globose. Plant usually
growing not (00 far frOmMmIhe COGSIcescccc sean: seeceanedaee aera H. irya®
Leaves various, usually not white-blotched. Fruit subglobose or ellipsoid;
seed ellipsoid. Plantcoastal Or NOP scac.c.ceewaseaeesiu esse eee eee eee eee 2
Perianth’ 2-valved 0.2. 25.300 2e ean dacthosaconbe snob ee eas Canas eee reese te eee EE ee 3
Perianth predominantly 3- (or 4-) valved <c2.:.2..---.2e- snes eee 5
Leaves coriaceous, the lower surface usually densely short-pubescent and
with scattered dots and streaks (lens!). Twigs grey-brown, not much con-
trasting with the colour of the dried petioles .................... H. crassifolia®®
Leaves membranous, the lower surface glabrous and without dots. Twigs
greyish or straw, rather contrasting with the blackish colour of the dried
PETION!’ . oo. aaaace Disha SiRahosa KA aoRaNeae gies On ae etc ELT es Cede 4
Female inflorescence + spike-like, 5-10 cm long. Perianth persistent under
the fruit: (always?) . SEs Sabah sees. oie eee H. sterilis”
Female inflorescence ramified, 1-2 cm long. Perianth not persisting under
the fruit. Sabah, E. & S. Kalimantan ............... H. sucosa subsp. bifissa**”
[eavesiin 3/0r more rows alone: theltwigsies.-csncee- ce stee cee eeee tear e eee eee 6
Leaves, GisticNouss. 5. sissies aclaacis oeinyetaoite sispie(paicielaslee os opeciene Sancta tae eee ee 8
Leaves generally bunched towards the top of the twigs. Petiole propor-
tionally long and slender, 25-50 mm long .....................e000: H. sabulosa”
Leaves not bunched. Petiole proportionally shorter ....................00e000+ 7
Leaves 10-30 cm long. Twigs pale, greyish or straw, contrasting with the
blackish colour of the dried petioles. Female perianth c. 2.5-3 mm long.
Fruit: 1, 5=4icm long ii ccpacasesiacce sonmaeeosneen etch anes seceneeee H. pallidicaula”
Leaves 20-45 cm. Twigs brown, not contrasting. Female perianth c. 4-5 mm
long,, Fruits.6-8) cit On Gig 53 ces icce wocatnotivcae cease sade ae eee H. fragillima*
Twigs in apical portion angular, i.e, distinctly lined or ridged from petiole
tO, POI ON. 6 once sa accesses per ne sospeame ater eeeteiers st ingstaa ls etaitn cnet ater H. brachiata”
Twigs terete or but faintly angular, neither distinctly lined nor ridged ...... 9
Leaves on lower surface with persistent tomentum (sometimes largely
plabrescent 1 Fl. WQILICHI), .ojcetsesjsintdaersisncins du dsaeisigqegsnesesnide sae eee eae 10
Leaves beneath glabrous or early glabrescent (midrib sometimes late-
plabrescent). 2) 0. .o be. ge. scee cgedete qansbieneecieee set mns ee ae renee cera ss neat aeeaee eae 19
New account of Horsfieldia 159
10a.
17a.
Plant stout; leaves large, c. 50 cm long, petiole c. 3 mm long only. Lowland
PORN Sete I lak cincis.na sxcne Secaeccs cdaceedeesecaiusanse H. sessilifolia~’
Older leaves on upper and lower surface with scabrous hair scars. Fruit c.
PE PRC TRNOSTNE Sire oat Pees Sor Peeks ohne coed vendecdaberstees H. grandis”
eaves nat scabrous. Fruits generally larger «..............0...ossesssecesssseeae 12
Leaves on lower surface with scattered dark brown or blackish dots and/or
S057 75 (ESR) ee ee ee ee ee ee ee H. wallichii’’
Mowemleasurtace without dots or streaks <....-ssceccccevswseseosceseete e+e on 13
Perianth with persistent tomentum; perianth not persistent under mature
ET eer nese ns once tea toce ta secducesgscecess sseeesies H. motleyi®’
Perianth glabrous or early glabrescent, persistent under the fruit or not .. 14
Leaves distinctly bullate due to deep-lying veins and with + revolute leaf-
margin. Female flower and fruit not KNOWN ............00.0000+ H. reticulata®’
82 ES CEC TRG Vil en 15
Ovary pubescent; fruit sometimes pubescent only towards the base ....... 16
MITE AIGA AATISI ORAL OUS oe 5 cctasdie ca's/daio aiias Sac ce Sasnapiode esluio otuienat aeseiewpiamsacciense 17
Twigs moderately stout, 3.5-5 mm diam. towards apex. Leaves 10-23 cm
long; nerves 11-16 pairs. Fruit largely glabrescent; perianth not persisting
_ con 0006600H0G0006 O00 CUBS AEB BEE ACE EEE Stee ae eee H. rufo-lanata®
Twigs stout, 4-7 mm diam. Leaves 18-45 cm long, nerves 18-25 pairs. Fruit
pubescent, with persisting perianth ..............:...ccseeeceeceees H. splendida®™
Twigs rather stout, towards the apex 3-6 mm diam. Leaves 18-35 cm long;
Mewes Mi-A0palrsebnuit2). 3-27 (Cn lONG, J.scsenssneecee-teacecescse eee H. affinis®
Twigs slender, 1.5-3 mm diam., leaves generally smaller. Fruit c. 1-1.5 cm
Re se ha), 2s ois je c Hem MSiwn oh G Ves sekinaieidanasee wees «ssi 18
Beavesmembranous; nerves 14-17 pairs ...20. 0c. cc. cede cee cee ee H. gracilis”
Leaves thinly chartaceous; nerves 5-9 pairs ................0005 H. paucinervis®’
Lower leaf surface with + regularly spaced, brown to blackish dots or with
dots and striae (lens!); dots originating from hair scars, not to be confused
with usually smaller and irregularly spaced dots or punctation of different
OOTEIID cnotucnodesd dSeunbGseL GORCEEC ap RDOM ince RESET CnC One cece Cn ccceac erent ae a een acete 20
Lower leaf surface without dots (sometimes with enlarged hair scars) ..... 23
Tomentum of leaf bud, the twig apex and young inflorescences composed
OlANAINSeO 22, MIMMONPOWMOLE. j.cecescecree ceeesetaeasee et aseee eco ae ceee lcs oe 21
Tomentum very short, with hairs c. 0.1 mm long or less ...................05 22
Twigs conspicuously hollow. Leaves on lower surface often with persistent
tomentum. Pedicel at base not articulate. Perianth generally persistent
Be STMT RIC Mt tie hye Masia Perel sn cia cabetirive aicneeitste eet hicclevincinimehOnetls H. wallichii*’
Twigs not conspicuously hollow. Lower leaf surface glabrescent. Pedicel at
160
22ae
Dan
26a.
29a.
30a.
31a.
Gard. Bull. Sing 37(2)(1984)
base articulate. Perianth not persisting under the fruit ....... H. borneensis*°
Twigs rather delicate, diam. towards the apex c. 1.5-2 mm. Leaves c. 5-12
cm long; nerves 8-11 pairs. Fruit 1.1-2.0 cm long, pericarp thin ................
ais anlaiva'sWinindio-w bie nie Waioleia eles Salcieta sta wrerelns NOAMSER GR See anes aaa eee H. penangiana”’
Plant stouter, diam. of twigs towards apex c. 2.5-4 mm. Leaves 9-21 cm
long, nerves 11-16 pairs. Fruit 4.5-8 cm long, with thick pericarp ..............
uginbss34 05RR0G LESSER E SEARS aE SR ON SEM DDE Ch Mei cate atelstmaraiale apietinete H. punctatifolia”
Bark of twigs drying to a pale colour, grey-brown or grey-yellowish, con-
trasting with the dark brown or blackish colour of the drying petioles .... 24
Twigs drying brown, not or but contrasting little with the petioles ......... 26
Leaves membranous, drying brown or blackish brown, the lower surface
somewhat paler. Perianth persisting under the fruit. Mixed forest.............
Jo 3Ss ASIA toads SRL PIER es 25 SE eee SE ee ry Cerne H. pallidicaula”
Leaves usually chartaceous, drying bright brown or olivaceous, lower sur-
face usually brown or chocolate. Perianth not persisting under the fruit .....
Pee mm meee eee eee eee EEE EEE HEHE EE EE EEE EH EEEEEE EEE EEE HEHE EEEE HEHEHE HEHE EEE EEE EEE EES
Twigs slender, diam. towards the apex c. 2-3 mm. Leaves 7-16 cm long, on
lower surface bright brown or chocolate, contrasting well with the grey-
olivaceous upper surface. Fruit 1.8-2.7 cm long. Kerangas forest, peat-
SOTES SAAS a Ee ee H. oligocarpa’’
Twigs stouter, c. 3-10 mm diam. Leaves 13-35 cm long, the lower surface
not conspicuously contrasting in colour. Fruit c. 1.5-2.0 cm long .......... 28
Tomentum of leaf bud and immature inflorescence very short, with hairs c.
0.1 mm long or less. Lateral nerves on upper leaf surface flat or sunken, or
but-little raised! 2.0.4.5. Wouticlveninecme « dees dotee eces cv Oats mega eey eee ee een 2H
Tomentum composed of hairs c. 0.1 mm long or more; lateral nerves raised
or not above; if hairs only c. 0.1 mm long, then the lateral nerves, at least in
the lower halifi<distinctly ralsediabOVGmrace seeteeceeer reece ratte eet errs 31
Leaves on upper side dull, i.e. on drying the surface finely wrinkling ...... 28
Leaves above not conspicuously dull, surface not finely wrinkling ......... 29
Leaves elliptic-oblong to oblong. Fruit c. 1.6-2.0 cm long. Heath forest,
DCGlE SWATD J OFS U ici: sin.o.<cigawcg ora hastene eee denen eee ee H. carnosa”’
Leaves elliptic-oblong to lanceolate. Fruit c. 1.5 cm long. Mixed forest
winaviaie « Genlaidtpland bilge aeaba tn slid patentee alana sain eae AS acre H tristis*?
Lowland species, up to c. 700 m alt. Leaves membranous. Fruit 5-5.5 cm
long; pericarp’ thick ‘Slisi.tecceceaeonootot ace amoee noes eae ree eeteeee H. obscura”
Mountainous species; c. 800-1800 m alt. Leaves membranous or coriaceous.
Fruit generally smaller «i... ciccisa ces scotia den eet cette nck te 30
Leaves on lower surface without distinct large hair-scars (lens!). Fruit 3-5
CMON GS 5. cece cssew sak se etnn re uke ser anne Re H. subalpina subsp. kinabaluensis””
Leaves on lower surface usually with distinct enlarged hair-scars. Fruit
(immature) c. 1-15 cmilOng. 2 2.ccccsasscsckedossnessonsmeeeteeee meen H. xanthina”
Pedicels at base not articulate (this character best seen in male flowers).
i
\
|
|
|
--> =r
ial Pan
New account of Horsfieldia 16]
32a.
35a.
36a.
sae
Fruit with persisting perianth or not
Pedicel at base articulate. Perianth not persisting under the fruit
Leaves generally large, 15-45 cm long. Fruit c. 6 cm long or more; perianth
persisting for a long time under it. Lowland or mountainous forest ....... 33
Leaves generally smaller, c. 5-20 cm long. Fruit c. 2-4 cm long; perianth
not persisting. Montane forest at 800-2000 Mm alt. ........c.cccccccecesesceceees 35
Nerves 11-22 pairs. Sabah, Mt. Kinabalu at 1000-1500 m .............c.cccceeees
_cosodbocStibot GGCUO EL ES COSC ne ECCOR DERE OC ECOSOC OORCRCEES AA AEEEe aaa amee H. amplomontana‘?
Female flower and fruit not known. Hallier 624, Mt. Damoes, W. Kaliman-
tan; apparently an undescribed species close to H. valida, see notes ...........
{0 on¢600S GEE CODE ODUM ORS EE Su abe BB EEOs SERRE DSAR cee SORE AEC eens inne aff. H. valida*
Female perianth 4-5 mm long. Fruit c. 6-8 cm long, pericarp 10-20 mm
(LIDS esos Ae ee ee H. fragillima®
Leaves membranous, apex acute-acuminate. Fruit 2.4-3 cm long.............
A a ieee ee ere he des ore wlaisitale evaded eels de clacg oeecte ea’ H. androphora*’
Leaves chartaceous to coriaceous, apex rounded to (sub) acute, not acute-
RU AGME BC P= ae 8 hs Fd oy. ENG oo Ss Se R's Ss whe Scceple’s Dawe es 36
Leaves chartaceous to coriaceous, lower surface without large hair scars
(lens!). Female perianth c. 2 mm long. Fruit c. 2.0-2.7 cm long
Leaves usually strongly coriaceous, usually with large and distinct hair scars
on lower surface. Female perianth c. 2.5-3 mm long. Fruit c. 3-4 cm long
ME MRS ETT cele Palas vig be ee laierels See eGis Seat shoes avae neeecc eben H. endertii*’
Female inflorescence, female flower and fruit not known. Male in-
florescence very stout, the rhachis towards the base 5-8 mm diam. W.
NAUTICA CLT) PR RT cs arcinlcisidle'elale! sala crereiatoateleralomrooblelele beans H. pachyrachis”™
NAL CHITIONESCENGCERMESSISTOUL ton ce nooo e oases tener e en ern re ee nee nclaasas 38
Midrib on upper leaf surface towards the transition to the petiole c. 3 mm
EL TES CIOS. Agee ae ee ae Re ee a ee oe H. laticostata”
Leaves 16-28 cm long, base short-attenuate to rounded; nerves 16-19 pairs.
SITTIN Cie nena Rane Deemer Seba ney Sara 328 858 DE I59. Rae dee mnls H. nervosa‘?
Leaves 5-28 cm long, base either rounded or short- or long-attenuate; nerves
eM DATOULES, PRE ah SSeE LL t 8 Sek oct .Seee ss. Bares whlemmntplseluicle qaiclsSencescssigedese vee 40
Lateral nerves on upper leaf surface sunken or flattish, or but slightly raised
ceed) Pea bes eat asin rc Ps Har ssicigaas ibe ss stared abindieeacielae o@esinep eaten sees 41
Ireral Merv es GIStiNGthy FAISEG ADOVE oc... ceca ee co ncnncnacacerareessenmncissc 42
Medi apex TOUNGed. ETUll MOL SEEM .. 2. ii ce ccc occ teres sceberessenaess H. obtusa”
Leaf apex acute-acuminate. Fruit c. 1.5-2 cm long .............. H. ridleyana”
43a.
2a.
3a.
4a.
Sa.
Gard. Bull. Sing 37(2)(1984)
Tomentum of leaf bud and young inflorescence with hairs c. 0.1-0.2 mm
long. Leaves drying dull, greyish-brown, the colours of the upper and lower
surface not much contrasting. Fruit c. 1.7-2.0 cm long ........ H. tenuifolia”
Tomentum of leaf bud c. 0.2 mm long or more; or if c. 0.1 mm long, then
the olivaceous to dark brown colour of the dried leaf above much con-
trasting with the cinnamon colour benealh....---.: 2a. e eee eee 43
Twigs delicate, towards the apex c. 1-3 mm diam. Leaves generally mem-
branous, 10-18 (-27) cm long. Fruit c. 2.3-2.4cm long ......... H. macilenta”
Twigs generally stouter, towards apex c. 2-5 mm diam. Leaves chartaceous,
of:various Size. Eruitsof various Size: ccasciesceesss see deta 44
Twigs early glabrescent; older bark + longitudinally cracking. Leaf apex
long acute-acuminate. Fruit c. 2.8-3.2 cm long; pericarp hard-woody, 8-10
mint thick’. (BYUMCIET actos.cg-0 ce. see oeeer eee ete eee eee ae eee H. disticha”
Twigs usually rather late glabrescent; older bark striate, not cracking. Leaf
apex acute-acuminate, the acumen not conspicuously long. Fruit variable,
1:9-6:cnmlongs Whole Of BOrneo.ss2eec escent eee H. polyspherula®’
(6) PHILIPPINES, CELEBES, MOLUCCAS
Leaves membranous, usually with irregularly shaped whitish blotches here
and there. Female perianth 2-valved; ovary glabrous. Fruit perfectly
globose, 1.5-2.0 cm diam., glabrous; pericarp 1-2 mm thick; seed globose.
Plant usually growing not too far from the COS ........0.ccceeeeeeees H. irya’
Leaves various, usually not white-blotched. Fruit subglobose or ellipsoid;
seed usually ellipsoid: Plant coastal OP NOL sisesccsn.-<cceaciosseeen ere te eee 2
Perianth °3-valved:'Ovary and fruit glabrous <:00...-6--cees-e cee eee eee 3
Perianth: -2-valvedss.i5..0022ohecewtes botentaneet snc ape -bOee ees nese e ee nee ee ene 5
Tomentum of leaf bud and inflorescence with hairs c. 0.2-0.6 mm long.
Pedicel at base articulated. Fruit c. 2.0 (-2.5) cm long. Philippines:
Mindana: 22285 Beak eee H. polyspherula var. polyspherula’/<
Tomentum composed of hairs c. 0.1-0.2 mm long. Pedicel not articulated
TOerrrrrrrrrrrrr rrr rere ee eee eee eee eee eee eee ee ee ee ry
Fruit 3.5-7 cm long; dry pericarp 8-15 mm thick. Leaves + membranous,
drying olivaceous to brown, midrib glabrous above; leaves sometimes with
whitish marks as in H. irya. Philippines, Celebes ..............4+ H. costulata®’
Fruit c. 4 cm long, dry pericarp 3.5-8 mm thick. Leaves membranous to
thinly coriaceous, drying brown; midrib towards the base in younger leaves
pubescent above: (C. Celebes icc scsterncesteoescensneceneseon sect eeeee H. coriacea”
Plant stout; twigs towards apex 4-14 (-20) mm diam., leaves c. 20-45 cm
long, petiole short, 2-7 mm long. Tomentum of leaf bud and inflorescence
with hairs c. 0.3-1.0 (-1.5) mm long. Female perianth c. 3.5-5 mm long,
glabrous; ovary glabrous. Fruit c. 3.5-5.5 cm long, glabrous .................++
Cece ccc reece ccc c cece cece ese e cece esereeseeeresesessesesesesessssereneeeseoneseS
Plant generally less stout, petiole relatively longer. Tomentum composed of
hairs up to 0.2 mm long. Female perianth c. 3.0 (-3.5) mm long or less. Fruit
up'to 3'cm long; in Hlancifoliaup to 3iS emilong te.c..-c.e--5- cece cere 6
New account of Horsfieldia 163
6a.
Ta.
8a.
9a.
10a.
lla.
12a.
13a.
14a.
Leaves + chartaceous, oblong-lanceolate to lanceolate. Female perianth at
anthesis cleft to c. 1/4; ovary pubescent. Fruit often + pear-shaped, 2.5-3.5
cm long, early glabrescent; dry pericarp 4-8 mm thick. Celebes ................
eRe rae ance as ak tawialad onias Sd vale’ s SeaewENeied. cea dacslgcaneees H. lancifolia*’
Leaves of various consistency, generally broader, oblong to oblong-
lanceolate. Female perianth at anthesis cleft to c. 1/3 or more. Fruit c.
1.0-3.0 cm long
Ovary and fruit pubescent; hairs on fruit may be very small and in-
conspicuous and only remaining at the very base near the insertion of the
EEAMIEHS PE CIICAL) CMICKSOM TIMIN ojo5e0:0 «2i0.dotefinje' «iso eroisisioeierho veiaidiore wis cianateur cles wnsc'al 8
Ovary and fruit glabrous; dry pericarp thin, c. 1-2 mm. (Female flower and
fruit not known in H. samarensis'4 and H. aruana?’) .............0cceeceeeeeee 11
Female perianth c. 2.5-3 mm long, at anthesis cleft to c. 1/3-1/2. Fruit (1.6-)
1.8-2.8 (-3.0) cm long; dry pericarp 2-3 mm thick ................ H. laevigata**
Female flower not known. Fruit smaller, pericarp thinner ..................... 9
Fruit 1.5-1.6 cm long, short-ellipsoid. (Male perianth transversely ellipsoid,
at anthesis cleft to c. 2/3-4/5). Leaves 8-30 cm long, membranous to char-
taceous; nerves flat, inconspicuous. Twigs terete, not ridged. Talaud Isls.
NMC IO SE esis abi Co sisiaginwbieisla vireiowe on winenisekeitee H. talaudensis”’
SSSA E ea OI UES IPS Io 18 (0) 7 ee 10
Fruit subglobose. Twigs towards apex + flattened, usually lined or lowly
ridged from petiole to petiole. Leaves c. 12-25 cm long, membranous;
nerves flat, inconspicuous. (Male perianth + pear-shaped, cleft to c. 2/3).
LLL OLGETS ajateedes ganda aoa cee areca eo eee H. decalvata**
Fruit short-ellipsoid. Twigs terete, not lined. Leaves 5-14 cm long,
chartaceous, nerves very inconspicuous on both surfaces. (Male perianth +
obtriangular, cleft to c. 1/2-way). Philippines: Luzon .... H. obscurinervia’’
Twigs towards apex angular or ridged. (Species distinctive only in male
EC ESPICCIIMETIS x seen oe eterna asic ace caeac sac swoadee ue soasaindesmsceaavenasene 12
Twigs terete or subterete, neither angular nor ridged; twigs sometimes faint-
ly angular or shallowly lined from petiole to petiole ....................2.2000+ 13
SW. New Guinea, possibly Aru- and Tanimbar Isls. (Female flower and
Le TN PMT O ba KIT OWN) Bret eece aac Sesinc sc ciscck sare sismuck aortedeseeiaac is cuiileiviere H. aruana’®
Moluccas: Ceram, Banda, Dammar IsI., possibly Ternate. (Ovary glabrous)
Po oie ora cin hielo ie ctelsiobs'els v's Scioto slobigaaiaw oddapialeas H. smithii’
Bark of twigs drying pale, grey-brown, contrasting with the blackish colour
of the drying petioles. Fruit drying blackish, 1.5-2.0 cm long. Moluccas
nn Fania RRO ORT AN WN So en cuile cate atilboliceatiscnewate dee sissitarcestate’ H. spicata’
Fruit globose to subellipsoid, 0.9-1.2 cm long; drying blackish. Aru Isls.,
INCWIGILINICCR ent tne sn cee hoses eciecnecoaissiesiasisesescess H. subtilis var. subtilis?”
Fruit c. 1.1-1.6 cm long. Fruit not known in H. samarensis .............0.0++ 15
la.
Dae
3a.
4a.
Dae
6a.
Wale
8a.
Gard. Bull. Sing 37(2)(1984)
Philippines: Samardsisi . Sass uo. ce eee ee H. samarensis’4
MOLUCCOS ee Gacdec RM Boe hc en Ee ee 16
Fruit ellipsoid, c. 1.5 cm long; drying blackish. Morotai, Obi Isl. ............
sisi scaled wf ote a a Oe eae eee ee H. moluccana var moluccana”’
Fruit subglobose or ellipsoid, 1.1-1.6 cm long, drying brown. Celebes,
Kabaena Isl... COrai scccatertcten tar ete en Ree H. parviflora!’
(7) NEW GUINEA, SOLOMON ISLS., CAROLINE (PALAU) !SLS., AUSTRALIA
Leaves membranous, often with irregularly shaped whitish blotches here
and there. Female perianth 2-valved; ovary glabrous. Fruit perfectly
globose, c. 1.5-2.0 cm diam., glabrous; pericarp c. 1-2 mm thick; seed
globose. Plant usually growing not too far from the coast ........... H. irya’
Leaves of various consistency, generally without whitish blotches. Fruit
either ellipsoid or globose; if globose either only c. 1 cm diam. (H. subtilis),
or the pericarp more than 2 mm thick, at least at one side; seed mostly ellip-
soid. Fruit glabrous or pubescent. Plant coastal Or NOL .......1.csceeceeeeeeees 2
Plant from Australia. Female perianth 2-valved; ovary pubescent. Fruit
ellipsoid, 1.8-2.2 cm long, drying orange to brown; pericarp 1-2 mm thick
sneianhe Jubs.oes apaniesiwnaawawinteiads canoe PeaSeEe BREE oee eRe eee H. australiana”’
Twigs towards apex distinctly angled or ridged from petiole to petiole. Aru
Isls... New: Guinea o25A .asits tide eolo a BRI BE ee 4
Twigs terete, sometimes lined in between the bases of petioles but neither
angled nor ridged. New Guinea to SOIOMOMN ISIS. .........cceceecenseceecencecees 9
Leaves on lower surface with regularly scattered blackish-brown non-
traumatic dots or cork warts originating from hair bases (lens!). Perianth
2-valved. Vogelkop to. W., Sepik DISt. — .s.2c25.te een eee eee H. inflexa*®
Leaves without blackish-brown'dOts ” <..0..c.6.scnccsec- ceconne cate eee eae 5
Perianth 3-(or 4-) valved «. ssesstsdeocascendtoace nce teat eee see eee ee eee 6
Perianth 2-valved’ ii: chi 2 ol ascahcasedachaeecehccte cach canteen eee EEE eee 7
Ovary glabrous (?). Fruit 10-16 mm long, glabrous. Leaves chartaceous,
7-14 cm long; petiole relatively long and slender, 11-20 mm long. SW. and
S. New Guinea (Digul VWesternyD Sta) nase-ceseteeene seen eee H. olens’’
Ovary pubescent. Fruit 17-20 mm long, pubescent at base. Leaves mem-
branous to thinly chartaceous, 10-27 cm long; petioles 7-15 mm. Vogelkop
DENINS Saichinna sigcenvawlswows ene ber siiilesteistad cae mrada daa ease nee ae mea H. angularis”’
Female perianth depressed-globose, valves nearly 1 mm thick; ovary pubes-
cent. Fruit 17-20 mm long, pubescent. Leaves membranous to thinly char-
taceous: .VogelKop) Denins; S.iGasccebans de eet Reena eeeiseeee H. angularis”’
Female flower andfnuit snot kmowiniesnesnsssceeeencneneencececanceccn tease 8
Leaves membranous. SW. New Guinea; possibly Aru and Tanimbar Isls.
ne eee eer i BAe Be BE Oe dob oupndo aces dosed cbodnd0GscuN000. H. aruana”*
Leaves thinlyiconaceousi Sia New Guinea ences eseesmaceeseee stare H. iriana’’
New account of Horsfieldia 165
9a.
10a.
lla.
14a.
Sai.
l6a.
17a.
18a.
Perianth 3-(or 4-) valved; at anthesis cleft almost to the base. East Sepik
nS te Bee el none pind eas ae Oe a H. sepikensis’’
Perianth 2-valved
RR MME AMAR ENUIEIP EA DEOUS aca nn Giso Sec atina kh raisinaisaanacned cs oxcaccasaawassasoecascess 11
Ovary and fruit pubescent; hairs on fruit either distinct or small and in-
conspicuous and only to be seen remaining at the base of the fruit near the
insertion of the stalk (lens!). Fruit ellipsoid, sometimes in H. sinclairii
globose
Tomentum of leaf bud, apex of twig, and inflorescence, with hairs 0.3-1.5
mm long. Leaves large, 17-45 cm long, often + parallel-sided, nerves 30-40
pairs. Fruit ellipsoid, 3.4-5.5 cm long, glabrous. Moluccas and W. & C.
MENS (GRATED). SocqepocASh GSN NC GOSH ODHEE COCR EE UE CTE ECE CECE EOE CE EERE H. sylvestris!’
Tomentum with hairs c. 0.2 mm long or less; hairs in H. moluccana’ and
H. tuberculata?? c. 0.1-0.3 mm long. Leaves generally smaller, nerves fewer
Ee ee Ee Seamer oh oeT anise aoiaie nies seleisiaeislelsienlesscinsle slovoed emenclleeisier {72
Fruit globose or subglobose, not beaked and without pseudostalk, c. 1.4 cm
SARC Sek eRe P eee Ln. SR cl Miac dete coke tecccc canes aeSSavs eat eds daw ee. 13
Fruit drying brown; dry pericarp c. 1.5-3 mm thick. Female perianth at an-
thesis cleft nearly to the base. Northern parts of Irian Jaya and of Papua
MASA ES Settee. eed, be sees... SF ede «AEM piSsa cos oda ssieistaad te. H. basifissa*’
Fruit drying blackish; dry pericarp c. 1 mm thick. Female perianth at
anthesis cleft to c. 1/3. Aru Isls. Whole of New Guinea ...........00cceeeeeccnees
eR eee nc tiban aciectiwsieae acess: see'esesisc sees H. subtilis var. subtilis?”
Fruit up to c. 2.0 cm long, drying blackish; top pointed/beaked or not, base
ManonGOL with iong-or short pseudOstalk ............s.--s000scesccnsnceeoneoees 15
Fruit 1.3-3.7 cm long, drying brown or dark brown; top rounded, base
SL ONT NERS SRC ICES 0 Oe ee ee ee ee ee 16
Pseudostalk of fruit (1.5-) 2-6 mm long. Jayapura Dist. (Irian Jaya), W.
QB aINe JADA eae sS 06 G0C Rot Scar B ECC OP BRE CE AREER er een Ere H. schlechteri?’
Twigs lined from petiole to petiole. Fruit 2.5-3.0 cm long; pericarp c. 2-3
mm thick, without coarse wart-like lenticels. Palau Isls. .... H. palauensis’’
Twigs lined or not. Fruit 1.3-3.7 cm long; pericarp thin or thick, with small
BmAnCeati-like IENLICElS OF NOL... ..c.nmcres «nczanssboalss eb seed eee obeseee 17
Female perianth c. 2 mm long, cleft at anthesis to c. 1/2-4/5. Fruit 1.3-2.8
cm long, dry pericarp 1-2 mm thick. Moluccas, W. New Guinea ............+..
Brame eamr sb ca eieeceichioitltec Aesakiccsuw sews slg ih shew enue. cot a NOlMCcAnAa’
Female perianth 2-3 mm long, cleft at anthesis to c. 1/2-2/3. Fruits 1.5-3.7
cm long, pericarp 1-8 mm thick. Caroline Isls. (incl. Palau), Bismarck
Arch., Papuan Isls. and E. New Guinea (Milne Bay Prov.).H. tuberculata?’
Arches of the submarginal nerve on the lower leaf surface distinct and very
166
19a.
20a.
21a.
26a.
27a.
Gard. Bull. Sing 37(2)(1984)
regularly looping; nerves often reddish brown. Leaves often + parallel-
sided. Tomentum of leaf bud composed of hairs c. 0.1-0.3 (-0.4) mm long.
Female inflorescence and infructescence up to c. 7 cm long. Fruit 1.7-2.5
(-3.4) cm long, drying orange to brown; pericarp 1-2 mm thick. Solomon
TSIS.. ctsasvasirthe stevie Shayeee de Resse eee H. whitmorei*’
Marginal nerve not very regularly looping. Plant not from the Solomon Isls.
Tomentum of leaf bud, apex of twigs, and inflorescence short, composed of
hairs c. 0.2 mm long or less; in H. psilantha*} hairs c. 0.1-0.3 mm long
a a\d-ai.aiia siknayd'a etd 6 ai cla siveldelsie nea raiminee cto tia ee eee erate ne ee Seto ae eee 20
Tomentum long, with hairs c. 0.5-1.0 mm long; hairs 0.2-0.5 mm long in H.
ampliformis?®; tomentum not known in H. ampla*s ..........0.cccecceceseeees 28
Tomentum of leaf bud composed of hairs c. 0.1-0.3 mm long. In-
fructescence (and 9 inflorescence) large, much branched, 10-16 cm long.
Fruit 1.7-2.2 cm long, pericarp thin, 1-2 mm thick, brown. Bismarck Arch.,
Bagabag. si) LOnZISIS .. cc. csccceutasuceoneeeeecee ete eee eee H. psilantha*’
Tomentum of hairs 0.1-0.2 mm long, or less. Female inflorescence and in-
fructescence c. 10cm long or less. Fruits various .................ssscesessees- 21
Female perianth incl. pedicel long-obconical. Fruit stalk tapering, thickened
towards the fruit. Mature fruit not known. E. New Guinea: Morobe Dist.
BR ee er O BTS Hee Lc HERB SSE DED Bo SOBUOR wTooDooeeodaedac H. crux-melitensis?’
Female perianth + ellipsoid, well marked off from the slender pedicel.
Fruit stalk not tapering: Ao..-.csecsee5o 5.00 en cone eee ee See 22
Fruit 1.6 cm long or less; pericarp 1-3 mm thick. Perianth pubescent ...... 23
Fruit 1.5 cm long or more; pericarp 2 mm thick or more; if fruit c. 1.5 cm
long then almost globose and perianth glabrous ..................ceceeeeeeeeees 25
Fruit without pseudostalk, top rounded! >. 2.2... carpe sceessncaees H. pilifera”’
Fruit with pseudostalk c. 1.5-5 mm long, top rounded or usually acute,
SOMEUIMES TOSILALE’ 5. cise ses scnues te cameencteeonsase mn anenens Merete eniee eee ae 24
Leaves drying olivaceous; midrib distinctly raised ................. H. clavata”’
Leaves drying dark brown; midrib slightly raised ............ H. squamulosa”’
Female perianth c. 2-2.4 mm long, glabrous. Fruit (sub)globose, or short-
ellipsoid, or obovoid, c. 1.5-2.5 x 1.5-2.0 cm; pericarp 4-6 mm thick.
Leaves 6-14 cm long. E. New Guinea (Papua New Guinea) .... H. sinclairii??
Female perianth c. 2.5 mm long or more, pubescent or glabrescent. Leaves
1O‘em long Or MOTE. cu. cssetkeeweceoeee eo ee eee Reet 26
Fruit 1.6-3.0 cm long, usually with coarse pale-coloured wart-like lenticels;
dry pericarp 2-6 mm thick. Moluccas, New Guinea, and Bismarck Arch.;
0-1000 m alt. (sometimes much resembling small-fruited H. pachycarpa‘*')
sudden belmen aaa cnmeGiict sabe csluo noe Sen hene CER ECE eee ORE RC LC EEE EEE een ae H. laevigata*’
Fruit (3.0-) 3.5-7.5 cm long; pericarp (4-) 5 mm thick or more. New
Guinea; (450-) TO00=2000 Mm GIONS), BOT dave enn sea oe anh oe Renee ee 27
Perianth pubescent. Fruit 3.0-4.5 cm long; pericarp 4-10 mm thick ..........
Ee ee ere Pete te SS oorabooo soDdonensodoadtbectoustbobcoct H. pachycarpa’’
New account of Horsfieldia 167
b. Perianth glabrescent. Fruit 6-7.5 cm long; pericarp 10-20 mm thick ..........
ees: ecb las aero 2: oe Ae ae eh H. corrugata”’
28a. Tomentum of leaf bud and inflorescence composed of hairs c. 0.2-0.5 mm
long. Female perianth c. 3 mm long, at anthesis cleft to c. 2/3. Fruit not
known. Tomentum of leaf bud, female flower and fruit also not known in
DUE oe 29
b. Tomentum composed of hairs c. 0.5-1.0 (-1.5) mm long. Fruit usually con-
SUSAN NU EITM CSE CINE ac ech ofa apne alaors cntsd pusieiarlorekie saa jaihienledundawdsacbeckioos 30
29a. Inflorescence glabrescent. Papua New Guinea: Sepik Prov. ...... H. ampla?4
b. Inflorescence pubescent. Sepik and Morobe Prov. .......... H. ampliformis25
30a. Leaves coriaceous, beneath with harsh hairs, when hairs shed leaving
thickened scars. Female perianth c. 4 mm long, at anthesis opening with a
narrow pore-like slit at apex. Fruit 3.0-5.0 cm long, pericarp 4-7 mm thick.
INN ENGD (GTA Thee axene ac ic AO OSS b See BDO EEE HARA E COE EEE RRAEE EacHE Ee H. pulverulenta”
b. Leaves membranous or chartaceous. Perianth at anthesis cleft to c. 1/4-1/2
ee renee en crte chit) nia Wee demir, Sls eth ha clldiciadine vw os Sd aesh adda 31
3la. Flowers (only the male known) entirely pubescent. Fruit 2.0-2.4 cm long,
Penieanp 4-7 mm thick: New Guined «-. 2.2. s22i2c.2..0.seesee dees H. leptantha*
b. Flowers largely glabrescent
32a. Fruit 2.5-3.0 cm long. Female perianth c. 4 mm long. Leaves generally
oblong-ianceolate, at apex caudate. New Britain ............... H. ralunensis*
b. Fruit 1.2-2.8 cm long. Female perianth c. 3 mm long. Leaves oblong to
oblong-lanceolate, at apex not caudate (always ?). New Guinea and New
TI PED Sosciss delaware clas dss a> sta onipinavonlvaneneaeaccetilgeshed esc H. hellwigii”
Enumeration and Description of Species
1. Horsfieldia iryaghedhi (Gaertn.) Warb. Figs. 1A(1); 2 I; 3.
Mpristica iryaghedhi Gaertn., De Fruct. et Sem. Pl. 1 (1788) 196, t. 41, f. 4 — Horsfieldia iryaghedhi
(Gaertn.) Warb., Mon. Myrist. (1897) 332, t. 21 fig. 1-4; Sinclair, Gard. Bull. Sing. 28 (1975) 68 —
Type: Gaertner’s drawing.
? Phelima Noronha, Verh. Batav. Genootschap Kunsten en Wetenschappen 5, art. 4 (1790) (3) et (edit.
1827) 66, nom. nud.
M. glomerata Thunb., Acta Holm. sive Vet. Akad. Nya Handl. (1799) 88, t. 2. f. 1 (non Migq.,
1852) — Type: Herb. Thunberg, n.v.; microfiche: fragment of male inflor.
H. odorata Willd., Sp. Pl. (ed. 4) 4, 2 (1805) 872 — Type: not known (see Sinclair, l.c.).
M. horsfieldii (first spelling horsfieldia) B., Bijdr. 2, 11 (1826) 577; Rumphia 1 (1837) 192, t. 63; King,
Ann. Roy. Bot. Gard. Calc. (1891) 296, pl. 122, 123 — Pyrrhosa horsfieldii (Bl.) Hasskarl, Cat. Pl.
Hort. Bog. (1844) 174. — Type: Blume s.n., Jave (L).
M. notha auct. non Wall. Koenig ex BI., Rumphia 1 (1837) 192, nom. nud. pro syn.
Myristica (Cnema) glomerata Migq., P|. Jungh. (1852) 170 (non Thunb., 1799) — Type: Junghuhn s.n.,
Java (U; iso L).
Myristica odorata Reinw. ex de Vriese, Pl. Ind. Bat. Or. 2 (1857) 95 (non odorata Willd.), nom. nud. pro
syn.
For further references see Sinclair, 1.c.
168 Gard. Bull. Sing 37(2)(1984)
Tree 5-25 m. Twigs terete, not ridged, towards the apex 2-5(-14) mm diam., at
first with woolly tomentum, of rust-yellow hairs c. 0.5 mm, twigs early to late
glabrescent, bark grey-brown, rather coarsely striate, lower down coarsely striate or
longitudinally cracking, rarely somewhat flaking; lenticels usually present but
sometimes inconspicuous. Leaves in 2 rows, chartaceous, ovate-elliptic to oblong-
lanceolate, broadest usually at about the middle, 10-28 x 4-12 cm, base rounded to
attenuate, top acute-acuminate, upper surface glabrous, usually drying dark brown,
or dark olivaceous, lower surface early or late glabrescent, without brown-black
dots, the epidermis always finely papillose; midrib above often late-glabrescent, flat
or slightly raised; nerves 9-16 pairs, above flat or sunken, the lateral arches not very
distinct; tertiary venation forming a coarse network, usually + trabeculate, above
distinct to hardly visible; petioles late-glabrescent, 15-20 (-25) x 2-3 mm; leaf bud c.
10-15 x 3-4 mm, densely pubescent with hairs c. 0.5 mm long. Inflorescences densely
woolly pubescent with hairs c. 0.3-0.5 mm, in d : c. 6-15 x 4-10 cm, usually twice
ramified, the branches rather few, thickish, common peduncle c. 0.3-2.0 cm, the
flowers united into c. 10-25 subglobose dense clusters or capituli c. 5-10 mm diam.,
each with c. 80-100 flowers, the capituli being rather spaced along the branches; 9?
inflorescences smaller, little-branched, c. 1.54- x 1.5-2 cm, the flowers solitary or a
few together; bracts broadly ovate or + triangular to elliptic, the larger ones with
distinct midnerve, c. 1-3 mm long, pubescent, caducous. Flowers in 6 : 3 (or
4)-valved, in 2: 3 (or 2)-valved; perianth glabrous except towards the very base;
pedicel short or absent, pubescent, at base not articulated. Male perianth narrowly
+ obovoid-oblong or obconical, 3-5 (or 6)-angular, c. 2-2.5 x 1-1.6 mm, top
broadly rounded, gradually tapering to the base, glabrous but with a few hairs c. 0.2
(-0.5) mm long at the transition to the pubescent pedicel: 0-0.2 (-0.3) mm long;
perianth at anthesis cleft at first to c. 1/5-1/4, in adult usually up to c. 3/4; valves c.
0.2 (-0.3) mm thick. Androecium elongate, narrowly obconical or blunt ellipsoid-
oblong, top + truncate, base tapering, c. 1.0-1.2 x 0.4-0.5 mm; anthers 3-5 (i.e.,
6-10 thecae), largely sessile, erect, free apices c. 0.2 mm, central column narrow,
hollow to about half-way, androphore narrow, c. 0.2-0.5 mm long. Female perianth
broadly ellipsoid, c. 3 x 2 mm, at anthesis cleft to c. 1/2-way, valves c. 0.3 (-0.5)
mm thick; ovary broadly ellipsoid, c. 2.0 x 1.5 mm, densely short-pubescent,
stigma minutely 2-4 (or more)-lobulate; pedicel absent or up to 0.2 mm. Fruits 3-8 in
a cluster, ellipsoid to ellipsoid-oblong, top and base broadly rounded, c. 2.5-4.2 x
1.7-2.4 cm, densely rust-yellowish pubescent with stellate-dendroid hairs c. 0.5 mm,
sometimes partly glabrescent, pericarp dark brown, not tuberculate, c. 1.5-2.0 (-3.0)
mm thick; stalk 0.5-2 mm long; perianth not persisting under the fruit.
Distribution: Ceylon, introduced in Malaya (Penang Isl.), Singapore, Java.
CEYLON: Balakrishnan NBK 935; Burman s.n., 33; Davidse & Sumithraarachchi 8539; herb.
Hooker s.n.; Huber s.n.; Jayasuriya & Bandaranaike 1869; Kostermans 23321; 24114, 26670, 27113;
Meijer 268, 276; Nooteboom (& Huber) 3164, 3188; herb. Pallas 51; herb. van Royen s.n.; Thwaites C.P.
221; Waas 743, 892; Walker s.n.; White & Arnott s.n.; Worthington 2130, 2291, 2325, 2345, 2346, 2582,
3533, 3535, 4124, 4832, 5201, 5228, 6023, 6474, 6580.
SINGAPORE (culta): Walker 267.
JAVA (culta, mainly W. Java): Backer 36342; Bakhuizen van den Brink 7319, 7409; Blume s.n.; van
Heurn s.n.; Junghuhn s.n. (42); Koorders 31019 3 ; Korthals s.n.; Martati (133); Radermacher 93; herb.
Reinwardtianum 154; Teysmanns.n.; de Vriese s.n.; Zollinger 3263 (? 3268).
Ecology. Lowland rain forest, wet evergreen forest, ‘‘intermediate’’ forest; also
in disturbed forest; 0-500 m. Flowers and fruits throughout the year.
Vernacular names. Ruk, Rukghedhi and Malaboda (Ceylon); Irie gaga (Ceylon,
Konig ms.); Tjampaka sélong (Java, Soenda).
Uses. Formerly in Java a wax was obtained from cooking the fruits. The wood
is moderately heavy and even-grained (see Sinclair, p. 72).
Horsfieldia iryaghedhi (Gaertn.) Warb.
a. habit of twig with leafy shoot and female inflorescences, x 1/2; b. twig portion with male in-
florescence, x 1/2; c. subspherical male flower head, flowers partly removed, one flower
opened, x 3; d, mature male flower bud, lateral view, x 12; e, ditto, longitudinal section show-
ing androecium, x 12; f, mature female flower bud, x 12; g, ditto, opened, showing pubescent
ovary and sessile, minutely lobulate stigmas, x 12; A. infructescence, x 1/2, i-/. seeds with and
without seed coat, x 1/2. — a, f, g from Anon., Hort. Bot. Sub. XIII-E-9; b-e from Jayasurya
& Bandaranaike 1869; h-j from Kostermans 26670.
169
170 Gard. Bull. Sing 37(2)(1984)
NOTES
1. Fieldnotes. Trees to c. 25 m tall, at base to c. 50 cm diam., branches droop-
ing, sometimes branched from the base. Stilt roots sometimes present when growing
in wet soil. Bark hard, outerbark light brown or black-brown or greyish-red,
fissured or gritty, or peeling off in large pieces, or smooth; living bark 5-10 mm,
beefy red or red brown, with white lines, exuding a clear light reddish brown sap;
wood white to light yellow with red streaks. Leaves grey-green beneath. Perianth
yellow or dark yellow, or orange brown; pistil with brown hairs. Fruits yellow or
yellowish brown with rusty tomentum; aril complete, orange to deep red. Fresh male
flowers c. 3 mm long; flowers strongly scented, reminiscent of that of Michelia
champaca, according to Sinclair emanating again on boiling the dried flowers.
2. This species rather deviates from all other Horsfieldias, and is now placed
by me in a separate section. It is mainly distinct by the leaves being papillate
beneath, male flowers arranged in compact heads, rather-many-lobulate stigma; and
according to Warburg (p. 334), the seed contains some starch, which seems to be
unique for this species of Horsfieldia.
According to Sinclair (l.c.) its closest relative might be H. wallichii because in
that species, before anthesis the flowers are almost sessile and clustered as well, and
because of similar leaves. However, when mature, the flowers of H. wallichii are
quite different, especially because of the much broader androecium; also, its leaves
have typical minute dots and stripes beneath, but are not papillose.
Warburg placed H. iryaghedhi and H. sylvestris in a section Orthanthera
because the flowers are similarly arranged into small dense capitula in the later
species. At full anthesis, however, the flower heads of H. sylvestris loosen con-
siderably and their inflorescences then link up with other Horsfieldias.
The arrangement of flowers in heads somewhat resembling dense capitula is
found in the African genus Pycnanthus and the genus Brochoneura from
Madagascar.
2. Horsfieldia kingii (Hook. f.) Warb. Figs. 1A(2); 4.
Mpristica kingii Hook. f., Fl. Br. Ind. 5 (1886) 106; King, Ann. Roy. Bot. Gard. Calc. (1891) 300, pl. 127;
Kanjilal & Das, 11. Assam 4 (1940) 43; C.Y. Wu, Fl. Yunnanica 1 (1977) 10, fig. 4 (9-10); Tsiang Li &
Li, Fl. Rep. Popul. Sin. 30, 2 (1979) 202, fig. 92 — Horsfieldia kingii (Hook. f.) Warb., Mon.
Myrist. (1897) 308; Sinclair, Gard. Bull. Sing. 28 (1975) 74. — Type: King s.n. (Sikkim, 19th June
1881) (CAL, n.v.; K, iso; BM: G, 7.v., lecto); Masters s.n. CAL, n.v.; K, iso; L, P; BO, DD, M,
PDA, 7.v.).
H. hainanensis Merr., Lingnan Sc. J. 11 (1932) 43; Tsiang, Li & Li, Fl. Rep. Popul. Sin. 30, 2 (1979) 199,
fig. 91 — Type: Tsang & Fung 17851 (NY, 7.v.; iso K).
H. tetratepalaC.Y. Wu, Acta Phytotax. Sin. 6 (1957) 218; Fl. Yunnanica 1 (1977) 12, fig. 4(1-8); Tsiang,
Li & Li, Fl. Rep. Popul. Sin. 30, 2 (1979) 197, fig. 90 — Type: Exp. Sino-ross. Yunnan 2770 (KNN,
PE 752):
Tree 6-25 m. Twigs terete (not ridged) towards the upper 4-10 (-12) mm diam.,
bark dark brown, rather early glabrescent, tomentum grey-brown to light brown,
hairs c. 0.2-0.4 mm long, bark lower down + coarsely striate, not flaking;
lenticels usually distinct. Leaves in (3-) 5 rows, membranous to chartaceous,
(obovate to) elliptic-oblong to oblong-lanceolate, broadest at or usually somewhat
above the middle, 12-35 (-55) x 5-17 (-22) cm, base attenuate, top acute-acuminate;
upper surface drying dark olivaceous-brown to dark brown; lower surface largely
early glabrescent but usually some tomentum vestigial fora while on and near the
midrib, without brown-black dots; midrib flat above; nerves 13-18 pairs, flat or
slightly sunken above, marginal arches indistinct; tertiary venation forming a coarse
network generally faintly visible above; petioles (9-) 15-25 x (2-) 2.5-4 mm, glabres-
New account of Horsfieldia 171
cent rather late; leaf bud c. 15-20 x 3-4 mm, densely grey brown to pale brown
pubescent with hairs c. 0.2-0.4 mm. Inflorescences rather thinly woolly-
pubescent with greyish to pale brown hairs c. 0.2-0.4 mm, in ¢ : rather slender to
broad, 2-3 times ramified, flowers moderate to rather few, c. (5-) 7-16 x 3-8 (-12)
cm, common peduncle c. 10-20 mm long; in 9 : 1-2 times ramified, rather few-
flowered, 2-8 cm long; bracts oblong to lanceolate, pubescent, 3-10 mm long,
caducous. Flowers in male (see notes to the synonyms) 2- or usually 3- or 4- (or 5-)
valved, in ? either 2- or 3-valved, in d arranged in loose or dense clusters of 3-10,
in 2 c. 1-3 together; perianths pubescent with soft hairs 0.1-0.3 mm long, or partly
glabrescent; pedicels pubescent to subglabrescent, at base not articulated. Male
perianth in mature bud globose or subglobose, usually slightly angled on the valve-
sutures, c. 2.5-3.5 x 3.0-4.0 mm, top rounded or slightly acute, base broadly
rounded; pedicel stoutish, 1.0-2.0 (-2.5) mm long; perianth at anthesis cleft to 2/3-
3/4, valves 0.5-0.8 mm thick, sometimes rather coriaceous. Androecium depressed-
globose or depressed and broadly obovoid, top broadly rounded to subtruncate, c.
1.0-1.6 X 1.5-2.0 mm, in transverse section subcircular to bluntly 3- (or 4-) angular;
anthers (12-) 14-16 (-20), largely sessile, free apices c. 0.1-0.3 mm, towards apex
curved over and into the rather broad apical cavity, cavity c. 0.3-0.8 mm deep;
androphore rather narrow, c. 0.1-0.3 mm long. Female perianth 2- or 3-valved,
ovoid-ellipsoid, 3.0-4 (-5) x 3.0-3.5mm, thinly pubescent or subglabrescent, at
anthesis cleft to c. 1/2-way, valves c. 0.6 mm thick, + coriaceous, ovary ellipsoid or
obovoid, c. 2.0-2.5 x 2.0 mm, pubescent, stigma minutely 2-lobed, 0.1-0.2 mm
long, pedicel 2-3 mm long. Fruits 1-6 per infructescence, ellipsoid to ellipsoid-
oblong, top rounded to (sub) acute, base narrowly rounded to rather long-tapering,
c. (3-) 4-6 x 2-2.7 cm, dry pericarp 3-4 mm thick with granulate-striate surface, dry-
ing dark brown, glabrescent early; stalk 3-8 mm; perianth persisting.
Distribution. India (Assam, Sikkim), E. Nepal, Bangladesh (?, no specimens
seen), Burma (?, no specimens seen), China: Yunnan, Kwangsi (no specimens seen),
Hainan.
INDIA. Bengal: Haines 842 — Assam: Masters s.n.; herb. Pierre 5462 — Sikkim: Gammie, King’s
Coll. s.n.; King 2380; Rogers s.n.
E. NEPAL: Stainton 6436, 6880.
CHINA. Yunnan: Henry 12.234 — Hainan: Tsang & Fung (317) L.U. 17851.
Ecology. Lower montane forest, in gullies of evergreen forest; 300-1200 m alt.
Flowers and fruits probably throughout the year.
Vernacular names. (See Sinclair, p. 75) Amol (Assam); Mijing-ikum-asing
(Miri); Pandikachoaphang (Kach.); Si/tui (Lushai); Bolong, Bolouchi (Garo).
Uses. Kernel used as a substitute for areca nut in betel chewing. The gum is
good against mouthsores.
NOTES
1. Fieldnotes. Bark grey or brownish, somewhat rough, flaking in small square
flakes, and longitudinally fissured; sap blood red. Male flowers recorded as orange-
red, fruits as yellow.
2. Synonyms. In the Flora of China (Fl. Rep. Popul. Sin.) (1979), besides
H. kingii two related species H. tetratepala and H. hainanensis are accepted. I have
seen no material of H. tetratepala and H. hainanensis only the sterile isotype in K,
and some separate flowers of the same collection kindly procured by the herbarium
at Berkeley, but after studying the descriptions and the figures in the Flora
of Yunnan (1977) and the Flora of China (1979) I see no reason to keep them
separate. In the Flora of China the three species are keyed out on vegetation
iWi2 Gard. Bull. Sing 37(2)(1984)
characters viz., the presence or absence of lenticels on the twigs, the degree of per-
sistence of the tomentum on the lower midrib, the number of lateral veins (which
strongly overlap), and the length of the infructescences: 6-12 cm in H. tetratepala
and 2-4 cm in H. hainanensis.
However, the following remarks should be made.
1. H. tetratepala: in the Flora of Yunnan, figure 4, 3-4, a distinct cup-like disk
under the androecium is visible, whereas in the figure of the same species in Flora of
China, figure 91, this cup-like disk is absent. If, in fact, a disk is present, com-
parable to the one present in the Malayan Knema plumulosa, then this would be a
strong argument to regard H. fetratepala as a distinct species.
2. H. kingii. In Flora of China, figures 92, 2, the male perianths were drawn as
glabrous and 2-valved, whereas for H. tetratepala and H. hainanensis they are
presented as pubescent and 3- or 4- (or 5-) valved. The perianths in the type of
H. kingii, however, being also pubescent and 3-valved indicate this species be
regarded as conspecific. If in the specimens used for the drawing of what is called
H. kingii in Flora of China the male flowers are truly glabrous, and 2-valved, this
would be a reason for further investigation on the status of these specimens, which
might represent a new taxon.
3. Henry 12.234 from Yunnan, a good male flowering collection, was iden-
tified by Sinclair as H. macrocoma, for reasons unclear to me, possibly because of
the rather slender male inflorescences.
3. Horsfieldia longiflora de Wilde, sp. nov. Fig. 1A(3)
Horsfieldia amygdalinae affinis, differt foliis distichis, perianthiis masculis 2-valvibus, + ellipsoideis
parum compressis, c. 2-3 mm longis, androecio c. 2.0 x 1.3 mm, antheris 10-12, fructibus breviter
ellipsoideis, c. 1.5 * 1.2 cm. — Type: Eberhardt 3050 (P).
Tree 6-20 m. Twigs terete, not ridged, towards the top 2.5-6 (-10) mm diam.,
bark dark grey-brown to blackish brown, tomentum with hairs c. 0.1 mm long or
less, grey-brown, early glabrescent, lower down the bark coarsely striate, not
flaking, lenticels distinct. Leaves in 2 rows, membranous to chartaceous, elliptic-
oblong to oblong-lanceolate, broadest at about the middle, 7-21 x 3-7.5 cm, base
attenuate, top acute-acuminate; upper surface drying olivaceous brown to dark
brown; lower surface glabrescent early, without larger dark brown dots; midrib
flattish above to slightly raised; nerves 9-13 pairs, flat to slightly raised, or sunken
above, marginal arches indistinct; tertiary venation forming a lax network, faint or
invisible above, usually faint beneath; petioles 9-12 x 1.5-2.5 mm, glabrous; leaf
bud slender, 10-14 = 2-3.5 mm, densely grey-brown pubescent with hairs c. 0.1 mm
long or less. Inflorescences with sparse tomentum of hairs c. 0.1 mm or
less, subglabrescent, ind: c. 3 times ramified, moderately- to many-flowered, c. 6-10
x 4-6 cm, common peduncle 6-20 mm; in?: 1-2 times ramified, c. 2-3 cm long;
bracts + elliptic, short-pubescent, 2-5 mm long, caducous. Flowers 2-, rarely with a
few 3-valved, the d in loose clusters of 3-6, the 2 c. 1-3 together, glabrous; pedicels
glabrous, at base not articulated. Male perianth in bud with frontal view broadly
ellipsoid-obovoid, laterally somewhat flattened, 2.2-3.0 x 1.8-2.0 mm, 1.2-1.5 mm
thick, top broadly rounded, base shortly rounded to subattenuate, outside faintly
longitudinally ribbed or not; pedicel slender, 1-2 mm long; perianth at anthesis cleft
to nearly 1/2-way, valves c. 0.2 mm thick. Androecium flattened, obovoid-ellipsoid,
top broadly rounded to subtruncate, 1.7-2.0 x 1.0-1.3 x 0.6-0.8 mm, in transverse
section ellipsoid; anthers 10-12, sessile, free apices c. 0.1 mm, suberect or little in-
curved; apical cavity narrow, 0.2-1.0 mm deep; androphore rather broad, tapering,
c. 0.2-0.3 mm long. Female perianth ovoid-ellipsoid, c. 2.5 x 2.0mm, glabrous, at
anthesis cleft to nearly 1/2-way, valves c. 0.3 mm thick, ovary ovoid, c. 1.8 x 1.5
mm, glabrous, stigma minutely 2-lobed, c. 0.1 mm long, pedicel c. 1.0-1.5 mm long.
toenar agent te
athe
a
Sm
Horsfieldia kingii (Hook. f.) Warb.
a, leafy twig with male inflorescences, note dispersed leaves, x 1/2; b, mature male flower bud,
lateral view, x 6; c, ditto, opened, showing androecium, x 6; d, androecium, longitudinal sec-
tion, schematic, x 6; e, mature female flower bud, x 6; f, ditto, opened, showing pubescent
ovary with minute bi-lobed stigma, x 6; g, twig portion with infructescence, note persistent
perianth under fruit, x 1/2. — a-d, from Haines 842; e & f, from Tsang & Fung 317; 2g, from
King’s Coll. s.n. (Sikkim).
173
Horsfieldia thorelii Lecomte
a, twig with leaves and male inflorescences, note dispersed phyllotaxis, x 1/2; b, mature male
flower, lateral view, x 12; c, ditto, opened, showing androecium, x 12; d, androecium,
longitudinal section, schematic, x 12; e, twig portion with female inflorescences, x 1/2; f,
mature female flower, laterla view, x 6; g, ditto, opened, showing glabrous ovary, x 12; A, twig
portion with infructescence with immature fruits, x 1/2; 7, mature fruit, x 1/2. — a-c, from
Poilane 19887, e-g, from Maxwell 75-212; h & i from Poilane 22394.
New account of Horsfieldia 7S)
Fruits 1-8 per infructescence, broadly ellipsoid, top and base rounded, c. 1.5 x 1.2
cm, surface granulate, not tuberculate, drying dark brown, glabrous, dry pericarp c.
1 mm thick; stalk 1.0-1.5 mm long; perianth not persisting.
Distribution: C. Vietnam (Annam)
VIETNAM. Annam: Clemens 3474; Eberhardt 3050; Polane 10224, 13529, 29315.
Ecology. Submontane evergreen forest, on rather poor soil; exact altitudes not
recorded; flowers March to May, fruits May-July.
Vernacular names. Lan Ham, Con na ham (Moi), Me tuong.
Uses. Leaves are used for bathing after childbirth. The wood is recorded as soft
and light.
NOTES
1. Flowers yellow, fruits red.
2. Closely related to H. amygdalina and H. thorelii, both differing by the
shorter and smaller male flowers with differently shaped androecium (but see note 3
under H. amygdalina), and by the larger fruits. In H. /ongiflora, however, fruits are
known only from one collection (Clemens 3474). All specimens of our present
species have the leaves distichous; in H. thorelii all specimens have the leaves
dispersed, in H. amygdalina both distichous and dispersed phyllotaxes can be
found.
3. Sinclair included the specimens of the present new species in his broadly con-
ceived H. glabra. The specimens were all collected after the treatment in the Flore
Génerale de |’ Indo-chine (1914).
4. Horsfieldia thorelii Lecomte Figs. 1A(4); 5.
H. thorelii Lecomte, Not. Syst. 1, 4 (1909) 99; Fl. Gén. I-C. 5, 2 (1914) 100; Sinclair, Gard. Bull. Sing. 16
(1958) 422 (See note, in syn. of H. amygdalina) — Type: Thorel s.n. and 1186 (P) (some of the
duplicates are H. irya).
Tree 4-20 m. Twigs terete, not ridged, towards the top 2.5-6 (-12) mm diam.,
bark dark grey-brown to dark brown, early glabrescent from a grey-brown to rusty
tomentum with hairs c. 0.1-0.3 mm, bark lower down finely to + coarsely striate,
not flaking, lenticels small, generally inconspicuous. Leaves in 3-5 rows, mem-
branous to chartaceous, elliptic-oblong to oblong, broadest at or somewhat above
the middle, 9-23 x 3-9.3 cm, base attenuate, top acute-acuminate; upper surface
drying olivaceous-brown to blackish brown; lower surface early glabrescent,
without brown dots; midrib slightly to much raised above; nerves 9-13 pairs, flat to
moderately raised above, marginal arches not distinct; tertiary venation forming a
coarse network, usually faint to invisible on both surfaces; petioles 6-17 x 1.5-3.5
mm, glabrous; leaf bud slender to rather stout, c. 8-16 x 2-4 mm, densely greyish
brown to rusty-pubescent with hairs c. 0.1-0.3 mm. Inflorescences + thinly pubes-
cent with hairs c. 0.2-0.5 mm, sometimes subglabrescent, in d : 3-4 times ramified,
many-flowered, 4-22 x 2-12 cm, common peduncle c. 10-30 (-75) mm; in ? : 1-2
times ramified, c. 2-6 cm long; bracts oblong to lanceolate, pubescent, c. 2-4 (?)
mm, caducous. Flowers either predominantly 2-valved or in mixture of 2- and
3-valved, the 6 in rather dense clusters of c. 5-12, the 2 c. 1-4 together; perianths
glabrous; pedicel glabrous, in ¢ at base not articulated, in 2 articulated or not (see
notes). Male perianth broadly obovoid to globose or depressed globose, top broadly
rounded, base rounded to broadly rounded, outside faintly longitudinally ribbed or
176 Gard. Bull. Sing 37(2)(1984)
not, c. 1.0-1.5 (-1.7) x 1.2-2.0 mm; pedicel 0.4-1.0 (-1.5) mm, slender; perianth at
anthesis cleft to c. 1/2-way, valves 0.1-0.2mm thick. Androecium depressed-obovoid
to (depressed)globose, c. 0.6-1.0 x 0.6-1.1 mm, usually somewhat laterally flat-
tened and hence subcircular to elliptic (not circular) in transverse section; anthers 7-9
(-10), sessile, free apices 0-0.1 mm, towards the apex incurved over a rather narrow
central cavity c. 0.1-0.4 mm deep; androphore narrow, 0.1-0.2 mm long. Female
perianth obovoid-ellipsoid, 1.8-2.5 x 1.6-2.2 mm , glabrous, at anthesis cleft
to c. 1/3, valves 0.3-0.4 mm thick, ovary ellipsoid, 1.4-1.7 x 1.0-1.4 mm, glabrous,
stigma minutely 2-lobed, c. 0.1 mm, pedicel 1-1.5 mm long, glabrous or thinly
pubescent. Fruits 1-5 per infructescence, broadly ellipsoid, top and base broadly
rounded, 1.8-3.2 x 1.5-2.4 cm, finely granulate, not tuberculate, drying dark
brown, glabrous, dry pericarp c. 1.5-2.5 mm thick; stalk c. 1-2 mm; perianth not
persisting.
Distribution. Vietnam (Annam, Cochin-China), Laos, Cambodia, S. and SE.
Thailand (not in peninsula).
VIETNAM. Annam: Poilane 8699, 18160, 18308, 18615, 19887, 22394 — Cochin-China: Chevallier
39134, Pierre 14, 1812, 5434, Poilane (56), Thorels.n., 1186.
LAOS. Poilane 13480.
CAMBODIA. Hahn 140, Pierre 680, Poilane 23253, 23341, Vidal 5044.
THAILAND. Maxwell 74-794, 75-212, 76-158, 76-429, Smitinand & Phengklai 10870.
Ecology. Evergreen forest, regenerating forest; on rich red soil; 200-1100 m alt.
Flowers and fruits throughtout the year.
NOTES
1. Latex from bark colourless. Leaves subcoriaceous. Flowers yellow; fruits
greenish-yellow; aril thin, orange.
2. Female flowers have only been seen from two rather differeing collections,
viz.,(1) an unnumbered specimen collected by Poilane (10) (Indo-China), which
deviates somewhat in habit from the other specimens by its rather narrow lanceolate
leaves c. 18 x 4.5 cm, and slender inflorescences of c. 6 cm long with immature
flowers with the pedicels glabrous and articulated at the base; (2) Maxwell 75-212,
from Thailand, with stouter and more condensed inflorescences, with stouter
flowers of which the pedicels thinly pubescent towards the base and not articulated;
possibly the articulation of the pedicels in the Poilane specimen is artificial and
caused by the drying of the immature flowers.
3. H. thorelii is mainly characterized by the leaves being always dispersed in 3-5
rows, and by the very small globose male flowers c. 1.0-1.5 mm diam. It is closely
related to the polymorphous H. amygdalina, a species with a larger distributional
area and differing by having larger, male perianths (c. 1.5-2.3 mm long), generally
more anthers (8-15), and slightly larger fruits (2.2-3.4 cm long); it is also closely
related to H. /ongiflora, but the latter species differs by its larger and more elongate
flowers, smaller fruits, and generally distichous leaves.
4. Poilane 23252, 23341, and Vidal 5044, from Cambodia, Prov. Kampot, alt.
c. 200-300 m, are specimens deviating by the poor and short pubescence (hairs c. 0.1
mm) of the inflorescences; similar short-haired, glabrescent inflorescences have been
collected in in Thailand e.g., Maxwell 74-794.
5. H. thorelii seems to replace H. amygdalina in Annam and Cochin-China
(Central and S. Vietnam).
New account of Horsfieldia 7
6. In 1956 Sinclair (p. 422) regarded H. thorelii as a synonym of
H. amygdalina; in 1975 (p. 42) both these names were reduced to his wide concep-
tion of H. glabra.
5. Horsfieldia amygdalina (Wall.) Warb.
Myristica amygdalina Wall., Pl. As. Rar. 1, 4 (1830) 79, t. 90; Cat. (1832) No. 6797; Hook. f. & Th., Fl.
Ind. (1855) 160, p.p.; King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 300, pl. 128. — Horsfieldia
amygdalina (Wall.) Warb., Mon. Myrist. (1897) 310; Sincl., Gard. Bull. Sing. 16 (1958) 422 (in syn.
and in notes to H. bracteosa); C.Y. Wu (Ed.), Fl. Yunnan. 1 (1977) 12, fig. 3, 5-6 — Type: Wallich.
Cat. 6797 (KW; K, iso; BM: CAL, G, 7. v.).
Mpristica floribunda Wall., Cat. (1832) no. 6805, nom. nud.
M. kurzii King, nom. nud. (sub M. glabra auct. non Bl.: King). — Type: Kurz s.n., 984 (CAL, n.v.; iso,
12).
Horsfieldia tonkinensis Lecomte, Not. Syst. 1, 4 (1909) 100; FI. Gen. I.-Chine 5, 2 (1914) 101 — Type:
Bon 4272 (4302) (P).
H. tonkinensis var. multiracemosa Lecomte, Not. Syst. 1, 4 (1909) 100; Fl. Gén. I.-Chine 5, 2 (1914) 102
— Type: Bon s.n. (4302) (P).
Mpyristica glabra auct. non B\.: King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 310, pl. 142 — Horsfieldia
glabra auct. non (Bl.)- Warb.: Sinclair, Gard. Bull. Sing. 28 (1975) 35, p.p. (as for most of the
specimens originating from continental Asia); Fl. Rep. Pop. Sin. 30, 2 (1979) 204. fig. 93.
H. prunoides C. Y. Wu, Yunnan Econ. PI. (1973) 74, fig. 56, mom. nud. — Type: publication not seen;
name cited in the synonymy of H. glabra auct. in Flora of China, 1979.
Tree 5-30 m. Twigs terete, not ridged, towards the top 1.5-3.5 (-8) mm diam.,
bark grey-brown to brown, tomentum greyish to brown, composed of hairs c.
0.1-0.2 mm (var. amygdalina) or c. 0.5-1.0 mm long (var. /anata), usually tomentum
early glabrescent, lower down the bark rather finely striate, not flaking; lenticels
small, + conspicuous or not. Leaves either in 2, or 3, or 5 rows, or phyllotaxes
mixed, membranous to subchartaceous, elliptic-oblong to oblong, broadest at about
or slightiy above the middle, 9-23 x 2.5-7.5 (-9) cm, base attenuate, tip acute-
acuminate; upper surface drying olivaceous brown to dark brown; lower surface
early glabrescent, without brown or blackish dots; above, midrib flat to moderately
raised; nerves 7-14 pairs, thin, either sunken or flat, or moderately raised, marginal
arches not distinct; tertiary venation forming a rather lax network, usually faint to
invisible on both surfaces; petioles 10-20 x 1.5-3.5 mm, glabrous; leaf bud slender
to stoutish, densely grey-brown to brown, pubescent with hairs 0.1-0.2 or 0.5-1.0
mm (see under the varieties), c. 10-15 x 1.5-4 mm. Inflorescences either very thinly
pubescent with pale hairs c. 0.1 mm (var. amygdalina) to early glabrescent or early
glabrescent and hairs densely set, rusty, c. 0.5 (-1.0) mm (var. /anata), ind: 3-4 times
ramified, flower-number moderate to many, 6-18 x 3-12 cm, common peduncle
7-40 mm; in ? : 1-2 times ramified, 1-3 (-6) x 1-1.5 (-4) cm; bracts + elliptic, tip
rounded, pubescent, 1-2 (-3) mm long, caducous. Flowers either 2- or 3- (or 4-)
valved, or + an even mixture of 2- and 3-valved, in 6 rather dispersed or in loose
clusters of 4-10, in 2 solitary or 2-3 together, glabrous; pedicels glabrous, at base
not articulated, or in 9 sometimes indistinctly articulated. Male perianth shortly
obovoid or short-ellipsoid to globose, tip rounded, base rounded to + tapering, out-
side often shallowly longitudinally ribbed, c. 1.5-2.3 x 1.7-2.0 (-2.2) mm; pedicel
(0.8-) 1-2 mm; perianth at anthesis cleft to c. 1/2-way, valves c. 0.2 mm thick.
Androecium globose or depressed-globose, sometimes truncate-ellipsoid (see notes),
0.8-2.0 x 0.8-1.3 mm, slightly laterally flattened or not and hence transverse section
subcircular to ellipsoid; anthers 8-12 (-15), sessile, free apices 0-0.1 mm, towards
apex curved over an apical cavity moderately broad to narrow, c. 0.2-0.4 (-0.5) mm
deep; androphore rather narrow, c. 0.1-0.2 mm long; central column broad, solid.
Female perianth ellipsoid, c. 2.5-3.0 x 2.0 mm, glabrous, cleft at anthesis to c. 1/3,
178 Gard. Bull. Sing 37(2)(1984)
valves 0.4 (-0.5) mm thick; ovary ovoid, glabrous, 1.5-1.7 x 1.2-1.5 mm, stigma
minutely 2-lobed, c. 0.1 mm high; pedicel c. 1.0 mm long, glabrous, at base +
articulated or not. Fruits 2-6 (-16, see notes) per infructescence, ellipsoid, top and
base (narrowly) rounded, 2.2-3.4 x 1.6-2.6cm, glabrous, drying glaucous-brown to
dark brown, finely granulate, not tuberculate, dry pericarp 2-3 mm thick; stalk 2-3
mm long; perianth not persisting.
Distribution. India to Indo-China, and S. China, not in Malaya.
Note. This species, and the closely related H. thorelii and H. longiflora (see also
there) have similar fruits. These three are akin to H. glabra from West Malesia, the
latter distinctive by a number of minor characters, but always easily ascertained by
the presence of blackish dots on the lower leaf surface. H. subalpina from
mountainous West Malesia is also closely related.
KEY TO THE VARIETIES
la. Leaf bud, upper portion of twigs, immature inflorescences and immature leaves with rusty woolly
tomentumiwith)hairs!0/5-1O;mmilong ei-c-eccesesecees so eceeeenereteeeeeeceeee eee eee eee b. var. lanata
b. Tomentum with greyish to dull brown hairs 0.1-0.2 mm long ......................55+ a. var. amygdalina
a. var. amygdalina
Leaf bud, young upper portions of twigs and immature inflorescences with
tomentum of grey to dull brown hairs 0.1-0.2 mm long, or less. Mature Leaves
usually membranous. Flowers in the same inflorescence either 2- or 3-valved or 2-
and 3-valved mixed.
Distribution. India (Assam, Andaman Isl.), Bangladesh (E. Pakistan), Burma,
S. China, N. Thailand, Laos, N. and C. Vietnam (Tonkin, N. and C. Annam).
INDIA (incl. E. Pakistan): Hooker & Thomson (coll. 11/50) s.n.; Jenkins (comm. Anderson) s.7.;
King’s Coll. 268; Masters (coll. Simons) s.n.; Wallich 6804, 6805 (belonging to the material of M.
exaltata, not the lectotype); de Silva & Gomes, Wall. Cat. 6805.
Andaman Isl. (S. Andaman(: Balakrishnan 1033; Nair 843, 3641; King s.n., Kings Coll. s.n. (several
dates).
BURMA: Beddome s.n.; Dickason 6556-bis, 6945; Falconer (comm. Anderson) 5446; Helfer 4358;
Keenan, Tun Aung & Rule 1569; Kurz s.n., 984; Wallich 6797.
CHINA. Yunnan: Huang Yun Wui 120; also recorded S. Kwangsi, Hainan.
VIETNAM. Tonkin: Bon 2669, 4272, 4302; Petelot 1070; Poilane 13041; W.T. Tsang 29198 —C. &
N. Annam: Chevalier 38164; Poilane 11106, 11119.
LAOS: Poilane 13700.
THAILAND. North: Hansen, Seidenfaden, Smitinand 11181, Put 796; Winit 1471.
Ecology: Evergreen forest, recorded from sandy soil; 0-1000 m alt. Flowers and
fruits throughout the year.
NOTES
1. Flowers greenish-yellow or yellow, evil-smelling; fruits yellow.
2. Rather variable in shape and size of the inflorescence, and size of the male
flower, including the androecium.
i et
New account of Horsfieldia 179
Wallich 6805 (de Silva & Gomes), from Sylhet (Bangladesh) rather deviates by
its large, longish, male flower, the perianth measuring c. 2.2 x 2.2 mm, with the
androecium exceedingly elongate, c. 1.8-2.0 x 1.0 mm; this specimen resembles
H. longiflora, a species with the leaves apparently always distichous and with still
larger flowers, to c. 3 mm long, and with the androecium with broader and longer
tapering androphore, c. 0.3 mm long.
3. The specimens Poilane 11106, 11119 (from Annam), and Poilane 13700
(Laos) seem to have a completely distichous phyllotaxis. Of these collections Poilane
11119 somewhat deviates by its large infructescence bearing up to 16 (still immature)
fruits; in most specimens only up to 6 fruits have been seen. In Poilane 13700 the
hairs on the leaf bud are relatively long for the variety, c. 0,2 mm.
4. Quite often the flowers in a specimen are in different stages of development;
also fruits in the same infructescence are sometimes (e.g., Balakrishnan 1033) in dif-
ferent stages.
5. All specimens of both var. amygdalina and var. /anata were lumped by
Sinclair in his large conception of H. glabra.
b. var. lanata de Wilde, var. nov. Fig. 1A(5)
A var. amygdalina differt tomento lanuginoso rufo-ferrugineo in foliorum gemmis et inflorescentiis
juvenilibus valde distincto, pilis 0.5 (-1.0) mm longis — Type: Kerr 8556 (L; iso BM, K, P).
Leaf buds, immature twig apices, immature leaves (partly), and young
inflorescences with dense conspicuous rusty woolly tomentum composed of hairs 0.5
(-1.0) mm long. Mature leaves chartaceous. Male perianths either 2-, or 3- or
4-valved. Female flowers and fruits not seen.
Distribution. E. Thailand, Cambodia.
THAILAND (E.): Kerr 8439, 8556.
CAMBODIA: Pierre 1812, 5469.
Ecology. Evergreen forest; 0-200 m. Flowers in February.
Vernacular name. Liat-nok (E. Thailand).
Note. The specimens Pierre 1812 (P), 5469 (BM) from Cambodia slightly differ
from the plants from Thailand by the somewhat shorter tomentum with hairs c.
0.4-0.5 mm long, and by the predominantly 2-valved perianths.
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The Reproductive Biology of Rambutan, Nephelium lappaceum L.
(Sapindaceae)
LIM AH LAN
Department of Botany, University of Malaya, Kuala Lumpur, Malaysia
Abstract
Nephelium lappaceum produces either male or bisexual flowers. The anther development in both
types of flowers is the same but those in the bisexual flowers do not dehisce even though their pollen is
viable. The anther is tetrasporangiate and its wall development conforms to the basic type. Cytokinesis in
microspore mother cells is simultaneous, forming tetrahedral tetrads. The mature pollen grains are
tricolpate and two-celled. Only pollen grains from the male flower germinate in the conventional media.
The ovule is anacampylotropous, bitegmic, crassinucellate and the micropyle is formed by the inner
integument only. The embryo sac development is of the monosporic Polygonum type.
Flower anthesis is at 0900-1100 hours and pollination is entamophilous. Preliminary flower
bagging experiments show that fruit formation is dependent on pollination. The average normal flower
and fruit drop are 40% and 90% respectively.
The endosperm development is ab initio Nuclear and cell formation commences at the micropylar
end, proceeding towards the chalaza. The outer layers of the outer integument differentiate into the
edible flesh of the fruit. The testa is formed mainly from the inner part of the outer integument and the
few remaining layers of the inner integument at the micropylar region.
The seed is non-endospermous and shows hypogeal germination. In fresh seeds, the average percen-
tage germination is 96%, of which 2% of the seedlings give multiple shoots.
Introduction
In Malaya, several species of Nephelium produce edible fruits. Those common
in cultivation are the Rambutan (N. /Jappaceum L.), Pulasan (N. mutabile Bl.) and
Mata kucing (N. malaiense Griff.).
Nephelium lappaceum L. is known as Rambutan to the Malays because of its
hairy fruits; ‘‘rambut’ in the Malay language means ‘‘hair’’. Whitehead (1959)
recognised different clones based on size and taste of fruits, nature, colour and
texture of the soft spines of the rind. The Rambutan is delicious when eaten fresh
but it can also be canned or made into an excellent jam. A wine made from it was
exhibited at the Colonial and Indian Exhibition in London in 1886 (Burkill, 1935)
but the practice never seems to have caught on. The fruit wall, roots, leaves and bark
are said to have medicinal value.
Materials and Methods
Field observations on phenology, floral anthesis and pollination, and collection
of plant materials were undertaken at weekly intervals or more often as required, in
the fruit-tree nursery of the Ministry of Agriculture in Serdang about 20 km from
the University of Malaya campus. A voucher specimen KLU 1861/0 was deposited in
the Herbarium, Botany Department of the University of Malaya. The buds and
flowers were fixed in formalin-propionic-alcohol while the fruits were fixed in Craf
III. Routine methods of microtechnique were employed to obtain sections 8-15
thick. Fruits in wax blocks had to be immersed for periods up to a month in a soften-
ing solution of Molifex (B.D.H. Co., U.K.) before they could be sectioned. The
181
Fig. 1.
Floral morphology,
microsporangium, microsporogenesis and microgametophyte. (ed,
endothecium; ep, epidermis; isp, inner secondary parietal layer; mic, microspore mother cell;
ml, middle layers; osp, outer secondary parietal layer; sc, sporogenous cell; fa, tapetum).
A & B, bisexual flower; C, bisexual flower after pollination; D, pollen grains germinating on
stigma; E, male flower, F, t.s. tetrasporangiate anther; G-J, anther wall at different stages of
development; K-O, meiosis in microspore mother cells; P-7, microspore and a two-celled pollen
grain.
Reproduction biology of Nephelium lappaceum 183
mounted fruit sections had to be bleached in a solution made up of potassium
dichromate (1 gm), chromic acid (1 gm), glacial acetic acid (10c.c.) and distilled
water (90 c.c.) before staining in safranin. Otherwise the whole embryo sac stains
dark purple and this is caused by tannins and other cell inclusions, which mask
the cellular organisation. Various concentrations of ‘‘Clorox’’, a commercial
bleach, were tried but proved unsatisfactory.
To test pollen viability, the pollen grains from the bisexual and male
inflorescences were germinated in various concentrations (2-20%) of sucrose and
lactose solutions. The pollen grains from the male inflorescence were used
immediately after anthesis while those from the bisexual flowers were released by
cutting open mature anther sacs. The pollen grains were examined and pollen tubes
measured at 12-hour intervals.
Seed germination and seedling morphology were studied by sowing fresh seeds
in garden soil.
Observations and Results
Floral and Fruit Morphology
Flowers are borne on axillary or terminal panicles. The greenish flowers are
minute (less than 3 mm wide), without petals (fig 1A, B) and have a pleasant scent.
The trees bear either male or bisexual flowers (androdioecious). In a male flower,
there are 5-8 anthers supported by white hairy filaments while the gynoecium is
small and rudimentary (fig. 1E). In the bisexual flower, the anthers are 5-7 in
number and the gynoecium is very well developed with a 2- or rarely 3-locular ovary
(fig. 1A, B, C).
The bunched fruits are borne on woody stalks. Depending on the clone, each
fruit is oblong to nearly round and it ripens to a red or, less commonly, yellow
colour. The rind is covered with thick, soft spines (fig. 4D, F). The edible pulp is
white and the taste ranges from sour to sweet. Sour taste and difficulty in detaching
pulp from seed are considered poor qualities.
The non-endospermous seed is normally rounded at the micropylar end and
pointed at the opposite end. It has a fibrous testa enclosing an embryo with two
unequal cotyledons.
Phenology
Depending on the clone, as well as soil and climatic conditions, the tree starts to
bear flowers and fruits after 3-5 years. Two distinct flowering seasons are observed
in the Malay Peninsula. The first season starts at the beginning of April and the
fruits are harvested at the end of July. The second begins mid-August and the
harvest is mid-December. The seasonality of the fruits can easily be upset by any
change in the pattern of wet and dry season. Most individual trees, however, pro-
duce fruits only once yearly; some during the first season and others during the
second. Some trees in Serdang are peculiar in that while one side can be laden with
ripe fruits, the other half can be just beginning of flower.
After the appearance of the inflorescence, the floral buds (average width 1 mm)
take only 3 weeks to develop into mature flowers (average width 3 mm). They open
acropetally. Both male and bisexual plants flower synchronously and the flowering
period lasts 2-3 weeks. Anthesis is between 0900 and 1100 hours. Ten days after
anthesis, the stamens in the bisexual flowers drop off and usually one of the ovules
enlarges to form a young fruit (average width 4 mm) (fig. 3C). These fruits mature
12 weeks later and by then the average width is 3.5 cm and length 5.0 cm. The
Fig. 2. Megasporogenesis and development of megagametophyte. (ac, archesporial cell; ap, antipodal;
e.g., egg; ii, inner integument; o/, outer integument; pn, polar nucleus; pp, primary parietal cell;
sc, sporogenous cell; sy, synergid).
A & B, development of integuments; C, D, formation of sporogenous cell; E-H, megasporo-
genesis; J, twin tetrads; J, functional megaspore; K, twin megaspores; L-O, development of
megagametophyte; P, embryo-sac after the degeneration of antipodals.
observation of 10 samples of randomly tagged floral buds, 200 per sample, showed
that the average flower abortion was 40% and fruit drop 90%.
Anther Development
The anther development in both male and hermaphrodite flowers is found to be
similar except that dehiscence does not occur in the latter. Hence the bisexual
flowers are found to be functionally female. The anther is tetrasporangiate and the
124
——aaaaEOEOoOoo
|
H
|
|
Reproduction biology of Nephelium lappaceum 185
anther-wall development conforms to the basic type of Davis (1966). The ar-
chesporial cells divide periclinally to form the primary parietal and sporogenous
cells. The latter differentiate into microspore mother cells while the former divide to
form the outer and inner secondary parietal cells. The emdothecium and a middle
layer are derived from the outer secondary parietal while the inner middle layer and
the tapetum are derived from the inner secondary parietal (fig. 1G, H, I). The
tapetum is secretory and includes at first uninucleate cells which become binucleate
just before the microspore mother cells undergo meiosis.
Simultaneous cytokinesis accompanies meiotic divisions in the microspore
mother cell (fig. 1K-O). The tetrads formed are usually tetrahedral but sometimes
isobilateral. Stages of development varying from late prophase to microspore
tetrad phase had been observed within the same flower. In addition, different
locules of the same anther may exhibit different stages of development such as a
locule with microspore mother cells at metaphase I while another locule shows
microspore mother cells at anaphase II.
Soon after the tetrads are formed, the microspores separate out and at this
time, the tapetum and middle layers begin to degenerate (fig. 11). The nucleus
migrates to the periphery of the microspore, giving the latter the characteristic signet
ring shape (fig. 1P, Q). It then divides to form a small lenticular generative cell and a
large vegetative cell (fig. 1R). At maturity, the pollen grain is binucleate (fig. 1S, T).
By the time the microspores become mature pollen grains, the tapetum and the
middle layers have degenerated, leaving only the fibrous endothecium and the
epidermis in the anther wall (fig. 1J).
The mature pollen grains are smooth, triangular, tricolpate and measure
15-18 in diameter. Both male and bisexual flowers produce similar pollen grains.
However, most pollen grains in the functionally female flowers are devoid of nuclei
and hence are non-viable.
Pollen Germination
Pollen grains from functionally female flowers did not germinate in the conven-
tional media of sucrose or lactose solutions. Those from male flowers showed
maximum germination of 39% in 10% sucrose and 56% in 10% lactose solution.
The control sample gave 19% germination in distilled water. Further, at optimum
sugar concentrations (10% sucrose and 10% lactose), the pollen tubes were longer,
i.e., an average length of 0.1 mm while after 24 hours’ incubation the average pollen
tube length in distilled water was 0.02 mm. Though the pollen grain is tricolpate,
germination is monosiphonous.
Anthers from functionally female flowers were squashed to release the pollen
grains, which were then dusted on to the stigma of another flower either of the same
or a different tree. The inflorescences were bagged before and after the transference
of pollen grains to prevent contamination by pollen grains from other flowers. This
was performed on flowers in vivo because cut inflorescences tend to dehydrate very
fast. After 4 hours the flowers were harvested, fixed in polyvinyl lactophenol and
stained in lactophenol cotton blue. The nucleate pollen grains could be germinated
on the stigmatic tissue (fig. 1D). However, the percentage germination was not
calculated as the sample used was small.
Development in Ovule
Within each loculus of the ovary only one ovule, mounted on a thick funicle, is
present. The ovular primordium develops from the placenta. The differentiation of
the inner integument is followed by that of the outer (fig. 2A, B). Simultaneously
one of the hypodermal nucellar cells enlarges to form the archesporial cell (fig. 2C).
wonen cere.
Ad
ube
CT]
Se y
ab, A q
Fig. 3. Development of embryo, fleshy part of fruit, and testa. (ce, cellular endosperm; em, embryo; 7,
fleshy part of fruit; i/, inner integument; me, nuclear endosperm; o/, outer integument; po,
placental outgrowth; fe, testa; vb, vascular bundle).
A-C, |.s. flower showing ovule development; D-F, ovule development; G & H, seed differentia-
tion; J, globular embryo in seed; J, K, dicotyledonous embryo; L, anatomy of integuments at
mature embryo-sac stage; M, anatomy of integuments at globular embryo stage; N-P, differen-
tiation of testa and fleshy region of seed; O, anatomy of outer integument near micropyle of H.
The mature ovule is anacampylotropous, bitegmic, crassinucellate and the
micropyle is formed by the inner integument alone (fig. 3E). A placental outgrowth
is present at the chalazal end of the ovule (fig. 3F).
The archesporial cell divides periclinally, giving rise to the sporogenous cell and
the primary parietal cell (fig. 2C, D) which undergoes further divisions resulting in
186
Reproduction biology of Nephelium lappaceum 187
the crassinucellate ovule. As its cytoplasm becomes denser and the nucleus enlarges,
the sporogenous cell becomes the megaspore mother cell (fig. 2E). Following the
first meiotic division in the megaspore mother cell, a pair of dyad cells are formed,
of which the lower divides further to give a three-celled stage (fig. 2F, G). Later, the
smaller, upper, dyad cell undergoes a second meiotic division to give a linear tetrad
(fig. 2H). Within an ovule, usually only one embryo sac develops; however, excep-
tional cases of developing twin embryo sacs had been observed (fig. 21, 2K). At the
completion of meiosis, only the chalazal megaspore differentiates to form the
female gametophyte while the other three megaspores degenerate soon after forma-
tion (fig. 2J). After three successive nuclear divisions, the megaspore gives rise to the
mature eight-nucleate embryo sac (fig. 2L-N). Its development conforms to the
monosporic Polygonum type. The free nuclei are at first arranged in two groups of
four each at opposite poles of the embryo sac. One from each group migrates to the
centre (the two polar nuclei) and the remainder organises into the egg, flanked by
two synergids at the micropylar end, and by the three antipodals at the other end
(fig. 20). Soon after formation, the antipodals degenerate while the two polar nuclei
fuse to form a large secondary nucleus (fig. 2P).
Pollination and Fertilization
N. lappaceum is entomophilous. Though the individual flowers are small, less
than 4 mm wide, they are grouped into prominent panicles. Bees and butterflies are
attracted by the nectar and presumably also by the sweet fragrance of the flowers.
The visits of these insects had been observed to be most abundant during floral
anthesis i.e., between 0900-1100 hours.
Preliminary bagging experiments, before and after anthesis, were conducted to
determine whether fruit production is dependent on pollination. As the anthers in
bisexual flowers do not dehisce, the inflorescences were bagged without the removal
of anthers. The flowers are very small and several attempts to remove the anthers at
the bud stage fatally damaged the buds. Ten random samples, each of 200 young
floral buds (average diameter 1.0 mm), were bagged in butter-paper. All the flowers
in these samples dropped off and no fruits were formed. Ten other random samples,
each of 200 flowers (average diameter 3 mm), were bagged 5 days after anthesis.
These samples did produce fruits and the average percentage of fruit abortion was
92%, very near to the average fruit drop (90%) in separate phenology studies. This
shows that there is no significant difference between the fruit set from flowers
bagged after anthesis and that from unbagged ones. Bagging thus has no marked
deleterious effect on fruit development. In addition, flowers bagged before anthesis
could not have been pollinated whereas those bagged after anthesis could have been
pollinated by pollen grains from the male trees nearby. This strongly suggests that
fruit production is dependent on fertilization or at least pollination stimulus.
Though actual pollen-tube growth and fertilization were not seen, the degenera-
tion of one synergid occurs soon after the fusion of the polar nuclei. Later, only the
egg and the secondary nucleus remain.
Endosperm
The development of the endosperm is of the ab initio Nuclear type. Just before
the primary endosperm nucleus divides, the embryo sac enlarges (fig. 3F). The
primary endosperm nucleus undergoes several mitoses and the resulting free nuclei
Occupy mainly the micropylar region and a position around the periphery of the
enlarging embryo sac (fig. 4A). Corresponding to the increase in the number of
endosperm nuclei, the embryo sac increases in size at the expense of the nucellus
and later of the inner integument. When the embryo sac is about 2 mm long, there
are only 3-4 layers of the inner integument at the sides but more at the micropylar
——
ZG\X Ss 2
ZLY NS
Fig. 4.
Embryo, fruit development and seed germination. (em, embryo; f/, fleshy part of fruit; //, inner
integument; me, nuclear endosperm; sd, seed; zy, zygote).
A & B, nuclear endosperm; C, degenerating inner integument cells; D, fruit development; E,
seed germination, F, |.s. fruit; G, multiple-shoot seedling.
and chalazal regions (fig. 3H, M).
At the chalazal end of the embryo sac, a few cells of the inner integument
enlarge to twice or thrice their normal size (fig. 4C). Some of these cells degenerate
and release into the embryo sac the contents, which stain darkly with safranin and
fast green. The contents are a mixture of tannin, resin and oil. When the embryo sac
is 5 mm long, cell formation in the coenocytic endosperm commences at the
periphery of the micropylar region and proceeds towards the chalazal end. At the
young, globular embryo-stage (+ 0.1 mm in width), only a quarter of the embryo sac
is filled with cellular endosperm (fig. 31). Most of the endosperm has become
188
)
Reproduction biology of Nephelium lappaceum 189
cellular when the embryo is dicotyledonous (fig. 3J), but it is completely absorbed
when the embryo matures (fig. 3K).
Embryo
The division of the zygote occurs much later, compared with the formation of
the nuclear endosperm. The earliest stage observed is a small globular embryo, 0.02
mm wide, surrounded by numerous endosperm nuclei (fig. 4B). The globular
embryo increases in size and later differentiates into a heart-shaped embryo which
then forms the mature straight embryo with two unequal cotyledons (fig. 3I-K).
Degeneration of the embryo sacs was frequently observed. Often, the fruits in-
creased in size while the embryo sacs remained small and eventually degenerated.
Seed Coat and Flesh of the Seed
The edible flesh of the fruit is derived from the outer integument while the testa
is derived from both the outer integument and the remaining inner integument. The
inner integument is initiated around the ovular primordium at the archesporial cell
stage (fig. 2A), followed by the development of the outer integument (fig. 2B). Both
integuments are already prominent at the megaspore mother-cell stage (fig. 3A, D).
At the mature 8-nucleate embryo sac stage, the inner and outer integuments are
respectively, 6-7 layers and 5-6 layers thick. The subepidermal cells of the outer in-
tegument near the chalaza and placental outgrowth have divided such that it is 10-12
cells thick (fig. 3B, E). The vasculature to the integuments is prominent and abun-
dant.
After the degeneration of the antipodals and just before the division of the
primary endosperm nucleus, meristematic activity spreads to the innermost cells on
the lateral sides of the outer integument (fig. 3F, L). All these cells divide
periclinally, hence they are arranged in rows which fan out radially. Simultaneously,
cells of the inner integument also divide in that it becomes 10-13 layers thick.
Periclinal meristematic activity of the outer integument continues from the chalazal
to the micropylar region of the seed during nuclear endosperm formation (fig. 3G).
The epidermis of the outer integument, so far, has only undergone anticlinal divi-
sions.
Before the zygote divides, when the seed is about 1 mm long, the outer integu-
ment is 15-30 cells thick. The embryo sac has enlarged at the expense of the nucellus
and the inner integument and by then the nucellus has been completely absorbed.
The outermost cells of the inner integument elongate while those next to the embryo
sac degenerate as this increases in size.
When the seed is about 5 mm long, the inner integument is only 3-4 cells thick
along the periphery of the embryo sac though it is much thicker at the chalazal and
micropylar ends. Meanwhile, rapid cell divisions, which may produce irregular
folds, occur at the micropylar end of the outer integument (fig. 3H, Q). The epider-
mis of the outer integument now divides both anticlinally and periclinally. These
cells contain tanniniferous materials. They then enlarge, become turgid with a juicy
fluid and the tannin content disappears during ripening.
The seed continues to grow at the expense of the inner integument which is now
only 2-3 layers along the wall of the embryo sac, and 5-6 layers at the micropylar
end. When the seed measures to 6-8 mm long (at the globular embryo stage) the cells
of the outer integument differentiate into an outer region of thin-walled
parenchymatous cells and an inner region of sclerenchyma cells, traversed by
vascular tissues (fig. 31). The outer 6-7 layers of cells increase in size and differen-
tiate into the fleshy part of the fruit (fig. 3M-P). The inner layers of the outer integu-
ment and the remaining inner integument become compressed to form the long
190 Gard. Bull. Sing 37(2)(1984)
fibres, tanniniferous cells and vasculature of the testa (fig. 3P).
The placental outgrowth of the ovule, very prominent at the mature embryo-sac
stage, contributes neither to the formation of the testa nor to the fleshy part of the
fruit.
Seed Germination
Fresh seeds take only 7-10 days to germinate and germination is hypogeal, in
which the hypocotyl is undeveloped and the cotyledons remain within the testa. The
first sign of growth is detected when the cotyledons split and the radicle emerges,
growing downwards, while the plumular shoot emerges above the soil level (fig. 4E).
It is recurved at first, later growing upright. Two axillary shoots develop from the
base of the shoot system but these die off one month after germination. When the
seedlings are about 2 weeks oldi.e., when the stem is about 8 cm long, the first pair
of leaves unfold.
The average percentage of germination of 200 fresh seeds sown in garden soil is
96%. Of the seedlings, 2% possess 2 or 3 shoots which emerge from the same point
of the seed and they share one root system (fig. 4G). Probably more than one shoot
apex are in the embryo.
Discussion
Nephelium lappaceum is a common, indigenous Malaysian fruit tree with great
economic potential. The external morphology has been described by Corner (1952)
and Allen (1967).
The plants are functionally dioecious and are either male or bisexual. The
stamens in the bisexual flowers have been called ‘staminodes’ (Ochse, 1961). The
term seems inappropriate since the present study shows that though the anthers do
not dehisce, they do produce viable two-celled pollen grains which could germinate
on a living stigma.
Corner (1952) reports that in Nephelium, the relation between the male and
bisexual trees is not known. He states ‘‘it seems therefore that the male trees are
useless though it is possible that the bisexual flowers must be cross-pollinated from
the male trees to set fruit’’. This present study on the bagging of the flowers before
and after anthesis strongly suggests that fruit production in N. /appaceum is indeed
dependent on fertilization or at least pollination. Self pollination cannot occur as the
anthers of bisexual flowers are not able to release its viable pollen grains. This seems
an effective barrier against inbreeding of the species.
Heterostyly is more elaborate in Cardiospermum halicacabum (Nair and
Joseph, 1960) and Litchi chinensis (Banerji and Chaudhuri, 1944). Both bear 3 types
of flowers, which differ from each other mainly in the degree of sexual develop-
ment. These flower types appear consecutively on the same panicle and are
designated as Type I, II and III according to the chronological order of the develop-
ment (Mustard, 1960). Type I functions as male since the gynoecium is rudimentary,
Type II as female as the androecium is underdeveloped and Type III, an in-
termediate of the two, normally functions as male. The only difference between N.
lappaceum and L. chinensis is that in N. lappaceum, the 2 types of flowers are borne
on different trees. Yet Whitehead reports (1959) that ‘‘the cultivated rambutan is
usually monoecious, flowers of both sexes being borne on the same inflorescence’.
A large proportion of the flowers and fruits of the Rambutan degenerate during
development. Khan (1939) and Mustard (1960) also noted such high percentages of
flower and fruit abortion in L. chinensis. Khan (1939) reported that only a very
Reproduction biology of Nephelium lappaceum 191
small ratio (1:78) of female flowers set fruit. Mustard (1960) was able to reduce
total degeneration of embryo sac from 63.1% to 41.0% by partial defloration. The
decrease in degeneration in partially deflorated as compared to non-deflorated
panicles indicates a competition among flowers. She suggested that water and
nutrient deficiencies may be contributing factors, not only to the degeneration of
megagametophytes but also the problem of fruit set in L. chinensis.
Studies on the embryology of the members in the Sapindaceae are scanty and
incomplete except in L. chinensis and Cardiospermum halicacabum. This present
study shows that the anther of N. /appaceum is tetrasporangiate and the wall
development conforms to the basic type. In the functionally female flowers,
degeneration of the pollen grain is common at the various stages of development.
The ovule is bitegmic and crassinucellate and the embryo sac development con-
forms to the monosporic Polygonum type as in other species in the family already
studied (Davis, 1966).
The details of embryogeny in N. /appaceum are masked by the dense, free-
nuclear endosperm and the darkly staining resinous and tanniniferous content of the
embryo sac. Within the family the embryogeny of C. halicacabum only has been in-
vestigated and reported as the Asterad type by Nair and Joseph (1960) but as the
Onagrad type by Kadry (1946).
The fleshy pulp of the fruit in Sapindaceae has generated much controversy
over the years. Radlkofer (1933) said that the edible fleshy part of N. lappaceum is
an aril which adnates to the testa. Van der Pijl (1957) and this study show that the
fleshy layer is the swollen outer integument and the dry, protective seed coat
originates from the innermost layers of the outer integument and the inner integu-
ment. In contrast, the edible, fleshy layer of Nephelium longana and L. chinensis are
considered as a free aril (Radlkofer, 1933) or an arillode (Van der Pijl, 1957). The
differentiation of the flesh around the micropyle observed here is the same as the
exotestal patch described by Corner (1976).
Acknowledgements
I express my sincere thanks to the Vice-Chancellor of the University of Malaya
for research facilities and grant which made this study possible. I am grateful to
Dr. N. Prakash, University of New England, Australia, for advice and comments.
Literature Cited
Allen, B. M. (1967). Malayan Fruits. Donaid Moore Press Ltd., Singapore.
Banerji, I. and K. L. Chaudhuri (1944). A contribution to the life history of Litchi
chinensis Sonn. Proc. Indian Acad. Sci. B 19: 19-27.
Burkill, I.H. (1935). A Dictionary of the Economic Products of the Malay Penin-
sula. Vol. Il. Crown Agents, London.
Corner, E. J. H. (1952). Wayside Trees of Malaya. Vol. 1. Government Printing
Office, Singapore.
. (1976). The seeds of Dicotyledons. Vols. 1 & 2. Cambridge University
Press, Cambridge, London.
Davis, G. L. (1966). Systematic embryology of the Angiosperms. John Wiley
and Sons Inc., New York, London.
Kadry, A. E. R. (1946). Embryology of Cardiospermum halicacabum L. Svensk
bot. Tidskr. 40: 111-126.
192 Gard. Bull. Sing 37(2)(1984)
Khan, A. R. (1939). Pollination and fruit formation in Litchi (Nephelium litchi
Camb.). Agric. J. India. 24: 183-187.
Mauritzon, J. (1936). Zur Embryologie und Systematischen Abgrenzung der Reihen
Terebinthales und Celastrales. Bot. Notiser 1936: 161-212.
Mustard, M. J. (1960). Megagametophytes of the Lychee (Litchi chinensis Sonn.)
Amer. Soc. Hort. Sci. 75: 292-304.
Nair, N. C. and T. Joseph (1960). Morphology and Embryology of Cardiospermum
halicacabum Linn. J. Indian bot. Soc. 39: 176-194.
Ochse, J. et. al. (1961). Tropical and Subtropical Agriculture. Vols. 1 & 2. Mac-
millan Press, New York.
Pijl, L. Van der (1957). On the arillodes of Nephelium, Euphoria, Litchi and
Aesculus and the seeds of Sapindaceae in general. Acta bot. Neerl. 6: 618-641.
Radlkofer, L. (1933). Sapindaceae. In Engler’s ‘Das Pflanzenreich’ IV: 165.
Whitehead, C. (1959). The Rambutan. A description of the characteristics and
potential of the more important varieties. Mal. Agric. J. 42 (2): 53.
A Revision of Rennellia (Rubiaceae) in the Malay Peninsula
WONG KHOON MENG
Forest Research Institute, Kepong, Malaysia
EFFECTIVE-PUBLICATION DATE: 7 MAR. 1985
Abstract
Rennellia (Rubiaceae) is revised for the Malay Peninsula. Four species are recognised, viz., R.
elongata (K. & G.) Ridl., R. paniculata K. & G., R. speciosa Hk. f. and a fourth, which resembles, ‘but
cannot be matched with certainty to, R. morindiformis (Korth.) Ridl. A key to the species and varieties is
provided, and the taxa are enumerated with descriptions. R. paniculata var. condensa Wong is newly
described.
Introduction
Rennellia Korth. is a small genus native to SE. Asia, with seven recorded
species. In the Malay Peninsula, it is represented by four species. Rennellia is most
closely related to Morinda L.; the two genera belong to the tribe Morindeae and
share the following characters: inflorescences terminal in origin; flowers
4-5-merous, connate by their calyx tubes and arranged in non-involucrate heads;
corolla aestivation valvate; ovary 2-celled, each cell with one ovule; fruits
syncarpous.
In the Malay Peninsula, Rennellia can be distinguished from Morinda by the
following key:
Shrubs, trees or lianes; flowering heads individually stalked, arranged in clusters of a few or as a
pseudoumbel (but never arranged along a main rachis); flowers many (more than 6) in each flowering
head; ovules inserted sub-basally in each locule ....................20e0ceceeeeee RRO 3 ce acewteasece Morinda L.
Shrubs or trees, never lianes; flowering heads sessile or individually stalked, arranged in clusters along a
main rachis; flowers few (3-6) in each flowering head; ovules inserted on the septum in each locule
Bae e ete oiireiiche vate's wo slots oisuienieceiente anieisiset vo cicat via ctdevianetisiseslev eve cass scla swleviocdiessectlassinesecweceree sss Rennellia Korth.
Rennellia was first recorded from the Malay Peninsula by Hooker (in Bentham
& Hooker, 1873), who recognised that a plant listed by Wallich as Morinda speciosa
(Wall. Cat. 8436, from Tenasserim in Burma) was identical with a species with
several collections from the Malay Peninsula, and that this species in fact belonged
to Rennellia. Subsequently, Hooker (1880) formally described it as R. speciosa, and
listed it as occurring in the Malay Peninsula. Later, King & Gamble (1904) described
a second species (R. paniculata K. & G.) for the Malay Peninsula and Ridley (1939)
elevated the status of R. speciosa var. elongata K. & G. to that of species,
R. elongata (K. & G.) Ridl.
However, because the keys presented in the accounts of King & Gamble (1904)
and Ridley (1923; 1939) rely mainly on the characters of habit and inflorescence
structure, the concept of R. speciosa came to be based on a mixture of species, and
thus the keys were difficult to use.
A fourth species represented by one specimen is here designated only as
Rennellia sp.; the difficulty in naming it precisely is discussed under the species
enumeration.
193
194 Gard. Bull. Sing 37(2)(1984)
Characters Useful to the Taxonomy of Rennellia
All four species in the Malay Peninsula have a habit ranging from shrub to
treelet. There is variation between species in the inflorescence structure, the number
of flowers in each flowering head, floral size and leaf venation.
Fig. 1. Inflorescence structure in Rennellia.
A, R. paniculata var. paniculata (3 flowers per head). B, R. paniculata var. condensa (3-6
flowers per head). C, R. speciosa (3-6 flowers per head). D, Rennellia sp. (3-6 flowers per
head). E, R. elongata (3 flowers per head). r: inflorescence rachis.
Plate 1. Rennellia elongata in flower (FRI 32013)
Rennellia in the Malay Peninsula 195
The flowering heads are individually stalked in R. paniculata var. paniculata,
R. speciosa and the species here designated as Rennellia sp., but are sessile on the
rachis in R. elongata and R. paniculata var. condensa. The heads are arranged
loosely in clusters, recognizable as tiers along the rachis (Fig. 1).
The number of flowers within each flowering head is consistent, 3 for
R. elongata and R. paniculata var. paniculata and 3-6 for R. paniculata var. conden-
sa, R. speciosa and the unnamed Rennellia species.
Floral dimensions distinguish the different species reliably. In R. paniculata the
corolla tube is 6-12 mm long, with lobes 4-9 mm long, while in the unnamed
Rennellia species the tube is 9-15 mm long with lobes 9-11 mm long. In R. elongata
and R. speciosa, the corolla tube length is 15 mm or longer, the lobes exceeding half
its length in R. elongata and not half as long in R. speciosa.
The species are functionally dioecious. In male flowers, the anthers are normal-
ly formed and protrude slightly from the corolla throat while the style is reduced,
hardly reaching half the length of the corolla tube. In female flowers, the anthers are
smaller, empty, and are completely included within the corolla tube, while the style
is slightly exserted from the corolla throat and bears a bifid stigma.
For leaf venation, R. paniculata has tertiary veins that are distinctly raised and
prominent as a dense network on the leaf undersurface. In R. elongata, R. speciosa
and the unnamed Rennellia species, the leaf undersurfaces have sparse tertiary veins
that are hardly distinct.
Key to the Species and Varieties of Rennellia in the Malay Peninsula
Leaf undersurfaces with a dense network of prominently raised tertiary veins; secondary veins 11-13 pairs
Inflorescence rachis 6-9 cm long, flowering heads with stalks 2-3 cm long, with 3 flowers per head,
arranged as 2-3 tiers along the inflorescence rachis ............... Rennellia paniculata var paniculata
Inflorescence rachis 1-3 cm long; flowering heads sessile, with 3-6 flowers each, arranged as 1-3 tiers
ALGMIPAENIE ITTIOTESCONCE LACINS (92-255. c282 se acececeteecesrcocceadeee-neee Rennellia paniculata var. condensa
Leaf undersurfaces with sparse, hardly distinct tertiary veins; secondary veins 6-11 pairs
Inflorescence rachis 1.5-3.5 cm long; flowering heads stalked, with 3-6 flowers each
Flowering heads with stalks less than 1 cm long, arranged as 1-4 tiers along the inflorescence
rachis; corolla with tube 15-23 mm long, the lobes less than half the tube length
a Ne ee NN Se oe ea aera casio owonssendes sovwcsontecoeedeseseves Rennellia speciosa
Flowering heads with stalks 1-3 cm long, arranged as 2-3 tiers along the inflorescence rachis;
corolla with tube 9-15 mm long, the lobes more than half the tube length ............ Rennellia sp.
Inflorescence rachis 8-15 cm long: flowering heads sessile, with 3 flowers each, (flowering heads
arranged as 2-12 tiers along the inflorescence rachis) ................20ceeeeeeeeeeeeees Rennellia elongata
Systematic Enumeration of Taxa
In the following enumeration, a list of specimens examined for each taxon is
provided wherein male and female specimens are marked ¢ and 9, respectively and
specimens with flowers too immature for sex determination are asterisked.
1. Rennellia elongata (King & Gamble) Ridley, Kew Bull.: 608 (1939) Plate 1
Basionym: R. speciosa var. elongata K. & G., J. As. Soc. Beng. 73: 90 (1904);
Ridley, Fl. Malay Pen. 2: 120 (1923).
Synonym: R. speciosa sensu Corner, pro parte, Wayside Trees of Malaya 1:
558 (1952).
Lectotype (here chosen): Ridley 5834, Pahang, Tahan River (SING!).
196 Gard. Bull. Sing 37(2)(1984)
Distribution & Ecology. Sumatra, Malay Peninsula, Borneo (Sarawak).
Lowlands to montane forest above 1200 m alt.
Shrub or treelet to 5 m tall. Leaves obovate to elliptic, (3-14) x (10-32) cm, with
6-10 pairs of secondary veins and sparse, hardly distinct tertiary veins. Inflorescence
rachis 8-15 cm long, with 2-12 tiers of sessile flowering heads. Each flowering head
with 3 flowers. Mature open flowers with corolla tube 15-19 mm long, and corolla
lobes 10-15 mm long.
Specimens examined
PERAK: Curtis s.n. 25. xii. 1901, Larut Hill (SING!, 2 ); Sow KEP 47232, Sg. Penuh (KEP!, 9).
SELANGOR: Ahmad KEP 94290, Ulu Gombak (KEP!, K,°); Gadoh & Millard K.L. 494, Ulu Langat
(KEP!2); Gadoh & Millard K.L. 2247, Ulu Langat* (KEP!); Hume 8980, Genting Simpah* (SING!);
Stone 5986, Fraser Hill (SING!, 6 ); Whitmore FRI 15721, Ulu Gombak (KEP!, 2 ); Wong FRI 32013,
KEP F.R. grds (KEP!, 9); Wong, s.n., 4. ili. 1981, Bt. Lagong (KEP!, 6 ); Wyatt-Smith KEP 60635,
Bt. Lagong (KEP! 6 ); Wyatt-Smith KEP 79199, Ulu Gombak (KEP!, 6 ). NEGRI SEMBILAN: Everett
KEP 104912, Jelebu (KEP!, K, 9). KELANTAN: Whitmore FRI 4081, Sg. Nenggiri nr Kg. Jenera
(KEP!, K, 2 ). TRENGGANU: Moysey & Kiah SFN 33827, Ulu Brang (SING!, 2 ). PAHANG: Ang FRI
23312, Taman Negara (KEP!, K,?); Henderson FMS Mus. 10548, Temerloh, Titi Bungor (SING!, 9);
Mad. Shah MS 1517, K. Kenyam (KEP!, K, 6); Md. Shah & Md. Noor MS 1768, Ulu Sg. Sat nr Kelepah
(KEP!, 2 ); Ng FRI 27272, Tekam F.R. (KEP!, 92); Ng & Beltran FRI 6447, Jerantut, Ulu Tekam
(KEP!, K, 9 ); Ridley 5834, Tahan River (SING!, 2 ); Ridley, s.n., 1891, Tahan (SING!, 2 ). JOHORE:
Corner, s.n. 9. ix. 1934, Mawai (SING!, d); Vethevelu FRI 25287, Kota Tinggi (KEP!, K,2); Whit-
more FRI 8764, NW. G. Blumut (KEP!, K, ¢ ).
2(a) Rennellia paniculata King & Gamble, var. paniculata.
J. As. Soc. Beng. 73: 89 (1904); Ridley, Fl. Malay Pen. 2: 119 (1923).
Synonym: R. speciosa sensu Corner, pro parte, Wayside Trees of Malaya 1:
558 (1952).
Lectotype (here chosen): King’s Coll. 2592, Perak, Larut, 3000-4000 ft. (K!).
Distribution & Ecology. Malay Peninsula (endemic). Montane forest at about
1000-1300 m alt.
Shrub or treelet to 8 m tall. Leaves obovate to elliptic, (4-9) x (10-22) cm, with
11-13 pairs of secondary veins and a dense network of distinctly raised tertiary veins
on the undersurface. Inflorescence rachis 6-9 cm long, with 2-3 tiers of stalked
flowering heads (the stalks 2-3 cm long). Each flowering head with 3 flowers. In
mature open flowers, corolla tube 6-12 mm long, and corolla lobes 4-9 mm long.
Specimens examined
PERAK: King’s Coll. 2164, Larut, 3000-3500 ft. (K! 9); King’s Coll. 2592, Larut, 3000-4000 ft.*
(K!); King’s Coll. 5432, Larut, 3000-3500 ft. (SING! K 9).
(b) Rennellia paniculata var. condensa Wong, var. nov.
Synonym: R. speciosa sensu K. & G., pro parte, J. As. Soc. Beng. 73: 89
(1904); sensu Ridley, pro parte, Fl. Malay Pen. 2: 120 (1923).
Varietas affinis Rennellia paniculata vat. paniculata sed inflorescentia 1-3 cm longis, capitulis
floriferis sessilis, 3-6 floribus, in 1-3-fasciculatis disposita differt.
Holotype: Curtis, s.n., Dec. 1895, Perak, Bujong Malacca (SING!).
Distribution & Ecology. Malay Peninsula (endemic). Lowlands to hill forest at
about 300 m alt.
Shrub ortreelet to 5 m tall. Leaves obovate to elliptic, (4-10) x (16-32) cm,
with 11-13 pairs of secondary veins and a dense network of distinctly raised tertiary
Rennellia in the Malay Peninsula 197
veins on the undersurface. Inflorescence rachis 1-3 cm long, with 1-3 tiers of sessile
heads. Each flowering head with 3-6 flowers. Mature open flowers with corolla tube
6-12 mm long, and corolla lobes 4-9 mm long.
Specimens examined
KEDAH, Kochummen FRI 2021, G. Inas F.R. (KEP! K, 2). PENANG, Abrams s.n. vi. 1890
(SING! 2). PERAK, Curtis 3345, Ipoh (SING!, 2); Curtis s.n. xii. 1895, Bujong Malacca (SING! 9);
Haniff & Nur SFN 6985, Sg. Siput (SING!, 2); Hashim KEP 9663, Ijok F.R. (SING!, 2°); Ng FRI 6083,
Bubu F.R., Sg. Wang (KEP!, 9); Ogata KEP 110224, Tapah, Changkat Jong F.R.* (KEP!); Wray 2897
(SING!, 2); Wray 4008, Larut, Relau Tuior (SING! c ).
3. Rennellia speciosa Hk. f., Fl. Brit. Ind. 3: 158 (1880); King & Gamble, pro
parte, J. As. Soc. Beng. 73: 89 (1904); Ridley, pro parte, Fl. Malay Pen. 2: 120
(1923); Corner, pro parte, Wayside Trees of Malaya 1: 558 (1952).
Holotype: Wallich, Cat. 8436, Burma, Tenasserim, Chappadong Hill (K!).
Distribution & Ecology. Burma, Thailand, Malay Peninsula, Borneo
(Sarawak). Lowlands to hill forest at about 500 m alt.
Shrub or treelet to 7 m tall. Leaves obovate to oblanceolate to elliptic, (3-12) x
(10-30) cm, with 8-11 pairs of secondary veins and sparse, hardly distinct tertiary
veins. Inflorescence rachis 1.5-3.5 cm long, with 1-4 tiers of stalked flowering heads
(the stalks less than 1 cm long). Each flowering head with 3-6 flowers. Mature open
flowers with corolla tube 15-23 mm long, and corolla lobes 6-7 mm long.
Specimens examined
LOWER THAILAND: Curtis 3000, Poongah (SING!, 2). KELANTAN: Cockburn FRI 7187, Ulu
Sg. Aring (KEP!, K, 2); Cockburn FRI 7462, Ulu Kelantan, Sg. Jenal (KEP!, K, 2); Md. Shah &
Ahmad MS 3211, Machang, Bt. Baka, Sg. Jeram Tinggi* (KEP!, SING!); TRENGGANU: Corner s.n.,
31.x. 1935, Kemaman, Bt. Kajang (SING!, 2); Md. Shah & Samsuri MS 3549, Jerteh, Bt. Yong, Sg. Tok
Barak* (KEP!, SING!); Md. Shah & Samsuri MS 3805, path to Bt. Bongkok, via Kg. La* (KEP!,
SING!); PAHANG: Md. Shah MS 1315, K. Tahan (KEP!, SING!, K, 2); Md. Shah MS 1348, Sg. Teku
(KEP!, SING! 2); Whitmore FRI 4805, K. Teku (KEP! K, 6 ); Wong & Wyatt-Smith W 47, Ascent G.
Tahan (KEP!, 2). MALACA: Alvins 562* (SING!); Alvins, 710, Selandar* (SING!) Alvins 2038,
Merlimau (SING!, 9); Al/vins s.n. 18.iv. 1886, Merlimau (SING!, 2 ); A/vins 210], Chabau (SING!, 2);
Alvins s.n. 9. ii. 1886* (SING!); Hervey s.n., 1891 (SING!, 9); Ridley s.n., 1891, Ayer Panas (SING!,9);
JOHORE: Corner SFN 29030, 14 m. Mawai-Jemaluang Rd (SING!, K, 2); Fox s.n., 1902, Muar
(SING!, 2); Hardial & Samsuri H.S. 1008, Datok Sawah* (SING!); Kiah SFN 32309, Sg. Kayu (KEP!,
?).
4. _Rennellia sp.
Shrub or treelet to 6 m tall. Leaves obovate to elliptic, (4-9) x (6-22) cm, with
8-10 pairs of secondary veins and sparse, fine tertiary veins. Inflorescence rachis
1.5-3 cm long, with 2-3 tiers of stalked flowering heads (the stalks 1-3 cm long).
Each flowering head with 3-6 flowers. Mature open flowers with corolla tube 9-15
mm long and corolla lobes 9-11 mm long.
Specimen examined
PAHANG: Purseglove P. 4311, Fraser’s Hill, Ring Road, 4000 ft. alt. (K! SING! @).
This species, represented only by one specimen in the Malay Peninsula, is
possibly the same as R. morindiformis (Korth.) Ridl. (Ridley, 1939), typified by the
specimen Korthals, s.n., from Mt. Singalang in Sumatra (holotype at Leiden,
isotype at Kew).
The Korthals specimen which typifies R. morindiformis has flowers represented
only in the bud stage and it is not possible to ascertain the dimensions of mature
198 Gard. Bull. Sing 37(2)(1984)
flowers without further collections of this species from the type locality in Sumatra.
The Purseglove specimen from the Malay Peninsula (where it is the only collection
of its kind) has mature flowers and matches the Korthals specimen in vegetative and
inflorescence structure. One disparity is the labelling of the Purseglove specimen as a
treelet and the labelling of the Korthals specimen as ‘‘pseudo-parasiticus.’’
However, the habit of species of Rennellia ranges from shrublets to treelets and I
have personally observed R. e/ongata to grow from hollows of tree boles as well as
to stand independently as treelets; the habit may therefore be confusing to the collec-
tor at times. I am not willing to name the Purseglove specimen as a distinct species as
present evidence allows this to be matched to R. morindiformis. Still, it is worthy to
remember that the structure of mature flowers of R. morindiformis can only be
elucidated with further collections from its type locality. Purseglove’s label on his
specimen identifies it as ‘‘R. longiflora Ridl.’’ but that is an unpublished name.
Acknowledgements
I wish to thank the Keepers of the herbaria at Bogor, Kew and Singapore who
have generously made available photographs of reference specimens end herbarium
loans for study. I am grateful to Diane Bridson at Kew, and Ruth Kiew at the
Agricultural University of Malaysia for their criticisms.
Literature Cited
Bentham, G. & J. D. Hooker (1873). Genera Plantarum 2: 118.
Hooker, J. D. (1880). The Flora of British India 3: 158.
King, G. & J. S. Gamble (1904). Materials for a Flora of the Malayan Peninsula.
J. As. Soc. Beng. 73: 89-90.
Ridley, H. N. (1923). Flora of the Malay Peninsula 2: 119-120.
. (1939). Notes on some Malayan Rubiaceae. Morindeae. Kew Bull.: 600-611.
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Keys to the Cultivars of Keladi (Colocasia esculenta — Araceae)
in Peninsular Malaysia
FARAH D.GHANI
Botany Department, National University of Malaysia
Bangi, Selangor, Malaysia
Abstract
Colocasia esculenta (Araceae), the Keladi or taro, is an ancient staple food crop of the Asian and
Oceanic Pacific populations. In Peninsular Malaysia, collections have reached 50 cultivars but only two
of them are commercially cultivated. Confusion exists in the identification of these cultivars, which is
attributed to loose usage of vernacular names. In this study a key to these cultivars is presented, one
which has been worked out from plants grown in sifu and replanted in experimental plots. Only stable
morphological and behavioural characteristics have been used in differentiating the cultivars.
Introduction
The Araceae, a family of about 115 genera and over 2000 species, is of worldwide
distribution, but 92% of the species are native to the Asiatic and American tropics.
The centres of greatest diversity are Brazil and the Indo-Malaysian region (Watt,
1889; Burkill, 1935; Chang, 1958; Keleny, 1962; Leon, 1976). In Peninsular
Malaysia alone there are some 23 genera and 120 species (Henderson, 1954).
The family is often an important component of tropical forests especially the
rainforest. There is considerable diversity of form in the family, as well as evident
adaptation to several rather different habitats. Some genera have become well-
known in cultivation either as ornamental plants, such as Aglaonema, Dieffen-
bachia, Caladium, Scindapsus, Syngonium, Philodendron, Raphidophora, and
genera with colourful inflorescences such as Anthurium, Spathiphyllum,
Typhonium, and Zantedeschia or as useful plants providing food, occasionally the
fruit (as Monstera) or, more usually, the enlarged subterranean storage organs or
corms (Alocasia, Amorphophallus, Colocasia, Cyrtosperma, and Xanthosoma).
One genus, Pistia, has become highly specialized as a floating aquatic plant.
The Araceae are divided into eight subfamilies on morphological and
anatomical grounds (Engler & Prantl, 1889; Engler & Krause, 1920). Each subfamily
is in turn subdivided into tribes. The genera with edible subterranean corms are in
two subfamilies, Lasioideae and Colocasioideae. The first of these consists of
terrestrial or paludose shrubs with sagittate, multiparted leaves with reticulate vena-
tion. They have either bisexual or unisexual flowers, anatropous ovules, and ex-
endospermous seeds. The fibrovascular bundles nave superposed simple laticiferous
cells. Cyrtosperma and Amorphophallus are included in the Lasioideae.
The Colocasioideae contains the useful genera Colocasia, Alocasia, and Xan-
thosoma. The first two are Asiatic, the last is an American genus. These plants are
usually found in wet or swampy ground, have spirally arranged long-petiolate leaves
with a sagittate lamina or with a pair of posterior lobes, and with reticulate vena-
tion. The petiole is peltately attached to the lamina. The flowers are unisexual and
the plants are monoecious, the ovules anatropous or orthotropous. The laticiferous
cells are fused, anastomosing in the fibrovascular bundles.
The genus Colocasia is one of the most important useful genera. There are
seven species, according to Engler & Prantl (1889), all indigenous in Asia (Bentham
199
200 Gard. Bull. Sing 37(2)(1984)
& Hooker, 1883; Conner, 1908). The best known species, Colocasia esculenta, was
first described as Arum esculentum by Linnaeus in 1775 (Hill, 1952), who also
described simultaneously Arum colocasia. Schott established the genus Colocasia in
1832 and renamed Linnaeus’s species as C. antiquorum and C. esculenta. While
some disagreement remains over these ‘‘species’’ many botanists now recognize one
species, though some divide this into subspecies or varieties (Purseglove, 1975). For
example, Colocasia esculenta may be regarded as consisting of two varieties, var.
esculenta (syn. var. typica A. F. Hill) and var. antiquorum (Schott) Hubb. & Rehd.
At the cultivar level, however, there are many more entities, and there has never
been a complete classification of the cultivars. Since Colocasia plants are normally
propagated vegetatively, each region or country tends to have and perpetuate its
own ‘‘forms’’ of which at least 1000 are thought to exist (Kay, 1973).
Owing to the plasticity as well as the morphological and behavioural diversity
of these plants, and to a prior lack of collections and descriptions, there are very few
adequate references to permit the accurate identification of cultivars.
Cytological and karyological studies of the Keladi have been carried out to
some extent but, although several workers have been involved, their findings are in-
conclusive (Ito, 1942; Rao, 1947; Delay, 1951; Yen & Wheeler, 1968; Marchant,
1971).
Local Cultivars of Colocasia esculenta
In Peninsular Malaysia the common name ‘‘Keladi’’ refers not only to
Colocasia but to all similar tuber- and corm-bearing plants including species of
Alocasia, Amorphophallus, and even unrelated plants such as Eichhornia crassipes
(Water Hyacinth; Watson, 1896; Burkill 1935; Furtado 1937, 1940). However, in the
present paper, ‘‘Keladi’’ is used strictly for referring to Colocasia.
Cultivars are usually provided with vernacular names and they are propagated
by vegetative means; hence each is a clone (Allen 1940; Chee and Low 1976; Ghani
1979). The cultivar name may be descriptive or not, and may be known only
regionally.
Though local names have proved helpful, much confusion exists because of the
numerous cultivars, their diversity and plasticity. It is believed that strict definitions
and careful use of the cultivar names is worth achieving, thus permitting better com-
munication between growers and research workers.
Collection and Identification
All plants located during collection trips were first observed in situ, and descrip-
tions of them drawn up. Later they were transferred to prepared experimental plots.
Repeated visits were made to the original collection sites to observe growth, and
these observations formed a basis for comparison with similar ones made on the
plot-grown plants.
Identifications were made using reference specimens preserved in the Her-
barium of the Botanic Gardens, Singapore (SING). From these, and primarily from
the living plants, the identification key was prepared. Characters were tested for
consistency and reliability over a four-year period of study.
A model for the key and characteristics used has been the classic study by
Whitney (Whitney ef a/., 1939) on Hawaiian taro varieties. Characteristics of the
developmental stages of the crop, descriptions of@the inflorescences (where
available) and longevity have all been utilised.
Keys to the Malaysian Keladi 201
The crop cycle of the cultivars is important in identifying the maturity phase i.e.
the mid-crop period. At this stage the colour and markings of the leaf lamina,
petioles, corm flesh, leaf shape and size, number, length and colour of stolons and
rhizome are stable characters for identification purposes.
During the first few months of growth, while the plant attains height and
enlargement of corm size, and produces new leaves in increasing numbers, the
morphological characters are in a state of flux. At mid-crop, maximum plant height,
leaf number and leaf size are attained, after which there is a decline, until at harvest
when leaf number is reduced to 2-3 and plant height reaches between 30 and 40 cm.
Corms of the Keladi cultivars observed over the 4-year experimental period ex-
hibited consistent differences in configuration, which are useful in grouping the
cultivars viz., corm occurring singly and surrounded by cormels of the same size or
different sizes, shapes of branching corms, and presence or absence of stolons.
These characters have been used to initiate a preliminary key for identifying the
Malaysian Keladi cultivars. Ongoing research and observations could be used to up-
date useful characters and contribute towards a better key.
Grouping of Cultivars
GROUP I. Crop cycle 9 to 11 months: — plants tall, 80-110 cm; petioles upright
or spreading, thick; leaves sagittate.
Subgroup Ia:— corms single, cylindrical with ends tapered; plants producing
stolons.
Subgroup Ib: — corms single, cylindrical or globose, surrounded by cormels.
GROUP II. Crop cycle 6 to 8 months: — plant-height medium, 50-70 cm; petioles
upright or spreading, slender, leaves sagittate or cordate.
Subgroup Ila:— parent corm globose, with stolons producing plants close to
the parent corm, giving rise to ‘‘thick’’ clumps of plants of uniform height.
Stolon plants produce 10-12 smaller globose cormels closely arranged around
the parent corm.
Subgroup IIb:— corms cylindrical, single, without stolons.
GROUP III. Crop cycle 5-6 months: — plants short, 30-50 cm tall, petioles
slender, outspreading, leaves small and cordate.
Subgroup II]a:— parent corm single, not branching, surrounded by 6-8 cormels
of similar size and shape.
Subgroup IIIb:— corm branching, appearing like clump of 6-8 corms.
GROUP IV. Crop cycle indefinite:— leaves and petioles not acrid (edible). Corms
reduced (rhizomatous), acrid (inedible).
(2)
Plates 1 & 2. Vegatative characters of some cultivars of Colocasia esculenta. 1, left, Keladi cina; 2, right, Keladi udang.
Plates 3 & 4. Vegetative characters of some cultivars of Colocasia esculenta. 3, left, Keladi minjak, 4, right, Keladi banjar.
203
204 Gard. Bull. Sing 37(2)(1984)
KEY TO PENINSULAR MALAYSIAN Keladi CULTIVARS
Key to Groups I — IV
1. Crop:cycle'5-G;mOnth esses. seca saiozamens sw ceisesucoeasacn sls swoseeceeawencsenckettesmeeeoncemerseecenes GROUP III
1’ Crop cycle longer or indefinite
2. Crop cycle.6-8 months) |.cccccoccs ceteresee ne ote see eee Sie aeons SEE OER C Oe ee GROUP II
2’ Crop cycle longer or indefinite
3.. Cropicycle 9-11 months: ..2.2.6.ceteescataaes cnc sores saeesesewccsncusvousseeeseecucecassonceneeenee GROUP I
3° Crop: cycle indefinite «$< jcc.cc oes oce est sene- ce ucbssaeeaeanecnsanuneemeneinee ce ccesaseesseneamanes GROUP IV
Key to Keladi Cultivars of Group I
1. Parent corms 20-24 cm long, 10-12 cm wide, single, cylindrical, tapered at both ends, producing
numerous (8-10) stolons
2. Leaf lamina not variegated or mottled over entire surface in white, yellow and green
3. Main veins on the under-surfaceof lamina not purplish black. Lamina in shades of green
4. Laminar sinus purplish black-pink, the colour extending only to the petiole, the piko*
green, ‘lamina Sagittate::.-coresc sect etee done pac en cs oscar ieee ene enean tees Keladi cina (Plate 1)
4’ Laminar sinus deep red, the colour extending to all main veins on lower surface, the piko
reddishigreens laminainotisagittate mss sss-ceetassaeeerereeseeeta eeaee nee ree Keladi udang (Plate 2)
3° Main veins on under-surfacepurplish black ..............0....csccecsccccccccoeses Keladi batang hitam
2’ Leaf lamina variegated or mottled in white, yellow and green ................:.cseeeeeeeeeeeees Keladi air
1’ Parent corms 15-18 cm long, 8-9 cm wide, single, cylindrical, surrounded by 5-7 smaller cormels,
without stolons at maturity
5. Lamina with upper surface glaucous dark green and lower surface green; veins and undulating
Margin purplish FEM cs sasiiens alee ecuisacse reseeainsseecees eaeates eee tale e seeeeeceeee acer Keladi nibong merah
5’ Lamina with upper and lower surface and margin otherwise
6. Lamina with upper surface waxy green and lower surface light green; veins conspicuously
ridged) white toicneative..c.cssseecueceeteecen see ce oe ce cece eee hoc eeaesoremeeerecr Keladi minyak (Plate 3)
6’ Lamina surfaces not as above and veins not ridged
7. Petioles purplish black throughout except for the white base ............. Keladi serakit hitam
7’ Petioles not entirely purplish-black
8’ Petioles otherwise
9’ Corm flesh yellow, flecked with pale brown fibres ................. Keladi serakit putih
9’ Corm flesh creamy white, flecked with brownish yellow fibres ...............c0eeeeeeeeees
RR Aa retire iehe sr csnaneistitel aaa pentina taanacnht aaxeBEncnedacdasand Keladi banjar (Plate 4)
*Centre of the leaf lamina
Keys to the Malaysian Keladi 205
fe
1”
1
Key to Keladi Cultivars of Group II
Parent corm globose, 10 cm diameter, surrounded by cormels (6-8) of same size
2. Leaf cordate and pendant
Selcaminasupper sutface wrinkled, pikO, STEEN) <a... <cewesacccveeeseseeesceeceseses Keladi cincang wangi
3’ Lamina: upper surface not wrinkled, piko black splashed ................. Keladi pinang (Plate 5)
2’ Leaf not cordate and not pendant, lamina broad sagittate and cupped .................... Keladi putih
Parent corm not globose, 8-10 cm diameter, 15-18 cm long; tapered at both ends, surrounded by 6-8
smaller cormels or stolons
4. Lamina: upper surface glaucous dark green; margin undulating and blackish purple; veins on the
PME ES HL ACCIDULPIIS MCC pea- nese schnse ce ahs aosiscesceelicccbascceetectccteums beset Keladi bantan (Plate 6)
4’ Lamina: surfaces, margin and veins not as above
5. Petiole reddish purple and with green longitudinal stripes throughout except for the purplish
RASC reels ce eons oo che casas Sose sascha see cee vecetieseessssiseterestaccvsesscvadeaceceeds Keladi songket ungu
5’ Petiole not reddish purple
6. Petiole creamy white with green longitudinal stripes ....................20- Keladi songket hijau
6’ Petiole otherwise
emG@ormiflesh lilac purple; without fibres: ........csnoccecscccevececcecvescessceceveesess Keladi dara
7 Corm flesh reddish orange, with brown fibres ...............0cccececeeeeeeeeeee Keladi kelapa
Key to Keladi Cultivars of Group III
Parent corm non-branching, cormels small, 10 cm diameter, globose (or cylindrical, 10 cm wide,
10-12 cm long)
2. Petiole light green with dark green striations
Seeenoleipaseand collar ring yellow/On DOWN .c..0..cccesc-.o.0esceereceesce eee cccc-cecnssesess Keladi lilin
Sie benole base andicollar ring pinkish purple:.. 2... .20..-.-cs0csnecencscscnevcscccecceeeensuees Keladi dara
2’ Petiole not entirely green, with purple and white striations ...................02.ee0eeeeee Keladi tongsan
PALE MMCORM DANCING ala DEX cmecae shone saee asc sae-eacateasencactesciseassceeseceerlee cesses vases Keladi peladang
Key to Keladi Cultivars of Group IV
Tall plants 80-100 cm high; rhizome underground
2. Petiole greenish yellow throughout, with the basal zone white
Semlecatisapittate: the vpper surface Wrinkled) ...2...5.0c..c.cecccccsccsosssocsececsevwevceseses Keladi huma
Sumleatnotsapittate. the upper surface SMOOth, PlOSSY......2.2..0ccceccenscecscccasecescesesces Keladi air
2’ Petiole colour not as above, but with purplish green streaks; petiole-lamina junction pinkish .......
MMe eee ee rernce estes se nee ctiaee rn cto enlnish sen telesainrsise sb adapledesasgesescesieshtestaass Keladi belang hijau
Shorter plants, less than 80 cm high; rhizome underground, acrid
4. Petiole with alternate bands of blackish green and light green
5. Leaf narrowly sagittate; lamina margin undulating .......................0..0.. Keladi hutan
5’ Leaf not narrowly sagittate; lamina margin smooth ....................cececeeeuees Keladi liar
“ue]URG IPelay sys ‘9 ‘Bueuid Ipelay ‘1fa/ ‘¢ DJuajnrsa DISDIOJOD JO SIAL[NDI IUIOS JO SI9JOBIVYD DATJLIBBIA “9 7 ¢ Saye[d
06
Keys to the Malaysian Keladi 207
4’ Petiole without markings, light green throughout, the basal zone white and the collar ring
AGIT OT COM a eerr teen tesco race sence cnc ac sete cade totes es eee ries eae ee oan cde BLESS Keladi kubis
Acknowledgements
I wish to thank En. Hassan Basir, En. Abd. Rahim Osman, En. Saad Hamid of
the Botany Department, Universitit Kebangsaan Malaysia for their assistance during
the numerous collection trips; Cik Normadiah Omar for typing this manuscript; and
Dr. Abd. Latiff Mohamed of the same Department for his valuable comments. I
am grateful to Dr. B. C. Stone for his constant encouragement and willing advice.
This work is supported by the International Foundation for Science, Sweden
(Grant R269).
Literature Cited
Allen, E. F. (1940). The Cultivation of Colocasia esculenta (L.) Schott in Malaya.
Malayan Agricultural Journal 28(9): 393-399.
Bentham, G. and Hooker, J. D. (1883). Genera Plantarum 3(2): 955-978. Reeves
and Company, London.
Burkill, I. H. (1935). Dictionary of the Economic Products of the Malay Peninsula.
Ministry of Agriculture and Cooperative. Kuala Lumpur, Malaysia: 2 Ed.
Vol. 1: 647-657.
Chang, T. K. (1958). Dispersal of taro in Asia. Annual Journal of the Association
of American Geography 48: 255-256.
Chee, K. S. and Low, A. L. (1976). The cultivation of Keladi China (C. esculenta)
in Johore. Proc. of the Crop Production Board Meeting 1976. Dept. of
Agriculture, Kuala Lumpur, Malaysia. Pp. 2-4.
Conner, J. E. (1908). Seeds and plants imported. United States Dept. of
Agriculture. Bureau of Plants, (Indiana)Bulletin 142: 36.
Delay, G. (1951). Nombres chromosomiques chez les phanerogames. Revue
Cytologique, Paris 12: 12-18.
Engler, A. (1889). Araceae. Genera Plantarum. In: Engler & Prantl, Naturlichen
Pflanzen familien 2(3): 102-153.
Engler, A. and Krause, K. (1920). Araceae. — Colocasioideae Pflanzenreich
IV: 23E: 1-129 Leipzig.
Furtado, C. X.(1937)._ A commentary on the laws of Botanical Nomenclature
Confusion with Colocasia and Xanthosoma species. Gard. Bull. Straits Settl. 9:
1-4.
. (1940). The Malayan Keladis and other edible aroids. Malay. Agri-hort.
Ass. Mag. 10: 11-17.
Ghani, F. D. (1979). The status of Keladi China (C. escu/enta) cultivation in Penin-
sular Malaysia. Proc. 5th int. Sympos. Tropical Root Crops (Manila): 35-54.
Henderson, M. R. (1954). Malayan Wild Flowers: Monocotyledons. pp. 213-243.
The Malayan Nature Society Kuala Lumpur. Reprint.
Hill, A. F. (1952). Economic Botany. 2nd ed. McGraw Hill & Co., New York,
Pp. 368-370 and 500.
208 Gard. Bull. Sing 37(2)(1984)
Ito, T. (1942). Chromosomen und sexualitat von der Araceae: Somatische chromo-
somenzahlen. Cytologia 12: 313-325.
Kay, D. E. (1973). Root Crops. Tropical Products Institute, Foreign & Commonw.
Office, London, TPI Crop & Digest No. 2, 245 pp.
Keleny, G. P. (1962). The origin and introduction of the basic food crops of New
Guinea people. Papua & New Guinea agric. J. 15: 7-13.
Leon, J (1976). Origin, evolution and early dispersal of root and tuber crops. Proc.
4th int. Sympos. Tropical Root Crops (Colombia): 20-31.
Marchant, C. J. (1971). Chromosome variation in Araceae: II. Richardieae to
Colocasieae. Kew Bull. 25: 47-56.
Purseglove, J. W. (1975). Tropical Crops: Monocotyledons. ELBS ed. Longman
Group Ltd., London, pp. 8-70.
Rao, N. S. (1974). A note on the chromosome number in Colocasia antiquorum
Schott. Curr. Sci. Bangalore, 16: 229.
Watson, J. (1896). Malay Plant Names. Malay. Forest Records No. 5. Reprint by
Forest Department, Federated Malay States, pp. 129-130.
Watt, G. (1889). A Dictionary of the Economic Products of India. Superintendent
of Government Printing, Calcutta. Vol. 2: 509-513.
Whitney, L. D., Bowers, F. A. I., and Takahashi, M. (1939). Taro varieties in
Hawaii. Hawaii Agric. Ext. Bull. 84: 1-86.
Yen, D. E. and Wheeler, J. M. (1968). Introduction of taro into the Pacific. The
indications of chromosome numbers. Ethnology 7(3): 259-267.
Notes on the Systematy of Malayan Species of Chionanthus
(Oleaceae)
R. KIEW
Department of Biology, Universiti Pertanian Malaysia, Serdang, Selangor, Malaysia
Abstract
Examination of Blume’s type specimens shows that Chionanthus platycarpus (K. & G.) Kiew is
synonymous with C. callophyllus Bl. and C. elaeocarpus (Stapf) Kiew is synonymous with C. macrocar-
pus Bl. Two new records of Chionenthus for Malaya are C. grandifolius (Elmer) Kiew (previously only
recorded from the Philippines) and C. curvicarpus sp. nov. (formerly known as C. cuspidatus Bl. sensu
Merrill).
Sabbatical leave gave me the opportunity of examining at the Rijksherbarium,
Leiden, Blume’s types (and other specimens determined by him) of Chionanthus,
which show that several of the names of Malayan species need to be changed.
1. Chionanthus macrocarpus Blume
Mus. Bot. Lugd. Bat. 1 (1850) 319. Typus: Java, Herb. Lugd. Bat. 908. 161... 213 (L, lecto, here
chosen).
Linociera macrocarpa (Bl.) Knobl. Bot. Centralbl. 61 (1895) 87; King & Gamble J. As. Soc. Beng.
ii 74 (1905) 267; Back. & Bakh. f. Fl. Java 2 (1965) 214, non (C. macrocarpus sensu Kiew Mal.
For. 42 (1979) 271).
Chionanthus insignis Miquel FI. Ind. Bat. Suppl. (1862) 559.
Typus: Teysmann. Palembang, Sumatra (BO, K, L, syntypes); pro parte Linociera insignis (Miq.)
C.B.Cl. in Hooker f. Fl. Brit. Ind. 2 (1882) 610; pro parte Ridley Fl. Mal. Pen. 2 (1923) 316.
Linociera elaeocarpa Stapf Kew Bull. (1915) 115; Typus: Beccari PB 725 (K, holo). Merrill J. Str. Br.
R. As. Soc., Spec. No. (1921) 488. Chionanthus elaeocarpus (Stapf) Kiew var elaeocarpus Mal.
For. 42 (1979) 267; 43 (1980) 373; 44 (1981) 150.
Notes
Examination of the specimens of Chionanthus macrocarpus at Leiden
annotated by Blume shows no differences between C. macrocarpus from Java and
the Bornean type specimen of Linociera elaeocarpa Stapf. Specimens cited as C.
elaeocarpus (Stapf) Kiew (Kiew 1980, 1981) are therefore also those of C. macrocar-
pus.
Backer & Bakhuizen f. (1965) described the fruit as more or less angular. Fruits
sometimes dry with sides flat but the angles are not ridged and these fruits would not
be confused with Linociera beccarii Stapf (from Sumatra) or C. porcatus Kiew
(from Borneo), which have ridged fruits where the ridges correspond to those on the
woody endocarp. The endocarp of C. macrocarpus, although hard and thick, is not
woody.
The geographic distribution of C. macrocarpus covers Java, Sumatra, Borneo
and Malaya (although it is less common in Malaya than elsewhere). It has been
collected from lowland and hill forest up to 1500 m, sometimes from freshwater
swamp or areas that flood periodically. In Malaya only the typical variety with
ovoid fruits is found; the variety with a globose fruit, var globosus Kiew, is
restricted to Borneo (Kiew, 1980).
209
210 Gard. Bull. Sing 37(2)(1984)
Chionanthus insignis was originally described from Sumatra by Miquel who
cited Teysmann’s specimen from Palembang. This specimen has leaves, in-
fructescences and fruits of the same size and shape as C. macrocarpus with which
C. insignis is thus synonymous. However, Clarke based his description of Linociera
insignis on specimens from Burma (Helfer 3688, K, L) which differ from those of
C. macrocarpus Bl. in their smaller fruits (1 cm long) and glossy leaves. These non-
Sumatran specimens are therefore not specimens of C. macrocarpus and must be
excluded from C. insignis Miquel, which is a synonym of C. macrocarpus.
2. Chionanthus callophyllus Blume
Mus. Bot. Lugd. Bat. 1 (1850) 319. Typus: Korthals s.n. Borneo Herb. Lugd. Bat. 908. 158... 933
(L, lecto — here chosen). Linociera callophylla (B\.) Knobl. Bot. Centralbl. 61 (1895) 319.
Linociera paludosa King & Gamble J. As. Soc. Beng. ii 74 (1905) 268. Typus: Wray 2424 (SING),
Ridley Fl. Mal. Pen. 2 (1923) 316.
Olea platycarpa King & Gamble |.c. 271. Typus: King’s Coll. 5541 (K), Ridley l.c. 319. Chionanthus
platycarpus (K. & G.) Kiew Mal. For. 42 (1979) 272 & Fig 5, 44 (1981) 149.
Notes
Chionanthus callophyllus_ is a distinctive species with large, obovate leaves
(23-38 by 7-13 cm) with a particularly thick petiole, the veins impressed above and
prominent below. In addition, it has short ramiflorous inflorescences and the base
of the fruit is flattened, the latter is a unique character for Malesian species of
Chionanthus. In addition, the fruit is often covered with a white bloom. Ramiflory
is known in only one other Malesian species viz., C. gigas (Lingelsheim) Kiew from
New Guinea. Blume described the inflorescence as equal to or longer than the petiole
(which they are) and as axillary or lateral. In fact the inflorescences are extra-axillary
i.e., ramiflorous.
Chionanthus callophyllus is more common in Malaya (18 collections) than in
Borneo (3 from Sabah and 3 from Kalimantan) or in East Sumatra (one collection,
Krukoff 310, US).
3. Chionanthus grandifolius (Elmer) Kiew comb. nov.
Linociera grandifolia Elmer Leaflets Philip. Bot. 5 (1913) 1657. Typus: Elmer 13425 (with fruits)
Mindanao, Philippines (K, lecto — here chosen, BM, Gray, L; US, isolecto).
Chionanthus macrocarpus non Blume sensu Kiew Mal. For. 42 (1979) 271.
Malayan specimen: Cockburn FRI 10578 (with flowers) Batu Biwa, Trengganu (K, KEP, L, SING).
Notes
The specimen FRI 10578 is outstanding among Malayan Chionanthus species
for its indumentum: no other species has leaves where the lamina is softly and dense-
ly hairy below. In this character it matches the single specimen of Linociera grandi-
folia from Mindanao, Philippines. Both these specimens have oblong leaves with a
rounded to acuminate apex and a cuneate base, the leaf is subcoriaceous and slightly
shiny above, the veins are impressed above and are prominent below and the twigs
are flattened at the node. The Malayan specimen is therefore considered to be the
same as Linociera grandifolia based on vegetative characters, as unfortunately the
Malayan specimen is in flower and the Philippine one in fruit. Linociera is now con-
sidered synonymous with Chionanthus (Stearn, 1971) and the appropriate combina-
tion is made above.
Amended description
Small tree to 7 m. Twigs pale grey, flattened at nodes, pubescent. Leaves
Systematy of Malayan Chionanthus spp. Dal
chartaceous to subcoriaceous, drying deep reddish brown, elliptic oblong, 22-33 by
9-13.5 cm, apex obtusely rounded to acuminate, base acute. Lower surface of
lamina and veins densely covered by long unicellular hairs, upper surface with
sparse, short unicellular hairs. Veins 11-13 pairs, slightly impressed above, promi-
nent below, tertiary veins inconspicuous. Petiole 1.5 cm long, 5 mm thick, shortly
pubescent. FRI 10578 — Inflorescence paniculate, 13-17 cm long, densely hairy,
axillary clustered, branched from base. Flowers sessile, yellow with pleasant scent.
Calyx lobes acute, 1.5mm _ long, densely pubescent. Corolla 3.5 mm long, lobes
joined in 2 pairs for 1 mm at the base, lobes narrowly linear, pubescent, valvate.
Stamens almost sessile, anthers 1 mm long. Ovary conic, stigma obscurely lobed.
Elmer 13425 — Infructescence 3-5 cm long, pubescent. Fruits ovoid to subglobose,
17 by 19 mm, subsessile. Pericarp thin and brittle. Seed 1, endosperm copious.
4. Chionanthus curvicarpus Kiew sp- nov.
Typus: A. Gibot SAN 35826 (SAN — holo, K, UPM — iso).
Linociera cuspidata (B\.) Knobl. sensu Merrill J. Str. Br. R. As. Soc. Spec. No. (1921) 488, Univ.
Cal. Publ. Bot. 15 (1929) 249; non Chionanthus cuspidatus Bl. sensu Kiew Mal. For. 43 (1980) 372 &
Fig 2J, 44 (1981) 152. Non Chionanthus cuspidatus Bl. Mus. Bot. Lugd. Bat. 1 (1850) 319. Typus:
Muller, Gunung Bahay Herb. Lugd. Bat. 908. 158... 996 (L, lecto — here chosen).
Chionanthus curvicarpus sp. nov. propter habitum arboris minoris cortice ramorum albo, folios
chartaceos lanceolatosque, nervis subtus prominentibus, C. oligantho (Merrill) Kiew affinis sed nervis
numerosioribus (8-15 paribus vice -9), inflorescentiis longioribus (1.5-6 cm vice 0.5-1.5 cm) et forma
fructu differt; fructus C. curvicarpi egregius, anguste-ellipsoideus, 2.5 x 1.75 cm, curvatus et porcatus
vice in C. oligantho globosus.
Notes
Chionanthus curvicarpus is common in Borneo where Merrill (1929) identified
specimen Elmer 21047 (BO) near Tawao as Linociera cuspidata (Bl.) Knoblauch.
This interpretation was followed by Kiew (1980, 1981). However, Blume’s type
specimen of C. cuspidatus although similar in leaf shape and size, differs in possess-
ing a shorter inflorescence (1.3 cm long as opposed to 1.5-6 cm in C. curvicarpus),
and the petals are longer (6 mm long as opposed to 2-3 mm long). The most striking
difference is that the leaves dry chestnut-brown (in C. curvicarpus the leaves of all
specimens examined dry pale grey). There are no fruits on the type specimen of C.
cuspidatus. Linociera cuspidata sensu Merrill is therefore not the same as
C. cuspidatus and so a new name is required for this taxon. ‘Curvicarpus’ is chosen
as the curved fruit (which is also ridged longitudinally) is unique among Malesian
Chionanthus species (Fig 2J in Kiew, 1980). Other species that have straight ridged
fruits are larger (6.5 by 3.5 cm in C. porcatus from Borneo, 3-5 by 2-3 cm in
Linociera beccarii from Sumatra and 3-7 by 1.5-3.5 cm in C. sessiliflorus (Hemsley)
Kiew from New Guinea). The fruit of C. curvicarpus is 2.5 by 1.7 cm.
Chionanthus curvicarpus most closely resembles C. oliganthus (Merrill) Kiew
of Borneo. Both species have white twigs, leaves which are thin, dry pale and have
prominent veins below. However, it differs from C. oliganthus in the number of
veins (8-15 as opposed to 6-9 in C. curvicarpus), longer inflorescence (0.5-1.5 cm
versus 1.5-6 cm) and fruit shape (C. curvicarpus has globose fruits).
In Borneo C. curvicarpus is found in the lowlands, sometimes on hillsides up
to 1000 m or near rivers. It is most common in Sabah. In Malaya it is known from
a single flowering specimen, Kiah 32325 (A, K, KEP, L, SING) from Sungai Kayu,
E. Johore and four specimens have also been collected from Sumartra. This same
distribution pattern, the Borneo element of Corner (1960), is also seen in C. /ax-
iflorus B\., which in Malaya is also rare (collected once from Fraser’s Hill and from
the Kuantan-Kluang-Mersing area), whereas it is more common in Borneo and
Sumatra and in addition its distribution extends through Celebes to New Guinea.
AND Gard. Bull. Sing 37(2)(1984)
Acknowledgements
The author thanks the Director of the Rijksherbarium, Leiden, The
Netherlands for permission to work there and to Dr J. Dransfield for help with the
latin diagnosis.
References
Corner, E. J. H. (1960). The Malayan Flora, in Purchon, R. D. ed, Proc. of
Centenary and Bicentenary Congress of Biology, Singapore, pp 21-24.
Stearn, W. T. (1976). Union of Chionanthus and Linociera (Oleaceae). Ann.
Missouri Bot. Gard. 63: 355-357.
A Revision of Aniselytron with Some New Combinations
in Deyeuxia in SE. Asia (Gramineae)
H. M. KORTHOF & J. F. VELDKAMP
Rijksherbarium, Leiden, The Netherlands
EFFECTIVE-PUBLICATION DATE: 7 MAR. 1985
Abstract
Aniselytron Merr. (Gramineae), better known as Aulacolepis Hack. (1934), non Ettingsh. (1893), has two
species in SE. Asia and Malesia. A key and descriptions are given. Anisachne Keng is reduced to Deyeuxia
Beauv. and some new combinations in that related genus are proposed. Deyeuxia abnormis Hook. f. is not
identifical with D. zenkeri (Trin.) Veldk., which is an enigmatic species known only from the type from the
Nilgiris, India.
Introduction
Aniselytron Merr. resembles Deyeuxia Beaux. best in the induration of the lemma
but differs in that its lemma is distinctly longer than the lower glume. This feature is
so aberrant in the Aveneae (including Agrostideae) in which, according to most
descriptions, the glumes might be as long as or longer than the only or lowest lemma,
that Aniselytron can not usually be identified with general keys. Its very short callus
hairs, its minute, glabrous rachilla process and the usual absence of an arista are causes
of the resemblance in the spikelets with those of uniflorous forms of Poa Linné. In
fact, before it was realized that such species occur in Poa, one of its forms was
included in Aulacolepis Hack. by which name the genus was known until recently.
That Aniselytron obviously belongs to the alliance of Agrostis Linné can be noticed
when callus hairs are seen; when present in Poa, they are abaxial and then usually
somewhat arachnoid or woolly; in Agrostis and its related genera, they form a ring
around the base of the lemma, often with the lateral hairs somewhat longer and giving
the impression that there are two tufts. Moreover, in Poa, the lemmas are more or less
keeled at the back while in the other genera, they are more rounded.
A similar reduction in glume size, though rarely as extreme as in Aniselytron,
occurs sporadically in the Aveneae as in some species of Deyeuxia e.g., D. abnormis
Hook. f. from India to S. China; D. exilis Veldk. and D. pusilla (Reeder) Jansen from
New Guinea, the Australian D. gunniana (Nees) Benth., the circumarctic genus Arcta-
grostis Griseb., and the New Zealand Simplicia Kirk (1897). Simplicia especially is very
similar to Aniselytron as was pointed out by Zotov (1971). Indeed, if its two species
had been found in Asia, Aniselytron of 1910 would probably have been included in
Simplicia. However, with the present great disjunction, we get the impression that we
are dealing with the results of two convergent lines in the reduction of the spikelet,
the cause of the apparent resemblance. The difficulty remains that there seem to be
no clear differences whereby the two genera can be separated satisfactorily at the
generic level. The most obvious distinctions are the width of the leaves: 1.5-3 mm
in Simplicia and 3-17 mm in Aniselytron; the lemma: 3-nerved in Simplicia and
5-nerved in Aniselytron (although Zotov has remarked that in S. /axa Kirk there may
be weak additional nerves...); the absence of callus hairs (at least in the single specimen
seen by us of the same and neither mentioned nor depicted by Zotov); and the sub-
terminal arista.
Zotov and the few other authors who have dwelt on Aniselytron repeated Hackel’s
remark (1907) that having the two proximate nerves of the palea would not make it
Pils
214 Gard. Bull. Sing 37(2)(1984)
distinct from Aulacolepis (means “fluted scale”). These nerves are indeed close to one
another, with a more or less narrow depression in between, hardly a furrow. The
minute rachilla process lies in this depression for which reason it is difficult to detect.
It seems to us that Zotov had not actually seen material of Aniselytron for such
a nervation is present also in at least the isotype of S. /axa in Kew. This proximity of
the nerves seems to depend on the extent the palea is covered by the lemma; such close-
set nerves are sometimes present in species of Agrostis and Deyeuxia as well.
~ It is therefore not surprising that Hitchcock (1934) was misled by the apparent
similarity when he distinguished between Aul. clemensae Hitchc. from Sabah and Au.
petelotii Hitchc. from North Vietnam. Again, in 1936, he fell into the trap provided
by the uniflorous Poa species when he proposed the combination Aul. epileuca for a
taxon from New Guinea, one which was described by Stapf in Deyeuxia. However, it
turned out that Awl. epileuca is not Stapf’s species but another uniflorous Poa (see
under “Excluded Names”). Awl. pete/otii has had a rather chequered history: up to now
the taxon has been included in Agrostis, Anisachne Keng and Deyeuxia. We regard the
species as a member of the last genus.
The first mention of a representative of Aniselytron was based on material from
Sikkim. Kuntze (1891) thought that it represented a Milium Linné. This was soon
corrected by Stapf (1895), admitting that he included the species in Deyeuxia(!) for
technically unsatisfactory reasons. He compared it with his D. epileuca, the Poa
intended by Hitchcock (see above). Hooker f. (1896) also regarded it as a “very
anomalous” species for Deyeuxia.
Hackel (1907) proposed a separate genus Aulacolepis for Kuntze’s species and for
some collections from Japan which he suspected might prove to be merely subspecies
of one taxon. This suspicion was affirmed by Ohwi (1941), who regarded them as
varieties while we think that they are merely two local races, not warranting any
nomenclatural status.
In a number of instances, some authors have used Jndex Nominum Generi-
corum to achieve results by proposing all the new combinations under another generic
name without any specialistic knowledge of the group concerned or without a
laborious search for relevant material and/or literature. For them entries in the Index
Kewensis apparently gave enough information. Thus three persons have made all the
“necessary” combinations under Aniselytron, and a fourth, in the blissful ignorance
of that name even proposed a new one, Neoaulacolepis (see also Wilson’s comment,
1983).
As more collections from different places became available, more taxa were added
to Aulacolepis. Ohwi (1935) included Aul. milioides(!) from Taiwan, which had
previously been described as a Poa(!) by Honda (1927). As mentioned above, Hitch-
cock (1934; 1936) distinguished between Au/l. clemensae, Aul. petelotii and Aul.
epileuca. Ohwi (1956) included Aul. pseudopoa(!), a new combination of a species
from North Sumatra, described previously by Jansen. The latter, as can be deduced
from his correspondence in Leiden, had described it as a Deyeuxia on Ms. Vickery’s
suggestion, who had revised that genus for Australia where some species also have
similar relatively short glumes.
The distribution of Aniselytron species is very scattered. The plants apparently
prefer shaded, moist, rocky ravine floors, places out of reach except perhaps in Japan,
and thus material is rare in herbaria. The form of Mt. Kinabalu, for instance, has been
collected only by Ms. Clemens along the Masilau River and the Silau basin. The first
locality seems close to the Paka Caves, a site frequently visited by botanists. The grass
probably grows in the deep gorge below the Caves. Merrill (1910) remarked that An.
agrostoides was uncommon on Mt. Pulog. As far as we know, it has been collected
only twice again [BS 44989 (Ramos & Edano); Santos 7854]. The scarcity could be
iB
Aniselytron & Deyeuxia in SE. Asia 215
attributed to its close resemblance with Agrostis rigidula Steud. as had been observed
by Merrill. That is a rather common grass on Mt. Pulog.
From the present study we think that Aniselytron has only two species: one, An.
agrostoides, occurring in Luzon and in Taiwan, the other, An. treutleri, ranging from
Sikkim through Assam, north Burma, China, north Vietnam and Taiwan to Japan,
Sabah and North Sumatra. The second species has a number of local races as is to
be expected in disjunctive populations scattered over such a wide area. In Japan, only
one form is rather widely spread but the situation in China is not clear because too
few collections are available. The races can only be distinguished by a certain set of
characters which occur in a different expression and/or combination in each individual
race. Provenance is usually the best guide but is most unsatisfactory. We have therefore
refrained from naming them officially. Although they are allopatric, the rank of
subspecies automatically and uncritically bestowed on such taxa by some seems to be
too honorific for the little-impressive differences.
Aniselytron
Aniselytron Merr.
In Merr. & Merritt, Philip. J. Sc. 5, Bot (1910) 329; Sojak, Cas. Nar. Muz. 148 (1980) 202; Raizada &
Bennett, Ind. For. 107 (1981) 433. — Type: Aniselytron agrostoides Merr.
Aulacolepis Hack. in Fedde, Repert. 3 (1907) 241; Hitchc., J. Washington Ac. Sc. 24 (1934) 290; Ohwi, Bot.
Mag. Tokyo 55 (1941) 361; Bor, Grasses (1960) 394; Hsu, Fl. Taiwan 5 (1978) 433; non Ettingshausen
(1893). — Neoaulacolepis Rauschert, Taxon 31 (1982) 561. — Lectotype: Aulacolepis treutleri (O. Ktze)
Rauschert = Aniselytron treutleri (O. Ktze) Sojak.
Tufted perennials, branching extra-vaginally at base. Culms simple, glabrous,
smooth, but cataphylls and lowest sheaths finely puberulous. Ligules membranous,
enlarging upward, the lower collar-shaped, usually rather thick, outside puberulous,
the upper triangular, thinner, glabrous or not. Blades flat, flaccid, slightly unequal-
sided, glabrous, smooth or scaberulous. Panicles of lax spikelets. Spikelets somewhat
laterally compressed, 1-flowered, articulating above the glumes. Glumes membranous
to slightly indurated in fruit, unequal to very unequal, at least the lower one (much)
shorter than the lemma. Rachilla process minute, glabrous. Lemma ovate-lanceolate,
indurated, scabrous. 5-nerved, muticous or (rarely) with an apical filiform arista,
callus hairs very small, inconspicuous, around the base of the lemma, the laterals +
longest. Palea as the lemma, nearly completely clasped by it, nerves 2, rather close-set
and slightly impressed in between. Lodicules glabrous, sometimes slightly fimbriate,
thin, veinless, free. Anthers 3. Styles 2, free at base. Caryopsis ellipsoid, + terete, not
furrowed; hilum subbasal, punctiform; embryo 0.2-0.25 times the length of the
caryopsis.
Distribution. Two species from Sikkim, N. Burma through China to Japan,
Taiwan, Luzon, Sabah, N. Sumatra.
Ecology. High mountains, in shaded, moist forest.
Chromosome number. X = 7 (see under A. treutleri).
KEY
1. Lateral spikelets with rather uniform, short, up to l-mm long pedicels. Lower
glume virtually absent or a nerveless scale, up to 0.75 mm long, up to 0.2 times
the length of the spikelet. Upper glume I-nerved .............. 1. An. agrostoides
Lateral spikelets with filiform pedicels of variable length. Lower glume 0.5-2.5 mm
long, 1-nerved, 0.25-0.71 times the length of the spikelet. Upper glume 3-nerved
MEINE ee eRe ie ators er ajatcts slaie « cies 5 ibieisls'sals\cis's lols osjeiaia see'sio's oisiaeia'eie @S'nsie 2. An. treutleri
216 Gard. Bull. Sing 37(2)(1984)
1. Aniselytron agrostoides Merr.
In Merr. & Merritt, Philip. J. Sc. 5, Bot. (1910) 329; Enum. Philip. Fl. Pl. 1 (1923) 82. — Aulacolepis
agrostoides Ohwi, Acta Phytotax. & Geobot. 4 (1935) 30; Bot Mag. Tokyo 55 (1941) 361, comb. ill.
— Type: Merrill 6483 (PNH, ¢ holo; K, L, W Hb. JV. Santos, n.v.), Philippines, Luzon, Mt. Pulog,
2400 m, May 1909.
Aulacolepis agrostoides (Merr.) Ohwi var. formosana Ohwi, Acta Phytotax. & Geobot. 4 (1935) 30;
Hsu, Taiwan Gr. (1975) 303, t. 46; Fl. Taiwan 5 (1978) 434, t. 1389. comb. ill — Type: Ohwi 2363
(KYO, holo; K), Taiwan, Ilan Co., (Mt.) Taiheisan (Taipingshan), 26 May 1933.
Culms erect to geniculately ascending, up to 90 cm high, sometimes rooting in the
lower notes. Ligule 0.3-1.5 mm high, margin erose to fimbriate. Blades 10-24 cm by
2.3-6 (-8.5) mm. Panicle erect to slightly nodding, usually + contracted, to effuse with
patent, sometimes slightly reflexed branches, 9-22 by 0.7-13.5 cm diam., axis glabrous,
smooth, branches smooth to scaberulous, lowermost 2-5 together, the longest up to
7.5 cm long, naked in the lower 0.5-0.7th, spikelets 3-15 (-many), pedicels of lateral
spikelets 0.3-1 mm long, of the terminal ones much longer. Spikelets 2.5-4 mm long.
Lower glume virtually absent to a small, scabridulous, nerveless scale up to 0.75 (-1)
mm long, up to 0.2 (-0.3) times as long as the spikelet; upper glume ovate-oblong to
ovate-lanceolate, very variable, 1-2.75 mm long, (0.2-) 0.5-0.75 times the length of the
spikelet, long-acuminate, I-nerved, glabrous, smooth. Rachilla process 0.35-0.8 mm
long. Lemma with the margins becoming scarious, attenuating upward into an acute
to acuminate and then sometimes hook-like apex, muticous; callus hairs 0.05-0.2 mm
long. Anthers (0.6-) 1.25-1.5 mm long, yellow. Caryopsis 1.25-1.5 mm long.
Distribution. Taiwan: Chiayi (Mt.) Arisan, ((Mt.) Morrison), Hsinchu ((Mt.
Taihasensan), Hualien (Mt.) Chilaisan), Ilan ((Mt.) Daikansan, ((Mt.) Loyehweisan,
(Mt.) Pahsiensan); Malesia; Philippines, Luzon (Mt. Pulog).
Ecology. Mixed montane forest, 2150-2850 m, or open grassland just below the
summit of Mt. Pulog (but notes usually lacking on the labels).
Note. Merrill (1910) reported that this species was uncommon on Mt. Pulog. In
fact it has been collected only twice again [BS 44989 (Ramos & Edano); Santos 7854].
There seem to be no collections present in PNH (Dr. Santos, Manila, in /itt.). A poor
juvenile isotype (W) and some scraps of the second collection and Santos’ specimen
were all that were available but these fall well into the range of variability shown by
the ample Taiwan material. When Ohwi proposed a distinct variety for Taiwan, as far
as can be ascertained, he never saw any Philippine material but relied on descriptions
only. His proposal cannot stand.
2. Aniselytron treutleri (O. Ktze) Sojak
Cas. Nar. Muz. 148 (1980) 202; Bennett & Raizada, Ind. For. 107 (1981) 434. — Milium treutleri O. Ktze,
Rev. Gen. Pl. 2 (1891) 780. -- Deyeuxia treutleri Stapf in Hook., Icon. Pl. 24 (1895) t. 2396; Hook. f.
Fl. Br. Ind. 7 (1896) 269. — Aulacolepis treutleri Hack. in Fedde, Repert. 3 (1907) 242; Honda, J. Fac.
Sc. Imp. Univ. Tokyo III, 3 (1930) 195; Schmid, l’Agron. Trop. 13 (1958) 484, f. 2; Bor, Grasses (1960)
394; Hsu, Taiwan Gr. (1975) 305, t. 47; Anon., Icon. Corm. Sin. 5 (1976) 104, t. 7037; Hsu, Fl. Taiwan
5 (1978) 434, t. 1390; comb. ill. — Lectotype: Treutler 486 (K, holo; L, W; CAL, nv.), Sikkim, Tanglo,
3050 m, 5 August 1874.
Aulacolepis japonica Hack. In Fedde, Repert. 3 (1907) 241; Honda, J. Fac. Sc. Imp. Univ. Tokyo III, 3
(1930) 195; Anon., Icon. Corm. Sin. 5 (1976) 104; comb, ill. — Aulacolepis treutleri (O. Ktze) Hack.
var. japonica Ohwi, Bot. Mag. Tokyo 55 (1941) 361; Fl. Japan (1965) 158; comb. ill. — Aniselytron
japonicum Bennett & Raizada, Ind. For. 107 (1981) 434. — Neoaulacolepis japonica Rauschert, Taxon
31 (1982) 561. — Lectotype: Faurie 6394 (W, holo; P, nv; KYO, L), Japan, Honshu, Prov. Shinano,
Mt. Komagetake, July 1905.
Aniselytron & Deyeuxia in SE. Asia PAG
Poa milioides Honda, Bot. Mag. Tokyo 41 (1927) 641. — Aulacolepis milioides Ohwi, Acta Phytotax. &
Geobot. 2 (1933) 161, 169; comb. ill. — Aulacolepis treutleri (O. Ktze) Hack. var. milioides Ohwi, Acta
Phytotax. & Geobot. 6 (1937) 151; comb. ill. — Aniselytron milioides Bennett & Raizada, Ind. For.
107 (1981) 434. — Lectotype: Faurie 205 (KYO, holo; L; P, mv.), Taiwan, Chiayi Co., (Mt.) Arisan,
2500 m, December 1914.
Aulacolepis clemensae Hitchc., J. Washington Acad. Sc. 24 (1934) 290; comb. ill. — Aniselytron clemensae
Sojak, Cas. Nar. Muz. 148 (1980) 202; Bennett & Raizada, Ind. For. 107 (1981) 434. — Neoaulacolepis
clemensae Rauschert, Taxon 31 (1982) 561. — Type: Clemens 34448 (US, holo; BM, G, K, L), Sabah,
Mt. Kinabalu, Masilau River, 26 December 1933.
Deyeuxia pseudopoa Jansen, Acta Bot. Neerl. 2 (1953) 363, f. 1. — Aulacolepis pseudopoa Ohwi,
J. Jap. Bot. 31 (1956) 137; comb. ill — Aniselytron pseudopoa Sojak, Cas. Nar. Muz. 148 (1980) 202;
Bennett & Raizada, Ind. For. 107 (1981) 434. — Type: Van Steenis 9135 (L, holo; BO, n.v.), Sumatra,
Aceh, summit Goh Lembuh, 3000 m, 21/22 February 1937.
Culms erect, up to 1 m high, rarely somewhat geniculate or decumbent and
rooting in the lower nodes. Ligules 0.5-7.5 mm long. Blades 10-26.5 cm by 3-17 (-21)
mm. Panicle effuse, erect, 9-26 by 0.7-15.5 cm diam.; axis smooth; branches scabrid,
the lowermost 2-7, usually 5 together, the longest up to 12.5 cm long, naked in the
lower 0.45-0.7th, spikelets 7-many; pedicels of the lateral spikelets filiform and variable
in length. Spikelets 2.35-4.6 mm long. Glumes usually distinctly unequal, ovate-
lanceolate, acute, scabrous; lower glume 0.5-2.6 mm long, 0.25-0.85 times as long as
the upper one, 0.2-0.7 times as long as the spikelet, 1-nerved; upper glume 1.6-3.8 mm
long, 0.6-1 times as long as the spikelet, 3-nerved. Rachilla process 0.2-1.3 (-2) mm
long. Lemma with an acute to acuminate apex, sometimes with a thinner tissue and
then sometimes incurved, rarely with an apical, filiform arista up to 0.25 mm long,
callus hairs up to 0.8 mm long. Anthers 0.75-1.5 (-2) mm long when chasmogamous,
0.75-1.35 mm long when cleistogamous. Caryopsis 1.25-2 mm long.
Distribution. (Most collections were summarily labeled with often just a rough in-
dication of provenance. Many localities could not be traced, e.g. in Japan. Field notes
were usually meager or absent). Sikkim (Jongri, Nanki, Shimong, Tonglo), Burma
(Kachin: Uring Bum), China (Guizhou, Fan Ching Shan; Sichuan; north Guangxi),
Taiwan (Chiayi, Ilan, Nantou, Taipei), north Vietnam (Fan Si Pan Mts., Chapu),
Japan (Honshu: North to Shina; Prov. Kyushu: Bungo, Mt. Kuju), Malesia (Sabah:
Mt. Kinabalu; Sumatra: Aceh, Mt. Leuser).
Ecology. Moist, shaded, often rocky places in midmontane to upper montane
areas, often in ravines (see also under the entities).
Chromosome number. 2n = 42 (here published with the kind permission of
Dr. T. Tateoka; vouchers: Tuteoka 9389, 9395, KYO).
Notes. Although local forms seem to be present, the collections were usually so
few that no clear idea of their variability could be formed. The differences between
the entities, moreover, are expressions of a mixture of recurring characters each time
with a slightly different value. We thought that it was therefore premature to describe
different taxa for the various localities and have not accepted the varieties japonica,
milioides and treutleri of previous authors. Short diagnoses should suffice here as an
aid in future research.
Sikkim, Burma (‘treutleri’). Lower glume 0.5-1.75 mm long, 0.25-0.8 time length of the
upper, 0.2-0.65 time length of the lemma; upper glume 1.6-2.5 mm long, 0.7-0.9
time length of the lemma. Lemma 2.35-2.75 mm long. Rachilla process 0.2-0.4 mm
long. Callus hairs up to 0.2 mm long. Anthers 0.75-1.25 mm long, often cleisto-
gamous, if so, then of same size. Alt. 2275-3660 m.
China (Guizhou, Sichuan?, N. Guangxi). Lower glume (0.65-) 1-1.5 mm long, (0.3-)
0.45-0.7 time length of the upper, (0.2-) 0.3-0.5 time length of the lemma; upper
218 Gard. Bull. Sing 37(2)(1984)
glume 2.15-2.5 mm long, 0.67-0.8 time length of the lemma. Lemma 2.75-3.3 mm
long. Rachilla process 0.35-0.75 mm long. Callus hairs 0.25-0.5 mm long. Anthers
c. 1.1 mm long. No altitudes given.
Taiwan (‘milioides’). Lower glume 1-1.9 mm long, (0.33-) 0.45-0.8 time length of the
upper, 0.35-0.55 time length of the lemma; upper glume 2.1-2.75 mm long, 0.7-0.85
time length of the lemma. Lemma 2.8-3.5 mm long. Rachilla process 1-1.35 mm
long. Callus hairs 0.25-0.3 mm long. Anthers c. 1.25 mm long. An altitude of
c. 2500 m was once mentioned, other ecological notes are lacking on the field labels
and in the literature.
North Vietnam (and China, Sichuan?). Lower glume 2-2.1 mm long, 0.8-0.85 time
length of the upper, 0.5-0.55 time length of the lemma; upper glume 2.35-2.55 mm
long, 0.6-0.67 time length of the lemma. Lemma 3.8-4 mm long. Rachilla process
1-1.25 mm long. Callus hairs c. 0.3 mm long. Anthers 1-1.25 mm long. N.B Only
1 collection seen (Pételot 8025, L, US), the duplicate in P, which could not be found,
was presumably described by Schmid as Au. treutleri. His dimensions differ thus:
‘lower glume 1.7-2 mm long, upper glume 2.2-3 mm long, lemma 3-4 mm long’,
measurements resembling those of ‘milioides’. The anonymous author(s) of the Icon.
Corm. Sin. may have described this form as Au. japonica; if so, it would also occur
in Sichuan Alt. c. 2900 m in Vietnam.
Japan (Japonica’). Lower glume 0.9-1.6 mm long, 0.4-0.65 time length of the lemma;
upper glume 2-3 mm long, (0.6-) 0.65-0.85 (-0.95) time length of the lemma. Lemma
(2.65-) 3-3.55 mm long. Rachilla process (0.6-) 0.75-1.3 (-2) mm long. Anthers 1-1.35
(-2) mm long when chasmogamous, 0.75-1.35 mm long when cleistogamous. Alt.
1300-2500 m. 2n = 42. N.B Some specimens have dorsally pilose lemmas.
Sabah (‘clemensae’) Lower glume 2-2.5 mm long, 0.7-0.8 time length of the upper,
0.55-0.7 time length of the lemma; upper glume 2.85-3.25 mm long, 0.8-1 time
length of the lemma. Lemma 3.25-3.5 mm long. Rachilla process 0.4-1.15 mm long.
Callus hairs 0.4-0.5 mm long. Anthers 1.25-1.5 mm long. Alt. c. 2100 m, with
Astilbe, Galium, Gunnera, Juncus. Only found by Ms. Clemens!
Sumatra (pseudopoa). Lower glume 1.9-2.6 mm long, 0.55-0.75 time length of the
upper one, 0.45-0.65 time length of the lemma; upper glume 3-3.8 mm long,
0.75-0.95 time length of the lemma. Lemma 3.75-4.6 mm long. Rachilla process
0.35-1.15 mm long. Callus hairs 0.25-0.35 mm long. Anthers 1-1.5 mm. Alt.
2500-3350 m. N.B The lemmas in Van Steenis 8660 and De Wilde & De Wilde-
Duyfjes 16816 are sparsely pilose on the midrib and the outer nerves, those in Van
Steenis 9135 and De Wilde & De Wilde-Duyfjes 1637] are glabrous.
Excluded names and some new combinations in Deyeuxia
1. Deyeuxia abnormis Hook. f.
Fl. Br. Ind. 7 (1896) 268; Keng, J. Washington Acad. Sc. 48 (1958) 188, in passim. — Lectotype:
Hooker f. & T. Thomson s.n. (Agrostis 12’), (K, holo, nv, L), India, Meghalaya, Mt. Khasia, 1525
— 1830 m. (Here designated).
Agrostis pleiophylla Mez In Fedde, Repert. 17 (1921) 301. — Syntypes: C.B. Clarke s.n. (B, +, holo; K,
ny.), Himalaya, Darjeeling; Khasia, Soynung (probably syntypes of D. abnormis).
Aulacolepis petelotti Hitchc. J. Washington Acad. Sc. 24 (1934) 291; Schmid, l’Agron. Trop. 13 (1958) 484,
comb. ill. — Aniselytron petelotii Sojak, Cas. Nar. Muz. 148 (1980) 202; Bennett & Raizada, Ind. For.
107 (1981) 434. — Neoaulacolepis petelotii Rauschert, Taxon 31 (1982) 561. — Type: Pételot 4743 (US,
holo; L; P, m».), north Vietnam, Fan Si Pan Mts, Chapu, 1900 m, August 1933.
Anisachne gracilis Keng, J. Washington Acad. Sc. 48 (1958) 117, f. 2; Anon., Icon. Corm. Sin. 5 (1976) t.
7036. — Type: Hou Hsueh-Yuh 2143 (N, holo, ny.), China, Kweichow, Pichieh Hsien, 1440 m, 1 June
1943.
Aniselytron & Deyeuxia in SE. Asia 219
That Deyeuxia abnormis is a rare but rather widely distributed species in SE. Asia
was unknown until now as the species in each area went by a different name. When
Keng described Anisachne gracilis he actually did think of D. abnormis but in the
absence of material had to rely on Hooker f.’s description (1896) of it. Moreover, he
remarked that he would have included his new species in Deyeuxia if the glumes had
been longer. We now know (see also the Introduction) that glumes shorter than the
lemma do occur sporadically in Deyeuxia. The additional presence of a relatively
thickish, subterminal aristule, a hairy rachilla process, and fairly long callus hairs led
us therefore to agree with Hooker f. that the species is better accommodated in
Deyeuxia. Although we have not seen Keng’s original material his description and
plate are a good match for the material of Au. petelotii and D. abnormis.
Agrostis zenkeri Trin. was equated with D. abnormis Hook. f. by Bor (1954) in
an extensive discussion on its provenance and identity. However, having seen the type
specimens of both, we have to disagree with his conclusion and it would seem that the
problem is even more complicated and enigmatic.
The description of Ag. zenkeri as given by Bor does not fit the actual type, but
refers to the true D. abnormis, which is a different species. Both belong to Deyeuxia
as accepted here and a new combination is necessary:
2. Deyeuxia zenkeri (Trin.) Veldk. comb. nov.
Agrostis zenkeri Trin., Mém. Acad. Sc. St. Pétersburg VI, 6 (1841) 363; Steud., Syn. 1 (1854) 170;
Hook. f., Fl. Br. Ind. 7 (1896) 258, sp. dub.; Bor, Kew Bull. (1954) 441; Grasses (1960) 392. — Type:
Schmid, s.n. (?) in Hb. Zenker (LE, holo; Kew neg. 2303), India, Tamil Nadu, Nilgiris.
The type of D. zenkeri consists of a single flowering culm, slightly geniculate, c.
35 cm long, with 5 leaves but without a base. Its life cycle can therefore not be deter-
mined. The internodes are glabrous, smooth, and also the sheaths, although the upper-
most is slightly retrorsely scaberulous. Ligules truncate, erose, scaberulous outside, the
lowermost longest, 2.25 mm long. Blades infolded, slightly expanded at the very base,
the largest 4.5 cm long, 2.25 mm wide, smooth, glabrous. Panicle lax, spikelets few, 6.5
by 4.5 cm. Axis, branches, and pedicels smooth, axils swollen, glabrous. Branches
short, filiform, wavy, the 2 lowermost together, the longest 3 cm long, spikelets 8,
nearly perpendicularly forked several times, the pedicels subequal, 3-7 mm long, apex
clavate, the spikelets apparently all terminal at the end of the branches. Spikelets
2.6-3.25 mm long. Lower glume as long as the spikelet, 0.1-0.3 mm the length of the
upper, 1.05-1.15 times as long as the lemma. Lemma 2.25-2.95 mm long, 5-nerved,
microscopically scaberulous, apex muticous, 5-dentate, callus hairs c. 1 mm long,
process c. 0.5 mm long with c. 1-mm long hairs. Palea as long as the lemma to slightly
longer. Anthers c. 1.2 mm long (1 seen).
The inflorescence of D. zenkeri is reminiscent of the plate of Ag. tenuis in Hub-
bard’s well-known ‘Grasses’ (1968, f. 300), but with much fewer spikelets and is quite
different from that of D. abnormis, as can be seen in Keng’s plate of Anisachne gracilis,
although that plate seems to have been based on a rather depauperate specimen. There
the panicle is contracted with nearly erect, rather long, more or less straight,
scaberulous branches with many appressed spikelets, usually with short, obconical
(0.75-1 cr more mm long) pedicels. The glumes and palea are distinctly shorter than
the 5- but not the 3-nerved lemma, which often has an inconspicuous, subapical,
straight arista hardly exceeding its apex. The anthers are 0.8-1 mm long.
The species was named after J. C. Zenker by Trinius and described from the
Nilgiris. We agree with Bor that the type was probably collected by B. Schmid, who
gave his material to Zenker to study (he never was in India) and that Zenker apparently
passed on the grasses to Trinius. Now, no matching specimens have been collected ever
since in the area, under this or another name and mention is not even made in the local
floras of Fyson (Fl. Nilg. & Puln., 1915) and Fischer (in Gamble, Fl. Presid. Madras
3, 1928). So either this represents a very rare or overlooked species, or a mix-up has
220 Gard. Bull. Sing 37(2)(1984)
taken place and a non-Indian species was instead represented. We have been unable,
however, to identify it with anything else. Because of the smooth, short, forked
branches of the panicle, the long pedicels, the smooth glumes, the relatively short
callus hairs (c. 1 mm long) and process (c. 0.5 mm long, overlooked by Trinius), and
the glabrous, dentate, at least the unawned lemma here, it seems to differ from any
American, Asian, Australian, European, or Indian species known to us.
3. Deyeuxia brachytricha (Steud.) Veldk., comb. nov.
Calamagrostis brachytricha Steud. Syn. 1 (1854) 189. — Calamagrostis brevipila Steud. ex Miq.,
Cat. Mus. Lugd. Bat. 1 (1870) 112, nomen (copied from label but the name had been changed
by Steudel, 1854). — Calamagrostis arundinacea (Linne) Roth var. brachytricha Hack., Bull.
Hb. Boiss. 7 (1899) 652. — Deyeuxia silvatica (Schrad.) Kunth var. brachytricha Hack. ex
Rendle, J. Linn. Soc. London, Bot. 36 (1904) 396; Ohwi, Fl. Jap. (1965) 147. — Type: Anon.,
s.n. (L, holo, ‘Calamagrostis brevipila’ Steud. ms.), Japan, probably ex Hb. Von Siebold in view
of the rice-paper label with Japanese handwriting (For further synonymy see Ohwi, 1965).
When regarded as a variety, the epithet should be taken from Calamagrostis varia
(Schrad.) Host var. /ongiaristata Korsh., Acta Hort. Petrop. 12 (1892) 420 (‘longearis-
tata’), which is older than Hackel’s use of ‘brachytricha?
4. Deyeuxia debilis (Hook. f.) Veldk., comb. nov.
Calamagrostis debilis Hook. f., Fl. Br. Ind. 7 (1896) 262. — Agrostis debilis Bor, Grasses (1960) 387,
non Poir, (1810). — Agrostis neodebilis Bennett & Raizada, Ind. For. 107 (1981) 433. — Type:
Hooker f, s.n. (‘1 Calamagrostis”), (K, holo; L), Sikkim, Chola, 3050 m.
Aniselytron agrostoides Marr. J. V. Santos no. 7854
Reduced xerox copy Actual size of specimen
of original illustration.
Magnification
a. Habit of plant
b. Junction between
sheath & blade
49 cm. tall
blade 2.3 mm wide
ligule 1.4 mm long
Spikelet (3 views)
c. lateral view:
Glume:: {9 ) weep Et [ile See ree ee 0.17 mm long
Glimies TIA) Pisgy PRR i Ra deececeres tectee eeteerreasteeceer ee 1.2 mm long
d. dorsal view
e. ventral view
2.9 mm long
Lemma
f. lateral view:
hairs of callus
g. spread out lemma
2.8 mm long
ab oe Noawiatguy dan tcdbanecien eta bine ease 0.05 — 0.2 mm long
Woincde snsersd occas ck cttoctecceeaedeee anes 2.8 mm long
1.4 mm wide
Palea (3 views) X 24.5 — 2.7 mm long
h. lateral view
i. dorsal view:
rachilla produced
Into‘an*awn itt Re EE Mea. Macee cccscectce thc messescnahacerent 0.5 mm long
ventral view
k. Pistil and stamens X 24.5 2.4 mm long
ovary 1.23 mm long
style 0.4 mm long
stigma 0.8 mm long
anther 0.86 mm long
filament 1.75 mm long
lodicules 0.3 mm long
uo.
1. Caryopsis X 24.5 1.35 mm long
0.35 mm wide
Fig. 1. Aniselytron agrostoides (Santos 7854).
a. Habit; b; ligule; c. spikelet lateral view; d id., dorsal view; e. id., ventral view; f. lemma, lateral view; g. id.,
flattened; A, palea lateral view i id., dorsal view /. id., ventral view; k. pistil and stamens; /. caryopsis. (a
Db 6 41's C1 XK, 16.
221
222 Gard. Bull. Sing 37(2)(1984)
5. Deyeuxia epileuca Stapf
Trans. Linn. Soc. London, Bot. II, 4 (1894) 247, t. 20c, 10-16. — Aulacolepis epileuca Hitchc.,
Brittonia 2 (1936) 177, pro comb, comb. ill. — Aniselytron epileuca Sojak, Cas. Nar. Muz. 148
(1980) 202 (epileucum’); Bennett & Raizada, Ind. For. 107 (1981) 434. — Type: Haviland 1401
(K, holo), Sabah, Mt. Kinabalu, 3200 m.
= Poa epileuca (Stapf) Stapf in Hook., Icon. Pl. 27 (1899) t. 2607, in obs.; t. 2608.
This has long been a misunderstood taxon, partly because the spikelet is uni-
florous, partly because it has generally been confused with Poa papuana Stapf (e.g.,
by Hitchcock, 1936), Poa wisselii Jansen (cf. Chase, J. Arn. Arb. 24, 1943, 84) and
other uniflorous Poa species (see also Veldk. in Van Royen, Alp. Fl. N. G. 2, 1980,
1100). For the differences with Aniselytron see the Introduction.
6. Deyeuxia mazettii Veldk., nom. nov.
Calamagrostis stenophylla Hand.-Maz., Symb. Sin. 7 (1936) 1298, t. 40, f.1, non D. stenophylla
Jansen (1952). — Type: Handel-Mazetti 5173 (W, holo, nv, L), China, Sichuan, Lungchushan Mts.,
Huili, 3100-3400 m, 16 September 1914.
Index of collectors
Only numbered collections have been included. Specimens cited in literature but
not seen are also included with their identifications (between brackets) when these
seemed acceptable; otherwise they have been deleted.
BS 44989 (Ramos & Edano): 1.
Ching 6062: 2; Chuang 316: 1; Clarke 26044, 26046, 27482, 27438, Clemens 29692,
34448, 34448-A: 2.
Faurie 205, 6394, 6397: 2; Fukuyama 4778: 1.
Henry 4777, Hosomi 8970: 2; Hsu 437: 1; 5942: (2); 5942-A, Huang ef al. 5766: (1).
Kao 5201: (2); 5868: (1); King 3101, Kingdon Ward 21568, Kunio 35886, Kuntze
2396: 2; Kuoh 1297, 7453: (1).
Matsuda Gram. 10, T-381: (2); Merrill 6483: 1.
Ohwi 2363, 2485, 3548, 3623: 1; Oka 35891: 2; Okamoto 32: 1.
Petelot 8025: 2.
Santos 7854, Shimizu & Chuang 20182, Shimada 5103-B: 1; Van Steenis 8660,
9135, Steward ef al. 514, Sun Yat Sen Univ. 51196: 2.
Tagawa 389, 3856: 2; Tamura & Koyama 23353: 1; Tanaka 15970, 25701, Tateoka
9389, 9395, Treutler 486: 2.
De Wilde & De Wilde-Duyfjes 16371, 16816:2.
Acknowledgements
This revision was started by Ms. H. M. Korthof during a course in advanced
Angiosperm taxonomy at the Rijksherbarium, Leiden. It was finished by Dr. J. F.
Veldkamp. The Keepers and Curators of BM, BO, G, K, KYO, L, MAK, NY, SING,
TI, UC, US, and W are gratefully acknowledged for making available the necessary
materials. From LE the type of Ag. zenkeri Trin. was received for inspection. No
specimens could be found in P by JFV and in PNH by Dr. J. V. Santos (Manila), but
Aniselytron & Deyeuxia in SE. Asia 223
Dr. Santos did have one in his private collection (dupl. in L now), from which Mr. L. M.
Gregorio prepared the drawing, which is reproduced here with their kind permission
and we are most grateful. Dr. T. Tateoka, Tokyo, kindly allowed us to cite his un-
published chromosome count for the Japanese form of An. treutleri.
Literature Cited
Bor, N. L. (1960). The grasses of Burma, Ceylon, India and Pakistan: 392, 394. Oxford.
Ettingshausen, C. von. (1893). Uber fossile Pflanzenreste aus der Kreideformation
Australiens. Sitzungsber. Kais. Akad. Wiss. Wien, Math-Naturw. Classe 102, 1: 135, 147.
. (1895). Beitrage zur Kentniss der Kreideflora Australiens. Denkschr. Kais. Akad.
Wiss. Wien, Math-Naturw. Classe 62: 12, t. 1. f. 10.
Hackel, E. (1907). Gramineae novae. II. In Fedde, Repert. 3: 241-244.
Hitchcock, A. S. (1934). New species of Aulacolepis and other grasses. J Washington
Acad. Sc. 24: 290-291.
. (1936). Botanical results of the Archibold Expedition. 1. Papuan grasses collected
by L. J. Brass. Brittonia 2: 117.
Honda, M. (1927). Revisio gramintim Japoniae. XIV. Bot. Mag. Tokyo 41: 641.
Hooker, J. D. (1896). Flora of British India 7: 269. London.
Jansen, P. (1953). Notes on Malaysian grasses. III. Acta Bot. Neerl. 2: 363-265, f. 1.
Keng, Y. L. (1958). Two new genera of grasses from China. J Washington Acad. Sc. 48:
117-118, f. 1.
Kirk, T. (1897). Trans. & Proc. New Zeal. Inst. 29: 497, t. 44.
Kuntze, O. (1891). Revisio genera plantarum 2: 780. Leipzig.
Merrill, E. D. (1910). In E. D. Merrill & M. L. Merritt, The flora of Mount Pulog. Philip.
J. Sc. 5, Bot: 328-330.
Ohwi, J. (1933). Symbolae ad floram Asiae orientalis. 9. Acta Phytotax. & Geobot. 2:
161-162.
. (1935). Symbolae ad floram Asiae orientalis. 11. Acta Phytotax. & Geobot. 4: 30.
. (1941). The tribe Agrostideae of Japan, excluding the genus Calamagrostis. Bot.
Mag. Tokyo 55: 361.
. (1956). Notes on some plants from Japan and its neighbours. (2). J. Jap. Bot.
51-0137;
Stapf, O. (1895). Deyeuxia treutleri Stapf, in Hook., Icon. Pl. 24: t. 2396.
Wilson, K. L. 1983. Publication of superfluous names. Taxon 32: 644-645.
Zotov, V. D. 1971. Simplicia Kirk (Gramineae). New Zeal. J. Bot. 9: 539-544.
viviemed) saxo lode b) qotteatiers eet
bhew thi tive eur scar eningas ie stoi,
Sh 2h vot Ste pret tose
vi VL Te tht! hime “A a
fort enmtepea.$ itis leone
, i a. th Pa japouge
scot a, Aah ae bet } Ace iB Ys . aie baad "OOeIE ES
Ge v4 mart” Alp. Fay Be “wh
ot eet ee | uli GRAD RY. ae ff
=) wwe NM Ge he: Sak ‘weg
7) ARS ea hie et macy. TUX ;
2 fd- a D? ¥)
wide = pee sohbet jd 4] seven swontmiore) .
jes i” tom en caukh lo nreKye wll wa @4
ee reliesina © - Is
a sao ht ayes Toru it ND sires i
ites: Spay ‘aT
avntifes’ wns PVaht 4 neon tr
ae 6 Ge aabusnasalary leh
coher J . aibnl sears Np matt
a & aml oa ah
\
\ ae mot cas, i Ferg Wait OWT
Se fad: Anat ae sath}
4 PSG ul bru al Shs
mi BPS worwonuley mere nd
t seml(al 4S line a aint ona
DEE
aA) | areas: nel.
wrlte -sokA fumoth Ge seloderge (engi
- »
Zz
ie A iraia i hi ere:
¥ er, Omir ta wear enh, ath Bly
Ide th ae
= i. y siycak surelg ane) 00 eer dat ui ;
Wey toot oy bget?t bei duet A an) & y
ets
i hem
La CF yung) sony mNenteae te. Bolimalide 901
Sih nll oa nite EB 1p
4 Di [ents : YO 1 MARS
knowles 1 ey makony
ie tonto Tri aan rispalved fer insp
-5 and in PNY by Dod. ¥, Sagas i
Preliminary Pollen Study of the Oleaceae in Malesia
R. KIEW
Department of Biology, Universiti Pertanian Malaysia, Serdang, Selangor, Malaysia
Abstract
Preliminary work on pollen of the Oleaceae shows that it is of limited diagnostic value at the generic
level. Small pollen grains (10-18, diameter) are recorded for Malesian species of Chionanthus, Myxopyrum
and Olea. The size of pollen grain is associated with style length.
1. Pollen morphology
The pollen morphology of the Oleaceae is reticulate and tricolporate (Wodehouse,
1965; Erdtman, 1971). A preliminary survey of Malesian species using JEOL 35 SEM,
where the pollen grains were acetolysed before coating with gold, confirms this and
also shows that there is variation between species for characters of the muri (whether
beaded or smooth, high or low) and for the size of the lacunae (Table 1). The large
lacunae are conspicuous under light microscopy in several species of Jasminum and
in Nyctanthes arbor-tristis (personal observation e.g., and in Ligustrum (Wodehouse,
ibid).
Some characters are constant for a genus eg., all the species of Olea examined
have large lacunae with spinules inside (Plate 1f); the muri of Myxopyrum are smooth
(Plate 1d & e) and the muri of Chionanthus are high (Plate la & b). However, since
only a single character is constant, a few species of different genera share the same
characters, e.g. Ligustrum confusum and the pollen of male flowers of Olea decussata
and the large grains of Myxopyrum ovatum all have large lacunae with high, smooth
muri (Table 1). Pollen morphology cannot therefore be used infallibly to identify
oleaceous pollen to genus. However, from this preliminary study certain character
combinations do indicate generic identity e.g., small lacunae with low, smooth muri is
characteristic only of Myxopyrum. A larger sample is needed to confirm this.
Variation of these characters may also occur within species, for example pollen
from the male flower of Olea decussata has smooth, not beaded, muri, the large pollen
grains of Myxopyrum ovatum have high muri instead of low (Plate le), and different
samples of Chionanthus ramiflorus differ in the size of the lacunae and whether the
muri are beaded or smooth (Table 1).
In addition, for the Oleaceae, pollen morphology also has limited value for
identification at the family level as reported by Erdtman (1971): that several other
families, such as Caprifoliaceae (Viburnum) and some species of the Celastraceae have
very similar pollen to that of the Oleaceae. As such, pollen morphology is of limited
taxonomic value above the species level in the Oleaceae.
Wodehouse (1965) suggested that the low muri of pollen grains of Fraxinus,
compared with the other genera he examined, was related to its pollination by wind.
Myxopyrum shows this same character but is very unlikely to be pollinated by wind
as the anthers are almost sessile and are included within the corolla tube.
223
226 Gard. Bull. Sing 37(2)(1984)
Table 1. Morphological characters of pollen of some Malesian Oleaceae
Species Lacunae
+ beaded — smooth + high — low} + large — small
}
+1 &S+rtttt
(Oleatibrachiatdienc-coasteen eee
QO} PQNiculaltiepaneasee eee ees
O. decussata (male flowers) .........
(bisexual! flowers)! ...........------
Chionanthus enerve ...........0...000+
(CL TU GHID US ssconendncnaccceanncscoosne
GR DPIUTIOTUS Reece ceeeeeec eee eee
Ligustrum CONfUSUM .......6.6002000-
Myxopyrum COriaCeumM ........ 2.0.00
M. ovatum (small grain) ............
(largegrain)) 32... 223edseneeececeeeels
+
tee + i +s
2. Size of Pollen
Wodehouse (1965) recorded the range of pollen grain diameter in the Oleaceae as
between 19.5 and 30 » and Erdtman (1971) gave the range of the longest axis as
between 20 and 63 y. These both fall within the medium-size class of pollen grains.
Wodehouse (ibid) found that most angiosperm pollen grains fall within the 20 to 40
p diameter range.
This study shows that while most of the genera of the Oleaceae do fall within this
range, the pollen of Chionanthus, Myxopyrum and some species of Olea is much
smaller, between 10 and 17 » diameter (Table 2) i.e., fall within the small-size class of
Wodehouse (1965) and Erdtman (1971).
Muller (1979) discussed possible causes for differences in size of pollen grains and
reported cases where pollen size correlated with flower size, style length, anther length
and latitude or altitude.
For the Oleaceae, the relationship is conspicuously between style length and pollen
size (Table 2). Genera with sessile stigmas or styles up to 1 mm long produce pollen
in the small-class size: Chionanthus (10-17.5 « diameter), Myxopyrum (13-16 y) and
some Olea species (12-15 y). Genera with styles 1 mm or more have in general pollen
grains larger than 20 yp. The exception is Osmanthus scortechinii with style length bet-
ween | and 4 mm but with pollen grains 17.5-22.5 » diameter.
Baker & Baker (1979) also found that for 147 species with style lengths of 10 mm
or longer they investigated, the mean pollen diameter was significantly larger than that
of their shorter-styled counterparts.
The relation between pollen size and style length is also found in heterostylous
species. Ganders (1979) recorded that in 50 out of 55 heterostylous genera, the thrum
anthers produce larger pollen than the pin anthers. (It is the thrum pollen that has to
grow down the longer pin style). It was first suggested by Darwin (1877, cited by
Ganders, 1979) that pollen grains that need to produce long pollen tubes to grow down
long styles would need greater food reserves and would therefore be larger.
The relation between flower size (corolla length) and pollen grain diameter is less
obvious and flowers longer than 10 mm (Jasminum, Forsythia, Nyctanthes and Syringa)
also have longer styles. For Chionanthus, where all species have sessile or short
stigmas, there is a considerable difference in corolla length between the temperate
species, C. retusus and C. virginicus with flowers more than 10 mm long, and the
‘tropical species with small flowers less than 5 mm long, but the range in pollen size
overlaps between these species with large and small flowers (Table 2). In addition,
Plate 1.
3b bBbE
bHbO O8 to .o0u
Pollen of Some Malesian Oleaceae
a: Chionanthus pluriflorus; b: C. ramiflorus; c. Ligustrum confusum;
coriaceum; e: M. ovatum (large and small grains); f Olea brachiata
oo
a
{a
Myxopyrum
297
228 Gard. Bull. Sing 37(2)(1984)
Table 2. Pollen Grain Size in the Oleaceae
SPECIES DIAMETER (u) STYLE LENGTH COROLLA
MEDIUM-SIZED POLLEN (MM) LENGTH (MM)
Menodora integrifolia ............... 63 AI REN ios ancontieeniceys |)-cas See eee eect
Jasminum crassifolium ...........++- (37.5) 43.5 (50) 35-55
ill Men eaeeeeaepeacsadt ne taeenee 46> 2 Wi U8 2a Ses oct aan all eset eee
Uh EPO DLL. casccosdneppanansaeesoo: (30) 34.5 (37.5)! 33
J. SQINDAC:. SR ee oo so (27.5) 29.5 (32.8)? 33
(B3):(90)* | Rudsenddccsadecbasseemetee |e checs sence eee
Nyctanthes arbor-tristis ............. 35-58 27-30
SCHYEDEraaNOISIIta aeaeeenee eee 353 Wdeaclvc'sssueBiucsectoes seri! neces eee eee
SAGE WAGE. ssacopacconnaaoa secs 263°, aioe 0 ea 8-15
Ligustrum CONnfUusuUM .........0.000+ (20) 22.5 (25) 3-4
Es SIOMCIOIUIN *. aisasmus cas ene (25) 28.3 (32.5) 2-3 (4)
L. tbota (28): 30.5.(B2)§ 2° - {| Resceickeooscceeeeseaec sol Weeceoeeaee een
EX Ovalifolium@en.ccoe ree oe 285-31 eRe. Seen 8
EASINGNSE. i aiuesc deaestass eee eos 25 2-3
De Vuloares 3a ene enero 28.5° 3
Fontanesia phillyraeoides ........... 7) | Sn in (PERE ner eer reper ral lider obozcubacckccoucoecs
Forsythiad SUSPeNs@ ........0.0..00005- (19.4) 28.5 (37.6)° 19
Fraxinus americanus ..............+.. )): (PEPER Een Ae sa Coenen | ssotnho soncodsdcoosexoe
ENCOriaCea sch nee nena eee eer y) | Jn WP PERRErrr EES sa acconces| | bovoscncddcadcnenoesces
Eexcelsior pee eae een ee 223 2-3
Osmanthus scortechinii ............- (17.5) 21 (22.5) 1.5-2.5
Phillyrea angustifolia ................ 2057 =, cscc RBs. cc. d ese s oc ei| c eee eee eee
SPECIES Diameter (,) STYLE LENGTH COROLLA
SMALL-SIZED POLLEN (MM) LENGTH (MM)
Olea CurOpaea Race correct eee ee 79h
OM DIACHIGIG sac sonsans ons we nntee eh ace 12.5 & 26
O. decussata (male flower) ......... 7-9 & 13
(DISExuale TlOWen)ecesesceeeeceaeeee 13
OO! JQVANICA ccs cotaee rece nee oes 12.5
O} DaNICUIAIG. Reeccsencsecenescectene 12-15
Myxopyrum Coriaceum ............++ 16
ME OVQIUIN AE ces aeteene a caceeeeee 13-15.5 & 21
Chionanthus retusus ..........0..0.+- (13) 14.1 (16)
GC Vinginicust..csest sce ee (13) 14.3 (16)°
GOACUFYICONDUS pace caicwa acess eeetee eee 12.5
GON wrecsirosenseevcinnictsnsinseeeoascs 17
CEIGAOFUS. esac caus ease neoet meee (10) 11.25 (12.5)
GS PlunflOvUsraieeccosescnecsre teers (11.25) 13.5 (17.5)
GS POFCONUS) scosicceucacceveseeeseceeser (11.25) 12.5 (13.75)
GOUTANTLOTUS cosh ch oomneace cease sen. 10-25
Gy PUDIOVERIA cucscc ces cacceeeeeenoet (10) 11 (12.5)
GUTUPICONMS Pesce ecco eee ree 10
: long-styled; 2: short-styled; 3: Erdtman; 4: Raman ef al; 5: Sohma; 6: Wodehouse)
~~
=)
Fraxinus, Ligustrum and Osmanthus, which have small flowers, have medium-sized
pollen in the same range as the large-flowered genera (Table 2).
Specimens of some species have pollen grains of two sizes, the difference in size
being almost double. This phenomenom I observed in Olea brachiata (with pollen
grains 12.5 and 26 » in diameter), O. decussata (with the majority being larger, 13 p,
than the other grains, 7-9 » in diameter), and in Myxopyrum ovatum (the majority
were 13-15.5 », with some larger, 21 y» Plate le).
Size differences have also been observed by Devi (1975) for Jasminum callophyllum
where he found the size difference corresponded to the state in which the grain was
shed — the smaller grains were 2- or 3-celled, the larger grains were multi-celled.
Pollen study of Malesian Oleaceae 229
Differences in size can also be expected to be observed in the heterostylous genera
of the Oleaceae: Forsythia, Jasminum, Nyctanthes and Schrebera (see above). Unfor-
tunately, for pollen sizes given by other authors it is not recorded whether these are
for the long- or short-styled flowers, so it is not possible to ascertain whether the
range of size within species of these genera may be ascribed to flower type or difference
between plants of the same flower type.
Polyploidy can affect the pollen size between individuals of a species. For example,
Sohma (1972) noted that tetraploid plants of C. retusus from Taiwan are almost twice
the size of normal diploid grains. Raman ef a/. (1970) reported a similar case for
Jasminum grandiflorum where triploid plants produced some grains twice the size of
those of diploid plants (Table 3). However, between species of Jasminum there is no
correlation between size of grain and level of ploidy (Table 3), the tetraploid species
falling within the range of other diploid species. The much larger grain of cv. ‘Iruvatchi’
of J sambac suggests that it is also polyploid.
Table 3. Level of ploidy and diameter of pollen grains in some species of Jasminum
(data from Raman ef al. 1970)
Jasminum Species Level of Ploidy Diameter of grain (u)
TYE TO in cCCC CER EE EEE OEE ECE EEEEE PELE E CEES 2n 33-52.8 (42-85.8 cv. ‘Iruvatchi’)
UICUNEITI Fae Se 58 nec Soa odes cee osteees ass 2n 29-52.8
ITEM ENTIT rcenin nae c= ann ats cs0tenndesscss 2n 33-46.2
3n 29-102
CONTMIIIMS en tena cians ceesesescascasecescesaes 3n 33-52.8
TECUINT IM ne eens sossse soeesscctssreceas.ses 4n 39.6-49.5
Variation on a geographical basis is small (Table 4) for wild Chionanthus material,
although it is larger for thecultivated Nyctanthes arbor-tristis. This latter species is also
heterostylous and its range in size may be ascribed to heterostyly.
Table 4. Variation of pollen grain diameter on a geographical basis
Species Locality Pollen Grain Diameter ()
Chionanthus ramiflorus Borneo 10-12.5
New Guinea 10-15
C. retusus China 13-15?
Japan 1416
Taiwan 13-15°
Nyctanthes arbor-tristis Java 42.5-52.5
Malaya 35-55
Singapore 3
Thailand 58!
(1: Erdtman, 2: Rao & Leong, 3: Sohma).
Within species, variation in size of the pollen grain in the Oleaceae can therefore
be attributed either to heterostyly, polyploidy or the state of the development of the
pollen grain when shed. Between species, variation in pollen size appears to be related
to style length. The occurrence of small pollen grains in the Oleaceae is here reported
for the first time and it is found in species of Olea, Chionanthus and Myxopyrum, all
of which genera have sessile stigmas or styles less than 1 mm long.
Acknowledgements
The author is indebted to A-AUCS for a research visit to the University of New
230 Gard. Bull. Sing 37(2)(1984)
England, Australia, where the SEM study was carried out and to the late Dr Jan
Muller of the Rijksherbarium, Leiden, for stimulating and helpful discussions on the
pollen of the Oleaceae.
APPENDIX
Material examined
1. SEM. Chionanthus enerve (Steenis) Kiew S30433; C. pluriflorus (Knobl.) Kiew
RK779; C. ramiflorus Roxb. NGF5969; Ligustrum confusum Decn. RK1I88; Myxopyrum
coriaceum Bl. S30037; M. ovatum Hill NGF46864; Olea brachiata (Lour.) Merrill Ahmad
s.n. Kuala Lumpur; O. decussata (Heine) Kiew $225/6 (male flower), Chew 939 (bisexual
flower); O. paniculata R. Brown NGFI1948.
2. Light microscopy. Chionanthus curvicarpus Kiew SAN35826; C. laxiflorus Bl.
SAN49386; C. pluriflorus (Knobl.) Kiew RK779; C. porcatus Kiew SAN43052; C.
ramiflorus Roxb. SAN64322; C. rubrovenius (Elmer) Kiew Ramos & Edano 38584;
C. rupicolus (Lingels.) Kiew Jacobs 9292; Jasminum crassifolium Bl. Elmer 20762;
J. multiflorum (Burm. f.) Andr. RKi237; J. sambac (L). Ait. RK1238; Ligustrum
confusum Decn. RK188; L. glomeratum Bl. SAN28544; L. sinense Lour. SA183;
Myxopyrum ovatum Hill NGF46864; Nyctanthes arbor-tristis L. Junghuhn 33
(Java), RK100] (Malaysia); Olea brachiata (Lour). Merrill T & P244; O. javanica
(BI.) Knobl. Kds10062; Osmanthus scortechinii K. & G. de
Literature Cited
Baker, H.G. and I. Baker (1979). Starch in angiosperm pollen grains and its evolutionary
significances. Amer. J. Bot. 66: 591-600.
Darwin, C. (1877). The different forms of flowers on plants of the same species. Murray,
London. (cited by Ganders, F-R.).
Devi, H.M. (1975). Embryology of Jasminums and its bearing on the composition of
Oleaceae. Acta Bot. Ind. 3: 52-61.
Erdtman, G. (1971). Pollen morphology and plant taxonomy. Hafner, New York.
Ganders, E.R. (1979). Biology of Heterostyly. New Zealand J. Bot. 17: 607-635.
Muller, J. (1979). Form and function in angiosperm pollen. Ann. Missouri Bot. Gard.
66: 593-632.
Raman, K.R., K.M.A. Khan and A. Shanmugam (1970). Palynological aspects of some
species and varieties of Jasminum. J. Palynol. 6: 73-77.
Rao, A.N. and EN. Leong (1974). Pollen morphology of certain tropical plants.
Reinwardtia 9: 153-176.
Sohma, K. (1972). Palynological notes on the genus Chionanthus (Oleaceae). Sci. Rep.
Tohoku Univ. Ser. IV (Biol.) 36: 203-209.
Wodehouse, R.P. (1965). Pollen grains. Hafner, New York.
Volume 37
Basionyms and synonyms appear in italics. Page numbers in italics indicate the presence of
illustrations.
ACACIA PYCNANTHA and soil compaction study:
influences of level of artificial soil
compaction 82
on
amended clay loam 87, 89
dry bulk density 84, 86
root penetration 86, 90
shoot dry matter 84
development of shoots in treated clay loam 88
total porosity 86
rate of oxygen diffusion 82
moisture content in improved clay loam 84,
87, 87
moisture content in straight clay loam 8&6,
87
methods of artificial compaction 83, 84
soil strength as a limiting physical attribute 84
methods to reduce soil strength 83
compacting methods of straight clay 82
Aglaia iloilo 62
Agrostideae 213
Agrostis 213, 214
(Agrostis) 12, 218
debilis 220
neodebilis 220
pleiophylla 218
rigidula 215
tenuis 219
zenkeri 219
Amorphophallus 199, 200
Anisachne 214
gracilis 218, 219
Aniselytron 213, 214, 215, 222
(and Deyeuxia) index of collectors 222
agrostoides 214, 215, 216, 22/
tabulated measurements of Santos 7854,
220
clemensae 217, 218
epileuca 222
japonicum 216, 218
milioides 217, 218
petelotii 218
pseudopoa 217, 218
treutleri 215, 216, 217
diagnoses of collections 217, 218
Japanese form 223
var. japonica 217
var. milioides 217
var. treutleri 217
Antelaea Gaertn. 54
azadirachta 62
canescens 62
Javanica 57
Antheeischima 1, 3
excelsa, 12
Anthelia (‘Anthelea’) canescens 62
Anthurium 199
Artagrostis 213
Arum colocasia 200
esculentum 200
Aulacolepis 213, 214, 215
agrostoides 216
var. formosana 216
clemensae 214, 217
epileuca 214, 222
Japonica 216, 218
milioides 214, 217
petelotii 214, 218
pseudopoa 214, 217
treutleri 215,216, 217, 218
var. japonica 216
var. milioides 217
AXILLARY BUDS
Acacia auriculiformis 68
Anacardium occidentale 74
Annona muricata 68, 74
reticulata 68, 74
squamosa 68, 74
Artocarpus 69
Bauhinia 69
Barringtonia asiatica 76
Bauhinia acuminata 75
blakeana 75
purpurera 75
Bixa orellana 70, 74
Brexia madagascariensis 77
Brownea capitella 75
grandiceps 65, 75
Callistemon citrinus 76
Calophyllum inophylum 75
Cassia 68
bakeriana 75
fistula 75
multijuga 75
sp. (hort. variety) 75
spectabilis 75
Cinnamomum iners 75
Citrus grandis 77
Coccoloba uvifera 73
Cochlospermum religiosum 74
Coffea sp. 69
Crataeva religiosa 74
Cratoxylon formosum 74
pruniflorum 74
Dalbergia oliveri 75
Delonix regia 68, 75
Dimocarpus longan Lour. var. malesianus 77
Diospyros discolor 75
Dryobalanops aromatica 75
Elaeodendron quadrangulatum 74
Elateriospermum tapos 75
Erythrina fusca 75
variegata 75
Erythrophleum suaveolens 68, 70, 75
Ervatamia dichotoma 74
Eucalyptus ficifolia 73
Eugenia 69
grandis 70, 76
javanica 76
421
232
Eugenia Cont.
longiflora 76
malaccensis 76
michelii 76
Ficus 69
benjamina 77
elastica 77
retusa 77
Filicium decipiens 77
Fragraea fragrans 70, 76
Gliricidia sepium 68, 75
Gnetum gnemon 75
Hibiscus tiliaceus 68, 76
Hopea mengarawan 75
Jacaranda obtusifolia HBK sp. rhombifolia 68,
74
Khaya grandiflora 76
Lagerstroemia speciosa 76
Lithocarpus urceolaris 68, 75
Mangifera indica 74
Melaleuca cajuputi 76
Michelia champaca 76
alba 70
Millettia atropurpurea 68, 75
Mimusops elengi 68, 77
Muntingia calabura 73, 77
Nephelium lappaceum 68, 77
number 68-70, 74-77
Palaquium obovatum 77
Peltophorum pterocarpum 68, 70, 75
Plumeria sp. 65
acuminata 74
Podocarpus 68
koordersii 76
neriifolius 76
polystachyus 76
Polyalthia longifolia 74
Pongamia pinnata 76
Psidium guajava 76
Pterocarpus indicus 76
Rhodamnia cineria 76
Rhus succedanea 74
Salix sp. 77
Samanea saman 68, 76
Sandoricum koetjape 76
shape 66-67, 69, 74-77
Shorea 68
curtisii 75
leprosula 75
sumatrana 75
size 70, 74-77
Spathodea campanulata 68, 74
Tabebuia pallida 68, 70, 74
rosea 784
spectabilis 74
Terminalia catappa 74
Triplaris americana 76
Vatica pallida 75
Aveneae 213
Azadirachta excelsa 62
indica A. Juss. 50, 56; 62, 63, 64
Azedara Raff. 55
speciosa 55
Azedaraca Raff. 55 and A. amena 55
Azedarach Dod 63
Gard. Bull. Sing 37(2)(1984)
Azedarach Mill. 54
commelini 60
deleteria 55
edullis, ‘edule’, 62
elegans 62
excelsa 62
florens Tourn. 64
fraxinifolia 62
nigra ‘nigrum’ 62
odorata 60
ramiflora, ‘ramiflorum’ 62
recisa, ‘recisum’ 62
sambucina 60
sempervirens 59
var. australasica 58
var. dubia 57
var. glabrior 55
tomentosa 62
vulgaris 55
Banisteria sericea 57
Brochoneura 121, 170
Caju lapia 6
Caladium 199
Calamogrostris
arundinacea var. brachytricha 220
brachytricha 220
brevipila 220
debilis 220
stenophylla Hand.-Maz. 222
varia var. longiaristata 220
Camellia 3
axillaris 1
Caria 4
Castanopsis-Quercus forest 9
Chionanthus
Malesian spp. 209, 210, 211
(Species)
callophyllus 210
curvicarpus sp. nov.
cuspidatus 211
elaeocarpus 209
var. elaeocarpus 209
var. globosus 209
insignis 209, 210
gigas 210
grandifolius comb. nov. 210
laxiflorus 211
macrocarpus 209, 210
macrocarpus sensu Kiew 210
oliganthus 211
platycarpus 210
porcatus 211
sessiliflorus 211
Chisocheton baccifera 62
patens 62
penduliflorus 62
tomentosus 58, 62
Cipadessa baccifera 50, 62
Closaschima 1, 3, 33
marginata & var. dasyophthalma 24, 26
lanceolata 26
obovata 26
ovalis 31, 33
vars. longifolia and minor 33
Crema glomerata 167
Compsoneura 121
Colocasia 199, 200
subf. Lasioideae 199
Colocasioideae 199
antiquorum 200
Index
Colocasioideae Cont.
esculenta 200 et seq.
var. antiquorum 200
esculenta 200
Penins. Malaysian cultivars 199, 200
grouping of, 200
keys 201, 204, 205
VERNACULAR NAMES: (see under Colocasia
esculenta); K. air 204, 205; K. banjar, 203,
204; K. bantan 205, 206; K. batang hijau 204;
K. batang hitam 204; K. belang hijau 205; K.
cina 204, 202; K. cincang wangi 200; K. dara
205; K. hutan 205; K. huma 205; K. kelapa
205: K. kubis 205; K. liar 205; K. lilin 205;
K. minjak 203, 204; K. nibong merah 204;
K. peladang 205; K. pinang 204, 206; K. putih
204; K. serakit hitam 204; K. serakit putih 204;
K. tongsan 205; K. udang 202, 204: K. songket
hijau 205, K. songket ungu 205
Cyrtosperma 199
Deyeuxia 213, 214, 219
abnormis 213, 218, 219
axilis 213
brachytricha comb. nov. 220
debilis comb. nov. 220
epileuca 214, 222
gunniana 213
mazettii nom. nov. 222
pseudopoa 217
pusilla 213
sylvatica var. brachytricha 220
treutleri 216
zenkkeri comb. nov. 219, 222
Dialyanthera 121
Dieffenbachia 199
Dipterospermum 4
Dysoxylon excelsum 62
nutans 62
Parasiticum 62
EMBRYOLOGY of Garcinia mangostana:
anther development 94
embryo development 98, 99, 101
embryo-sac type 96
endosperm formation & type 96, 98, 102
floral & fruit morphology 94, 95
fruit morphology 94
fruit season in Malaysia 93
megagametophyte 96, 99
megasporangium 96, 99
megasporogenesis 96, 99
microsporangium 94, 97
microgametophyte 94, 97
microsporogenesis 94, 97
ovule characters 191
phenology 183
pollination and fertilization 96
seed coat development 98, /0/
seed formation 96, 102
seed germination 100
seedling morphology 95
Endocomia 116, 118, 119, 120, 124, 130,
macrocoma 119
Eucalyptus savanna 56
Eucalyptus-Gironniera forest 9
233
Fraxinus ornus 63
Garcinia mangostana see EMBRYOLOGY
Gleichenia scrub 42
Gordonia 1, 2, 3, 9
in Malesia
index to scientific names 47
key to Malesian species 4-6
VERNACULAR NAMES: adekelp 9, alimp 9, bado
9, baif 9. benelemonde 9, bwabwa 9,
damak 24, dapiri 9, dimi 9, djirok bantjoh
31, entuyut, 36, iniaili 9, kawal-gugn 9,
kayu patjat 31, kayu kelat asam 41, kayu
kelat putih 41, kelak merah 28, kerkebo 9,
kilimdan 9, kimkaroo 9, koka kaber 9,
kuku 9, la, naningning 9, obah 11,
oOyvtungo 9, palembang putih 31, penagit 25,
rawali batu 11, eik 9, samak pulut 28,
samak samak 28, sawak pulot, 41, sjioe 9,
tawan 9, timor 9, titup 24, titup tiup 24,
tiup 24, toani 9, tokoi 9, totona rombo 9,
tugera 9, ubar lilim 31.
(SPECIES)
acuminata 12
amboinensis 6, 7. 9. 46. 47
benguetica syn. nov. 21, 23
borneensis sp. nov. 2, 3, 8, 11
brassii syn. nov. 6
brevifolia 45
concentricicatrix syn. nov. 26, 28
densiflora syn. nov. 42, 43
excelsalla 2 10:.10> 12
var. macrocarpa 12
var. sincapuriana 39
excelsa auct. non Blume 33
fragrans 2]
grandiflora /2, 13
grandifolia 11
grandis King 39
havilandii /4, 15
hirtella 16, /6
imbricata 2, 17, 18, 39
integerrima comb. nov. 19, 20
lasianthus 1, 3
lanceifolia 45, 45
lobbii 46
luzonica 21, 22, 23, 45
variants of, 23
maingayi 2, 23, 25, 39
marginata 24, 26
multinervis 2, 26, 27, 36
oblongifolia 2, 11, 15, 28, 29
ovalis 1, 2, 3, 30, 31, 33
forms of, 33
papuana syn. nov. (& incl.
vars. acuminata & montana) 6
penangensis 32, 33
polisana 2, 23, 34, 35
rumphii 6
sarawakensis sp. nov. 2, 3, 36, 37
scortechinii 3, 38, 38
singaporeana 2, 39, 40
sp. of Kobuski 6
sp of Burkill 21, 23
subclavata sensu Burkill 21, 23
taipingensis 41, 43
vulcanica comb. nov. 2, 19, 42, 44
var. vulcanica 42
var. buxifolia stat. nov. 43
welbornei 21
234
Gymnacranthera 119, 120
Haemocharis 3, 4
amboinensis 6
aromatica syn. nov.
(& excl. vars. minor &
longifolia) 31, 33
buxifolia 43
integerrima 19
aromatica syn. nov. (&
excl. syn. Lapl. semi-serrata
Mig.) 31, 33
marginata 24
ovalis 31
serrata syn. nov. 19
‘serrata’ 21
subintegerrima syn. nov. 31
vulcanica 42
Heath & peat swamps forests 134
Heteropterys laurifolia 57
Heynea trijuga 62
Horsfieldia Willd. 115, 116, 118, 119, 120, 121,
122, 123, 124, 170
distribution, 136, /37
VERNACULAR NAMES:
Amol 171, Bolong 171, Bolouchi 171,
Con na ham 175,
Lan ham 175, Malaboda 168; Me Tiong 175,
Luat nok 179,
Irie gaga 168; Pandekachoaphang 171, Ruk 168;
Rukghedhi 168; Lan Ham 175, Siltui 171,
Tjampaka se’long 168
Sect Bivalves Sncl. 127
Sect. Horsfieldia 126, 127, 136,
Sect. Irya 120, 121, 124, 126, 127, 128, 136, 137
subsect. Euirva Warb. p.p. (incl. H. irya) 127,
130
subsect Triva/ves pp. (excl. H. irya) 130, 132,
134
Sec. Iryaghedhi 121
Sect. Pyrrhosa 120, 121, 124, 126, 127, 128,
130, 132, 134, 136, 137
subsect. Bivalves Warb. incl. ser. Smithii &
ser. Globularia 127, 128
subsect. Eupyrrhosa 130
Sect. Orthanthera Warb. 126, 132
Sect. Trivalves
subsect. Orthanthera (Warb.) Sincl. 126
subsect. Triva/ves 132
(GROUPING OF SPECIES):
affinis-gp, spp. 61-67, 133
amygdalina-gp, spp. 2-5, 132, 125
carnosa-gp, sp. 69, 133
crassifolia-gp. sp. 68, 133
fulva-gp, spp. 52-54, 132
glabra-gp, spp. 97-100, 132, 133, 135
grandis-motleyi-gp, spp. 56-60, 133
clavata-gp, spp. 21-23, 130, 133
helwiggii-gp,. spp. 42-45, 130
irya-gp, sp. 6: 128
laevigata-gp, spp. 24-41, 130
pallidicaula-group, spp. 47-50, 132, 133, 134
parviflora-gp, spp. 7-16, 128
polyspherula-gp, spp. 71-82, 132, 134, 135
sabulosa-gp, sp. 46, 132
sterilis-gp, spp. 70, 134
triandra-gp, sp. 51, 133
Gard. Bull. Sing 37(2)(1984)
Grouping of Species Cont.
sessilifolia-gp, sp. 55, 132
spp. 83-93-gp, 134
xanthina-gp, spp. 94-96, 134
(SPECIES)
affinis 125, 133, /35
ampla 119, 130
ampliformis 119, 122, 130, /3/
amplomontana 124, /35
amygdalina 119, 128, 136, 172, 176, 177
var. amygdalina 177, 178, 179
var. lanata de Wilde var. nov. /29, 177, 179
androphora 124, 134, /35
angularis 120, 127, 130, /3/, 136
ardisiifolia 121, /29
aruana 124, 130, /3/
atjehensis 125, /3/, 133
australiana /29
basifissa 120, 130, /3/
basifissa 125, 132, 133, 134, 135
brachiata 118, 134, /35
bracteosa \77
carnosa 125
clavata 120, 127, /29
coriacea 125, /37
corrugata 125, 130, /3/
costulata 125, /37
crassifolia 120, 13, /35, 136
crux-melitensis 117, 120, 121, /29
decalvata 121, 130, /3/
disticha 118
endertii 125, 133, 134, /35
flocculosa 125, 133, /35
fragillima 125, 134, /35
fulva 125, /3/, 133
glabra auct. non (BI.) Wall. 177
glabra (BI.) Warb. 119, 121, 127, 130, 134,
EKG, VA/S5 Was TAS WA
var glabra /37
var. javanica 137
grandis 125, 133, /35
gracilis 125
hainanensis 170, 171, 172
hellwigii 125
var. hellwigii /3/
hirtiflora 125, 134, /35
inflexa 124, 127, 129
iriana 124, 130, /3/
irya 116, 118, 120, 121, 122, 126, 127, 128, 129,
130, 133, 136, 169, 170
distribution map /37
distribution /37
iryaghedhi 117, 119, 122, 123, 127, 129, 132, 136,
167
kingii 122, 129, 170, 171, 172, 773
laevigata 116, 119, 122, 130, 131, 136
lancifolia 121, 125, 130, /3/
laticostata 125, /35
leptantha 125, /3/
longiflora de Wilde sp. nov. 128, /29, 132, 136
macilenta 125, /35
macrocoma 116, 118, 124, 130
macrocoma s.I. 120, 124, 172
macrocoma-complex 116
macrothyrsa 125, /37
majuscula 125, 135, /37
moluccana 127
var. moluccana /29
montana 125, 133, /35
motleyi 125, 133, /35
Index
Species Cont.
nervosa 125, /35
obscura 120, 134, /37
obscurinervia 124, /29
obtusa 125, /35
odorata 124, 167
olens 124, 127, 129, 136
oligocarpa 118, /35
palauensis 124, 128, /29
pallidicaula 118, 122, 133
var. pallidicaula /3/
pachycarpa 125, 130, /3/
pachyrachis 125, /35
parviflora 116, 119, 120, 121, 127, 129
Paucinervis 125, /35
parviflora 116, 119, 120, 121, 127, 729
paucinervis 125, /35
penangiana 125, /37
pilifera 125, /3/
polvantha 116
polyspherula 121, 122
var. polyspherula /35
prunoides 177
psilantha 125, 130, /3/
pulcherrima 125, 133, /35
pulverulenta 120, 130, /3/, 132
punctata 125, 134, 135, /37
punctatifolia 119, 122, /37
ralunensis Warb. 125, /3/, 132
reticulata 125, 133, /35
ridleyana 125, 134, /35
rostrata 120
tufo-lanata 125, /35
sabulosa 118, 119, 123, 132
samarensis 124, /29
schlechteri 124, 130, /3/
sepikensis 122, 127, 129, 136
sessilifolia 119
sinclairii 122, 130, /3/
smithii 118, 128, /29
sparsa 125, /3]
spicata 118, 119, /29
splendida 122, 135
squamulosa 120, 122, /29
sterilis 120, 130, 132, 133, 134, /35
subalpina 125, 178
subsp. subalpina /37
subsp. kinabaluensis 160
subtilis 124, 130
var. rostrata 120
var. subtilis 122, /3/
sucosa (King) Warb. 118, 133
subs. bifissa 133
subsp. sucosa /3/
superba 125, 130, /3/, 133
sylvestris 117, 119, 127, 130, /29, 132, 170
talaudensis /29
tenuifolia 125, /35
tetratepala 170, 171, 172
thorelii 118, 120, 122, 128, 129
tomentosa 125, /34
triandra 120, 130, /3/, 133
tristis 125, /3/
tuberculata 121, 130, /3/
valida 125, 134, /35
wallichii 122, 132, 133, /35, 170
whitmorei 119, /33
xanthina 118
subsp. xanthina /37
Hypericum lasianthus 1
235
Jacaranda fraxinifolia 61
Knema 117, 119, 120, 122, 123
plumulosa 172
Koelreuteria elegans 62
Laplacea |, 3, 4
amboinensis 6
aromatica syn. nov. 31, 33
incl. vars minor & longi-
folia 31, 33
buxifolia 43
integerrima 19
marginata 24
ovalis 31
Sarasini 46
semiserrata 33
serrata syn. nov. 19
sepeciosa |
subintegerrima syn. nov. 31, 33
vulcanica 33, 42
Lasianthus Adans. 1, 3
Leptospermum scrub 19
Lignum mucosum 6
Lindleya 3
Linociera
beccarii 209, 211
callophylla 210
cuspidata sensu Merril (non Kiew) 211
elaeocarpa 209
insignis pp. 209, 210
grandifolia 210
macrocarpa 209
palludosa 210
Malpighia dubia 57
Melia 49 ef seq., 53, 54, 55
COMMON/VERNACULAR NAMES:
abor pareiso & arbor sancta 50; Azadiracht 63;
Azadraxt (also azadiraxt) 50; Azedarach 50;
Azedaraeth 50; bead-tree 50, 59, 63;
Chinaberry 59; kirikohomba (form of Persian)
Lilac) 63; k’u-lian 52; Mana Ash (= Fraxinus
ornus) 63; mindi kechil 50, 59, 63; neem or
neem tree 50, 56, 63, see also Azadirachta
indica; panukohumba 63; Persian lilac 50, 5/,
53, 54, 58, 59, 60, 63, 64, see also under Melia;
azedarach var. glabrior, forms of, 58; Texas
Umbrella Tree, 51, 54, 61, see also under Melia
azedarach cv. Umbraculifera; To sedan 51,
§2, 59; White Cedar 53, 57-58, tropical
Australian forms of, 49, 53; (is) zanzaliqu 50
(SPECIES)
angustifolia 52, 60
argentea 57
arguta 60
australasica 58
australis 57
azadarach 49
azadirachta L. 50, 62, 63
azedarach L. 50, 52, 53, 54; 56, 63
lectotype proposed 65
azedarach sensu Blanco 58
azedarach sensu Lour. 60
azcdarach sensu stricto 59
azedarach
cultivars 61
cultivars of wild trees 57-60
Chinese, names of, 54, 59
Indian, names of, 54, 59, 60
236
Common Vernacular Names Cont.
cv. Floribunda 61
cv. Umbraculifera 61
ecology of wild trees 56;
systematic arrangement 55;
past & present uses 56 see a/so under Melia
japonica
toxic fruits of, 52, 60
var. acuminatissima 50; 60
var. australasica 49, 53; 58
var. cochinchinensis 53; 58
var. floribunda 61
var. ‘gigantea’ 58
var. glabrior 53, 55
var. glandulosa 53; 58
var. incisa 50, 60
var. japonica 59
var. javanica 53, 58
var. sambucina 60
var. semperflorens 59
var. sempervirens 51, 52; 59
var. squamulosa 53; 58
var. subtripinnata 51; 60
var. toosendan 59
subvar. toosendan 59
var. umbraculifera Knox 61
var. umbraculiformis Hort. 61
f. albiflora 59
f. umbraculifera Knox 61
baccata 62
baccifera 62
birmanica 49, 58
bogoriensis 49, 58
bombolo 53
bukayun 60
bukheim 60
candollei 49, 57
chinensis 59
cochinchinensis 60
commelinii 60
composita 49, 57
comp var. biglandulosa 58
comp var. cochinchinensis 49, 58, 60
dubia 49, 53, 57
elegans 62
baccifera 62
excelsa 62
flaccida 58
floribunda 51, 61
var. sempervirens L. 51
florida 55
fraxinifolia 62
guineensis 52, 60
hasskarlii 59
indica (A. Juss.) Brandis 62
iloilo 62
integerrima 62
japonica G. Don 59
var. albicans 59
var. albiflora Mak. 59
var. semperflorens Mak. 59
var. toosendan 59
Japonica Hassk. 52, 59
cited in Japanese poetry 52
past economic uses in China 52
javanica 59
koetjape 62
latifolia 62
lobata 60
montana 62
neilgherrica 62
Gard. Bull. Sing 37(2)(1984)
orientalis 60
parasitica 62
parviflora 62
pendula 62
penduliflora 62
pinnata 62
pubescens 62
pumila 62
robusta 49, 57
sambucina 50, 60
sempervirens (L.) Sw. 59
sempervirens sensu Roxb. 60
‘sempervirens’ 52
superba 49, 57
tomentosa Roxb. 62
tomentosa sensu Migq. 58
toosendan 52, 59
var. semperflorens 52
f. albiflora 52
(hybrid) subvarieties japonica & toosendan
subvar. intermedia 52
volkensii 53
Melica latifolia 62
Milium 214
treutieri 216
MULTIPLE AXILLARY BRANCHING at
nodes of Symingtonia populnea:
Attim’s Model compared 105
development of higher order axillary lateral
meristems 107
factors influencing 105
frequency based on vertical orientation 106
frequency and pattern of branching between
leader shoots and lower limbs 109
compared with Acer saccharum 109
on apex of main axis /07
on main axis /06
vascular tissue in transv. sections /08
leader shoots branching pattern 107
twice-folded stipules 106
Munronia neilgherrica 62
pinnata 62
pumila 62
Myristica 119, 120, 122
Sect. Caloneura 130
Sect. Eumyvristica 124, 126, 130,
subsect. Horsfieldia 126
Sect. Horsfieldia 126, 127
Sect. Jrva Hook.f. & Th. 120, 121, 124, 127
Sect. /rva auct. p.p. 126, 130
Sect. Odorata Willd. 126
Sect. Pyrrhosa Bl. 124, 126, 130
(SPECIES)
amegydalina 177
floribunda 177
glabra auct. non BI. 177
glabra Bl. 130
glomerata Miq. 167
glomerata Thunb. 167
horsfieldii (‘‘horsfieldia’’) Bl. 126, 127, 167
irya 127
irvaghedhi 167
Javanica 127
Index
Species Cont.
kingii 170
kurzii 177
notha auct. non Wallich 167
superba 130
Nabiasodendron 2, 4
Nephelium lappaceum see REPRODUCTIVE
BIOLOGY
Nothofagus forest 9
Olea platycarpa 210
OLEACEAE POLLEN OF MALESIAN SPECIES: 225 ef
seq.
material examined 230
tables 226, 228, 229
(TAXA)
Caprifoliaceae 225
Celastraceae 225
Chionanthes 225, 226, 229
enerve 226, 228
pluriflorus 226, 227, 228
ramiflorus 225, 227
“wild spp.’’ 229
retusus 226, 228, 229
virginicus 226
Forsythia 226, 229
Fraxinus 225, 228
Jasminum 225, 226, 229
callophyllum 228
grandiflorum 229
sambac 229
Ligustrum 225, 228
confusum 225, 227
Myxopyrum 225, 226, 229
coriaceum 226, 227
ovatum 225, 226, 227, 228
Neoaulacolepis 214, 215
clemensae 217
Japonica 216
petelotii 218
Nyctanthes 225, 226, 229
arbor-tristis 225, 229
Olea 225, 226, 229
brachiata 226, 227, 228
decussata 225, 228
paniculata 226, 228
Osmanthus 228
scortechinii 226
Schrebera 229
Syringa 226
Verbenum 225
Orthanthera 170
Parkia biglobosa 52
Phelima noronha 167
Philodendron 199
Pistice 199
Ploiarium oblongifolium 28, 31
Poa 213, 214
milioides 217
papuana 222
wiselii 222
Polyspora 1, 3
Pycnanthus 170
Pyrrhosa 124
horsfieldii 67
odorata 167
Raphidophora 199
Rennellia in the Malay Peninsula 193, /94
198
key to species and varieties 195
key to distinguish from Morinda 193
(SPECIES NAMES)
elongata 193, 194, 195, 198
‘longiflora’ 198
morindiformis 197, 198
paniculata K. & G. 193, 195, 196
var. condensa var. nov. 195, 196
var. paniculata 195, 196
speciosa 193, 195, 196
speciosa sensu Corner p.p. 195, 196, 197
speciosa var. elongata 193, 195
species 193, 195
REPRODUCTIVE BIOLOGY Of Nephelium
lappaceum:
anther development 184
embryo development /86, 189
endosperm development 187
floral morphology, /82, 183
fruit:
formation of flesh /86, 189, 191
production 190
heterostyly 190
megagametophyte 184
megasporogenesis /84
microsporangium /82
microgametophyte /82
microsporogenesis /82
micropyle formation 185
ovule characters 191
ovule development 185, 186
phenology 183
pollen germination 185
pollen grains, degeneration 191
poilination (self) not possible 190
pollination and fertilization 187
231
seed coat and seed flesh development /86, 189,
191
seed germintion 190
sexuality of 190
viable pollen 190
Sandoricum koetjape 62, 64
Schima brevifolia 45
excelsa 2, 12
superba 46
wallichii ssp.noronhae
var. noronhae 47
Scindapsus 199
Simplicia 213
laxa 213, 214
238
Spathiphyllum 199
Symingtonia populnea see MULTIPLE AXILLARY
BRANCHING
Syngonium 199
Tamarindus woodland 56
Thysanolaena latifolia 62
maxima 62
Toona sereni 62
Trichilia connaroides 62
Trichomanes:
minutum 111, 113
minutum-group 112
Gard. Bull. Sing 37(2)(1984)
Trichomanes Cont.
motleyi 111
proliferum 111, 7/2, 113
first record for Singapore 111
morphological notes 112
Turraea pubescens 49
virens 49
Typhonium 199
Vitis vinifera 54
Xanthosperma 199
Zantedeschia 199
Zederachia 54
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PARKS AND RECREATION DEPARTMENT
PUBLICATIONS FOR SALE
1. The Gardens’ Bulletin, Singapore (Series IV).
Vols. 13-36, 1949-1983.
Price.
13(1) new impression: $12 31(1): $10, 31(2): $12.50
17(3): $12.50 32: $15.50
18 & 19: $25 per vol. The Freshwater Swamp-forest of S. Johore
20(1): $8 and Singapore by E.J.H. Corner (Gard.
25(1): $9, 25(2): $12 Bull. Sing. Suppl. 1) $35
26(1): $18, 26(2): $18 33(1): $21.50, 33(2): $12.50, Index: $1.90
.27(1): $18, Z7(2): $18.50 34(1): $21.50, 34(2) with Index for 34: $11.50
28(1): $18, 28(2): $15 35(1): $17.50, 35(2) with Index for 35: $21.50
29: $30 36(1): $21.50, 36(2): $14.00
30: $48 37(1) with Index for 36: $18.50
37(2) with Index for 37: $16.50
2. Materials for a Flora of the Malay Peninsula, Monocotyledons.
Parts 1, 2 and 3 remain available.
Price: $10 per set, $5 per part.
3. Selected Plants & Planting for a Garden City - Forty Shrubs, $1.20.
4. Selected Plants & Planting for a Garden City - Forty Climbers, $3.00.
5. A Guide to Tree Planting, $4.00.
6. A Pictorial Guide to the Singapore Botanic Gardens, $1.20.
7. Malayan Orchid Hybrids by M.R. Henderson and G.H. Addison, $15 (1969).
8. A Revised Flora of Malaya.
(a) Volk. 1, Orchids, by R.E. Holttum, $50 (3rd ed. 1980 Impr.).
(b) Vol. 2, Ferns, by R.E. Holttum, $20 (2nd ed. 1968).
(c) Vol. 3, Grasses, by H.B. Gilliland, $30 (1971).
9. Boletus in Malaysia by E.J.H. Corner, $50 (1972).
Items 1-6 can be purchased from the Commissioner, Parks & Recreation Department,
Botanic Gardens, Cluny Road, Singapore 1025; tel. nos. 4741165, 4741134.
For overseas orders, payment should be by bank draft or International Money Order and
made payable to the Commissioner of Parks & Recreation, Singapore.
Items 7-9 can be purchased from Singapore National Printers (Pte) Ltd, Upper Serangoon
Road, Singapore 1334, tel. no. 2820611 and their Sales Division, #01-29 International Plaza,
10 Anson Road, Singapore 0207, tel. no. 2230834.
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