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LIBRARY
OF THE
ARNOLD ARBORETUM
HARVARD UNIVERSITY
ISSN 0374-7859
THE GARDENS’ BULLETIN
SINGAPORE
Volume 40
(1987)
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
Cluny Road, Singapore 1025
CONTENTS
Volume 40
Pages
PART 1 — Ist June 1987
KATO, MASAHIRO:
A Phylogenetic Classification of Ophioglossaceae ................ ccc eceeeeeeeeeeeees 1-14
WONG, K.M.:
The Herbarium and Arboretum of the Research Institute of Malaysia
ee A EVISEQTICAL PEPSPECLIVE | 220.5252 i2- 0-25. 22s sar eaessttescnecscecsscdacces 15-30
WHITMORE, T.C., K. SIDIYASA, T.J. WHITMORE:
Tree Species Enumeration of 0.5 hectare on Halmahera .....................00000. 31-34
WONG, YEW KWAN:
Additions to ‘The Use of Tifgreen and Tifdwarf Bermuda Grasses in
eC AENE COE MOOIIESCS ce oes vic... coca ecee 82 Cee awics Bo Wins shan saccencees 35-36
CLASSEN, REGINE:
Anatomical Adaptations for Bird Pollination in Nicolaia elatior (Jack)
eee PAINE CSECN oie ce bce ged ee ns Seam eee Bee aAs es Suk wwaiveeis nnsee 37-43
WONG, YEW KWAN:
Ecology of the Trees of Bukit Timah Nature Reserve .....................c00eeeeee 45-76
PART 2 — Ist December 1987
BIDIN, Aziz:
A Preliminary Survey of the Fern Flora of Langkawi Islands ................... 77-102
LEONG, CHEE CHIEW:
MSMA for Controlling Cyperus kyllingia, Axonopus compressus and
Brachiara distachya in Tifgreen Bermuda Grass Turf ................ccceeeeeeeeeees 103-112
KENG, HSUAN:
Annotated List of Seed Plants in Singapore (XI) .............c cc cceceeeeeeeeeeeeeees 113-132
FOONG, THAI Wu and YANG, CHENG NOI:
Resolving Iron Deficiency in Wrightia religiosa by Foliar Analysis and its
Amelioration Using an Iron Chelate as a Soil Additive ..................ccccceeees 133-137
LEONG, CHEE CHIEW:
Chemical Growth Retardation of Baphia nitida with PP333 .................008. 139-144
VERHEIJ, E.W.M.:
Growth and Yield of Mango cv Golek in Java Over 25 Years ................00. 145-152
et ae a Ee Oe ec ec iba daca nebo accewedecedace 153-157
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INDEX
Volume 40
Basionyms and synonyms appear in italics. Page numbers in italics indicate the presence of illustrations.
Abdul Rahman bin Mohamad Ali, 27 Aspidium variolosum, 95
Acrostichum, 80 Asplenium, 81
aureum, 80, 93 macrophyllum, 94
speciosum, 80, 93 nidus, 80, 82, 93, 94
Adiantum, 101 paradoxum, 94
capillus-veneris, 82, 98 pellucidum, 94
caudatum, 81, 97 phyllitidis subsp. malesicum, 93
latifolium, 80, 98, 100 phyllitidis, 82, 93
mathewsianum, 82, 98 salignum, 93
peruvianum, 82, 98 spathulinum, 93
philippense, 81, 98, 101 spp., 80
polyphyllum, 98 tenerum, 94
stenochlamys, 81, 82, 97 Axonopus compressus, 35, 104, 107, 109,
tenerum 82, 98 LIC, Tit
trapeziforme, 82, 98 Aziz bin Budin, 24, 25
Adinandra, 57 Azolla pinnata, 80, 99
dumosa, 53, 55, 57
Aechmea bracteata, 126 Bain, V.L., 24, 25
Agave americana, 117 Baphia nitida, 139, 140, 142, 143
germinoflora, 117 Baccaurea, 55
sisalana, 117 kunstleri, 55
Allium cepa, 120 Barnard, R.C., 27
fistulosum, 119 Bayas, 131
ramosum, 120 Begonia, 139
sativum, 120 Belamcanda, 123
tuberosum, 120 Belvisia mucronata, 86
Aloé barbadensis, 116 Blechnum orientale, 80, 94
vera, 116 Blumeodendron tokbrai, 55
Amischotolyte, 124 Blyxa alternifolia, 114
Amphineuron, 81 auberti, 114
immersum, 81, 89 malayana, 114
opulentum, 89 Bolbitis, 81
terminans, 89 appendiculata, 95
Ananas comosus, 126 heteroclita, 94
Aneilema nudiflorum, 125 malaccensis, 81, 95, 101
vaginatum, 125 virens, 95
Angiopteris evecta, 82 Botrychium, 1, 3, 4 5, 6, 7, 8, 9, 10, 11, 12
Anisoptera curtisii, 47, 55, 61 lanceolatum, 6
Anthreptes, 41 lunaria, 4
malacensis, 38, 40, 41 paradoxum, 8
Antrophyum callifolium, 98 sens. lat., 3
Aporusa, 55 sens. str., 3
bracteosa, 57 sensu lato, 3, 10, 111
Archontopheonix alexandrae, 127 sensu stricto, 3
Arcypteris irregularis, 96 Botrypus, 1, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12
Areca catecu, 127 cicutaria, 9
triandra, 127 strictus, 8, 9
Arenga pinnata, 127 virginianus, 9
saccharifera, 127 Brachiara distachya, 35, 104, 110, 111
Artocarpus lanceifolius, 47, 55, 57, 59, 61 Bukit Timah Nature Reserve, Ecology of the
rigidus, 59 Trees, 45
Asam payah, 132 Burmannia championii, 123
Asparagus officinalis, 116 coelestis, 123
plumosus, 116 tuberosa, 123
sprengeri, 116 Burn-Murdoch, Alfred M., 15, 16
153
154
Calamus aquatilis, 127
densiflorus, 127
diepenhorstii, 127
erinaceous, 127
insignis, 127
javensis, 127
lobbianus, 127
laevigatus, 127
laxiflorus, 128
lucidus, 128
mitis, 128
ornatus, 128
oxleyanus, 128
pallidulus, 127
ridleyanus, 128
Calophyllum, 126
Campnosperma, 56, 57, 59, 61
auriculatus, 55, 57, 59
Ceratopteris thalictroides, 80, 96
Cheilanthes tenuifolia, 81, 97
Cheiroglossa, 1, 11
Chlorophytum comosum, 116
Christella, 81
dentata, 89
papilio, 81, 89
parasitica, 89
subpubescens, 81, 89
Chrysalidocarpus lutescens, 128
Chrysanthemum, 139, 143
Cocos nucifera, 128
Coleus, 139
Colysis acuminata var. angustata, 87
pedunculata, 87
Commelina attenuata, 124
benghalensis, 124
diffusa, 124
Cordyline fruticosa, 117
stricta, 117
terminalis, 117
Crinum asiaticum, 120
giganteum, 120
Crypsinus trilobus, 88
Ctenitis mannii, 96
subobscura, 96
Cubitt, G.E.S., 16
Curculigo /atifolia, 121
orchioides, 120, 121
recurvata, 121
villosa, 121
Cyanotis barbata, 124
cristata, 124
Cyathea borneensis, 85
contaminans, 85
glabra, 85
Cyclopeltis crenata, 96
Cyclosorus interruptus, 88
Cymodocea rotundata, 115
isoetifolia, 115
serrulata, 115
Cyperus kyllingia, 35, 104, 106, 107, 110, 111
radians, 35, 111
Cyrtostachys lakka, 128
renda, 128
Gard. Bull. Sing. 40 (1987)
Daemonorops angustifolia, 128
didymophylla, 128
grandis, 128
hystrix, 129
leptopus, 129
longipes, 129
micracantha, 129
periacantha, 129
Danaidae, 42
Davallia denticulata, 80, 81, 91
divaricata, 91
trichomanoides var lorrainii, 91
trichomanoides, 91
Dianella ensifolia, 116
Desch, H.E., 21
Dicamba (Fez PE 400), 104, 107, 110
Dicranopteris linearis, 80
linearis var. linearis, 84
speciosa, 84
Digitaria didactyla, 35
Dioscorea alata, 121
bulbifera, 121
esculenta, 121
glabra, 121
hispida, 121
laurifolia, 122
polyclades, 122
prainiana, 122
pyrifolia, 122
sansibarensis, 122
stenomeriflora, 122
Diplanthera uninervis, 115
Diplazium bantamense, 96
esculentum, 96
malaccense, 96
Dipterocarpaceae, 22
Dipterocarpus
caudatus ssp. penangianus, 55, 56, 57, 59, 61
penangianus, 55, 56
Doryopteris ludens, 80, 97
Dracaena aurantiaca, 117
brachystachys, 117
elliptica, 117
fragrans, 117
godseffiana, 118
granulata, 118
maingayi, 118
porteri, 118
sanderiana, 118
singapurensis, 118
terniflora, 118
umbratica, 118
Drymoglossum piloselloides, 80, 86
Drynaria, 88
bonii, 81, 87
rigidula, 80, 87
sparsisora, 80, 81, 87
Dyera costulata, 62
Egenolfia appendiculata, 95
Eichhornia crassipes, 122
Elaeis guineensis, 129
Elaphoglossum callifolium, 94
Index to Volume 40
Eleiodoxa conferta, 132
Elettaria speciosa, 37
Eleusine indica, 103
Endospermum diadenum, 55
malaccense, 55
Enhalus acrocoides, 114
koenigii, 114
Enhydrias angustipetala, 114
Eragrostis, 35
Eriocaulon longifolium, 126
sexanulare, 126
truncatum, 126
Etlingera elatior, 37
Eucharis grandiflora, 120
Euphorbia thymifolia, 104, 107, 108, 110, 111
Eurycles amboinensis, 120
sylvestris, 120
Fern Flora, Langkawi Islands, 77
Flagellaria indica, 125
Floscopa scandens, 124
Foresters’ Manual of Dipterocarps, 21
Forest Research Institute, Kepong, Malaysia. 15
Forrestia gracilis, 124
Forrestia marginata, 124
Foxworthy, 15, 17, 18, 19
Freycinetia, 31
Gloriosa superba, 116
Gluta wallichii, 55, 56, 57, 59
Golf Course Grasses, 35
Goniophlebium persicifolium, 80, 88
Gymnosiphon aphyllus, 123
Haemanthus multiflorus, 120
Halmahera, Tree Species Enumeration, 31
Halodule tridentata, 115
Halophila minor, 114
ovalis, 114
spinulosa, 114
Hanguana malayana, 125
Helminthostachys, 1, 3, 4, 5, 6, 7, 8, 9, 11, 12
zeylanica, 80, 82
Hemionitis arifolia, 97
Heterosmilax indica, 118
Hevea brasiliensis, 80, 85, 86, 87
Hibiscus, 139
Hippeastrum equestre, 120
puniceum, 120
Humata pectinata, 81, 91
repens, 91
vestita, 91
Hydrilla verticillata, 114
Hydrocarpus, 87
Hymenocallis littoralis, 120
Hymenophyllum, 81
denticulatum, 84
Hyophorbe verschaffeltii, 129
Iguanura ferruginea, 129
geonomaeformis, 129
Imperata cylindrica, 20
155
Japanobotrychium, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12
Kabong, 127
Kelapa bali, 129
Kelapa, 128
Kerintin, 132
Knema laurina, 93
Kochummen, K.M., 27
Koompassia malaccensis, 55
Korthalsia, 129
echinometra, 129
grandis, 129
polystachya, 129
rigida, 129
scaphigera, 129
wallichiaefolia, 129
Lecanopteris sinuosa, 88
Licuala ferruginea, 130
grandis, 130
spinosa, 130
triphylla, 130
Limnocharis flava, 113
Lindsaea, 81, 82
ensifolia, 90
heterophylla, 90
lucida, 90
parasitica, 91
repens var. pectinata, 90
repens, 90
Liriope muscari, 116
spicata, 116
Livistona chinensis, 130
kingiana, 130
rotundifolia, 130
saribus, 130
Lomariopsis lineata, 94
Loxogramme avenia, 86
Lycaenidae, 42
Lygodium, 82
circinnatum, 84
flexuosum, 83
linearis, 80
microphyllum, 80, 83
polystachyum, 83
salicifolium, 83
Mangifera, 55
lagenifera, 55
Mango cv Golek, 146, 147, 148, 150, 151
Growth and Yield, 145
Marsilea crenata, 80, 99
Mascarena verschaffeltii, 129
Melanorrhoea woodsiana, 55, 57, 59
Menon, P.K. Balan, 25
Mesophlebion chlamydophorum, 88
Metribuzin (Sencor WP 70), 107, 104, 110
Metroxylon sagu, 130
Microlepia speluncae var. hancei, 90
speluncae var. villosissima, 90
speluncae, 80
strigosa, 81, 90
156
Microsorium heterocarpum, 87
punctatum, 82, 86
Mitchell, B.A., 27
Minor Forest Products, Malay Peninsula, 18
Mohamed Nur bin Mohamed Ghous, 19
Monochoria hastata, 122
vaginalis, 122
Moraea, 123
MSMA for Controlling Weeds, 103
Murdannia nudiflora, 125
vaginata, 125
Myrialepis scortechinii, 130
Najas graminea var. angustifolia, 115
graminea, 115
Kingii, 115
Nenga, 131
pumila, 130
wendlandiana, 130
Nephrolepis biserrata, 80, 92
biserrata var. exaltata, 82, 92
hirsutula, 92
Ng, Francis S.P., 27
Nibong, 131
Nicolaia elatior, 37, 38, 39, 40, 41, 42
Nypa fruticans, 131
Oleandra pistillaris, 92
Oncosperma filamentosum, 131
horridum, 19, 131
tigillarium, 19, 131
Ophioderma, 1, 11
Ophioglossaceae, 1
Ophioglossum, 1, 3, 4, 5, 6, 7, 8, 9, 11, 12
bergianum, 7
crotalophoroides, 3
kawamurae, 8
lineare, 8
palmatum, 6, 7
pendulum, 3, 6, 7
petiolatum, 3
termale var. nipponicum, 4
Oreodoxa oleraea, 132
regia, 132
Osmundopteris, 1, 3
Palaguium, 59
wutia)-33,.55, 57, 59
hexandrum, 59
Pancratium littorale, 120
Pandanus, 40, 86
Paspalam conjugatum, 35
Peliosanthes teta ssp. humilis, 117
violacea, 117
viridis, 117
Pellacalyx saccardianus, 57
Pentacalyx, 98
Phaeomeria magnifica, 37
Phaseolus, 143
Philydrum lanuginosum, 123
Pholidocarpus kingiana, 130
Phymatodes nigrescens, 87
scolopendria, 88
Gard. Bull Sing. 40 (1987)
Pimelodendron griffithianum, 55, 57
Pinang, 127
raja, 128
Pinanga, 131
disticha Bl., 131
limosa Ridl., 131
malaiana (Mart.), 131
simplicifrons, 131
singaporensis, 131
subruminata, 131
Piper betel, 127
Pitcairnia integrifolia, 126
Pityrogramma calomelanos, 97
Platycerium, 80, 86, 101
coronarium, 80, 82, 85
holttumii, 80, 85, 101
platylobum, 80, 85
Plectocomia griffithii Becc., 131
Plectocomiopsis annulatus, 130
scortechinii, 130
Pleomele fragrans, 117
Polianthes tuberosa, 118
Pollia sorzogonesis, 125
Polystichum prolificans, 95
Polytrias amaura, 35
Pronephrium 81
asperum, 88
repandum, 89
Pseudodrynaria coronans, 101, 81, 87
Pteris, 81
biaurita, 92
ensiformis, 92
longipinnula, 93
mertensioides, 93
scabripes, 92
venulosa, 92
vittata, 92
Ptychoraphis singaporensis, 132
Ptychosperma macarthurii, 132
sanderianum, 132
Pyrrosia adnascens, 86
lanceolata, 86
longifolia, 86
penangiana, 81, 86
spp., 80
stigmosa, 86
varia, 86
Raphia farinifera, 132
ruffia,132
excelsa, 132
flabelliformis, 132
humilis, 132
Rhizoglossum, 1, 11
Rhodamnia, 57
cinerea, 57
trinervia, 57
Rhoeo spathacea, 125
Rotan bakau, 129
batu, 127
chichi, 127
chochor, 129
dahan, 132
Index to Volume 40
hudang, 128
kertong, 130
sabite, 129
sepah, 129
sepat, 128
simut, 129
Roystonea oleracea, 132
regia, 132
Sagittaria sagittifolia L., 113
Salacca, 132
affinis, 132
conferta, 132
edulis, 132
Salak hutan, 132
Sansevieria cylindrica, 118
trifasciata, 118
Sceptridium, 1, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12
daucifolium, 7
japonicum, 6
multifidum, 3
ternatum, 6
Schizaea dichotoma, 83
digitata, 80, 83
Sciaphila affinis, 115
maculata, 115
Seed Plants in Singapore (XI), Annotated List,
113
Setten, G.G.K., 27
Shorea curtisii, 47, 55, 57, 59, 60, 61
glauca, 55, 60
gratissima, 55
leprosula, 60
macroptera, 59
maxwelliana, 60
Sloetia, 59
elongata, 57, 59
Smilax bracteata var. barbata, 119
calophylla, 119
leucophylla, 119
megacarpa, 119
myosotiflora, 119
woodii, 119
Sphaerostephanos heterocarpus, 88
penniger, 88
Stenochlaena palustris, 80, 83
Streblus, 59
elongatus, 57, 59
Susum malayanum, 125
Swintonia, 55
schwenkii, 55
spicifera, 55
Symington, C.F., 21, 22
Tacca cristata, 121
integrifolia, 121
leontopetaloides, 121
157
Taenitis blechnoides, 97
Tapeinidium pinnatum, 91
Tectaria, 81, 82
angulata, 96
brachiata, 81, 95
maingayi, 95
oligophylla, 96
rumicifolia, 96
semipinnata, 95
variolosa, 81, 82, 95, 101
Thalassia hemprichii, 114
Thismia aseroe, 123
fumida, 123
Tillandsia usneoides, 126
Timonius wallichianus, 57
Tradescantia fluminensis, 125
Trema, 84
spp., 62
Trichomanes, 81
bipunctatum, 84
digitatum, 84
javanicum, 84
latemarginale, 84
maximum, 85
obscurum, 85
pallidum, 84
proliferum, 84
Trimezia martinicensis, 123
Tsuji, Yukio, 24
Tukas, 128
Vincent, A.J., 27
Vittaria, 88
angustifolia, 99
ensiformis var. latifolia, 99
ensiformis, 80, 98
Watson, J.G., 19, 20, 21, 25
Whitmore, T.C., 27
Wijasuriya, J.S., 25
Wrightia, 133, 137
religiosa, 133, 136, 137
Wyatt-Smith, John, 25, 26, 27
Xyris anceps, 126
complanata, 126
pauciflora, 126
Yucca aloifolia, 118
gloriosa, 118
Zebrina pendula, 125
Zephyranthes candida, 120
carinata, 120
flava, 120
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THE GARDENS’ BULLETIN
VOL. 40 (Part 1) 1 June 1987 {
es er C‘(C
x
: CONTENTS
PAGES
es
_) KATO, Masauiro:
A Phylogenetic Classification of Ophioglossaceae ............. 0... cece eceeceeeeeeeeeneeeenenenes 1-14
WONG, K.M.:
} The Herbarium and Arboretum of the Research Institute of Malaysia at Kepong —a
CN ee EC 8 cS acdc cae te 2 os dees MES Selim oe See Maa dda cmepansebons 15-30
WHITMORE, T.C., K. SIDIYASA, T.J. WHITMORE:
\ Tree Species Enumeration of 0.5 Hectare on Halmahera ............... 0. ccceeecceceeeeeeeee ees 31-34
:
' WONG, YEw Kwan:
\ Additions to “The Use of Tifgreen and Tifdwarf Bermuda Grasses in Two Singapore
SSE TS Ee a AUD ec ese eee ec Renee 35-36
\
3 CLASSEN, REGINE:
\ Anatomical Adaptations for Bird Pollination in Nicolaia elatior (Jack) Horan
s BE a SE tr eared ne thnss Cee Se ear ne en. ee 37-43
8 WONG, Yew Kwan:
@ Ecology of the Trees of Bukit Timah Nature Reserve ..................ccececeeeecececeeeeeeeeees 45-76
N
. r)
Published by the Botanic Gardens
‘ Parks and Recreation Department
Ministry of National Development
Cluny Road, Singapore 1025.
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es -28
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THE GARDENS’ BULLETIN
SINGAPORE
VOL. 40 (Part 1)
CONTENTS
KATO, MASAHIRO:
A Phylogenetic Classification of Ophioglossaceae ....................0c0eeeeeee: gs lath Pate ous
WONG, K.M.::
The Herbarium and Arboretum of the Research Institute of Malaysia at Kepong — a
Sg RS OP ean Re Cry cf 2 Pe ae oe ee
WHITMORE, T.C., K. SIDIYASA, T.J. WHITMORE:
Tree Species Enumeration of 0.5 Hectare on Halmahera ..................00..0060. A Lake
WONG, Yew Kwan:
Additions to ‘The Use of Tifgreen and Tifdwarf Bermuda Grasses in Two Singapore
Golf Courses’
CLASSEN, REGINE:
Anatomical Adaptations for Bird Pollination in Nicolaia elatior (Jack) Horan
(Zingiberaceae)
WONG, Yew Kwan:
Ecology of the Trees of Bukit Timah Nature Reserve
Published by the Botanic Gardens
Parks and Recreation Department
Ministry of National Development
Cluny Road, Singapore 1025.
ee ns ae eee Rien, «ee aera aes wee eS a8 Seni hp oeials s a'd.d bein, Sie Vpiew ais els = a'elew «le wees «ce ton wade coacedewe
ee ee ee Oe See See SS Ss a ea aap pe a6 se dt 80 619 6o.o ea .e 6a €n vine vig oe. eclems = pe cee ecucasscetacwoecuevecsesc
ee er es
1 June 1987
PAGES
15-30
31-34
35-36
37-43
45-76
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A Phylogenetic Classification of Ophioglossaceae
MASAHIRO KATO
Botanical Gardens, Nikko, Faculty of Science, University of Tokyo, Hanaishicho 1842, Nikko 321-14
Japan
EFFECTIVE PUBLICATION DATE: 8 SEPTEMBER 1987
Abstract
A cladistic approach was undertaken to construct a phylogenetic classification of the Ophioglos-
saceae. The characters used are: rhizome, stele, leaf sheath, size and cutting of trophophyll, vascular
branching in leaf axes, venation, position of sporophyll on trophophyll, form and construction of
sporophyll, position and dehiscence of sporangia, and suspenscr. Character state, primitive (plesiomor-
phic) vs. advanced (apomorphic), was estimated for each character. Based on presumed character
phylogenies, a cladogram representing phylogenetic relationships was drawn. Although in previous
classifications (e.g. Clausen, 1938) the Ophioglossaceae were classified into three genera, Botrychium,
Helminthostachys and Ophioglossum, it is contended here that the genus Botrychium sensu Clausen is
not a monophyletic group. In this paper I propose a new classification recognizing six genera and
arrange them as in Cracraft’s (1974) sequence as follows: 1 (most primitive), Botrypus; 2, Japano-
botrychium; 3-1, Sceptridium; 3-2, Helminthostachys; 4, Botrychium; 5 (most advanced), Ophio-
glossum.
The Ophioglossaceae have a combination of unique or rare morphological char-
acters among the extant lower vascular plants: three-dimensional leaf architecture,
collateral vascular bundles (sympodial vascular system; Stevenson, 1980) with
secondary growth in many species, hair-lacking roots with endophytic fungi, non-
circinate vernation of leaves, a soft and fleshy plant body devoid of sclerenchyma,
massive eusporangia, and subterranean, mycorrhizic, massive gametophytes of
fundamentally axial organization (Bierhorst, 1971; Foster and Gifford, 1974).
Because of those characters the family is regarded as an isolated group composing
the monotypic order Ophioglossales.
The usual systematic treatment is that the Ophioglossales, along with the Marat-
tiales, constitute a eusporangiate group in Filicopsida or ferns (e.g. Christensen,
1938; Eames, 1936; Smith, 1955), or the order is one of three or more orders
composing Filicopsida (e.g. Copeland, 1947; Foster and Gifford, 1974; Pichi Ser-
molli, 1973: Tryon and Tryon, 1982). However, a real relationship of the Ophio-
glossales with the Marattiales and also Filicales is doubted by some authors
(Bierhorst, 1971; Kato, 1982, 1983). To determine the phylogenetic position of the
Ophioglossaceae in the vascular plants, it is needed to better understand the
phylogenetic relationships within the family.
The currently accepted classification of the Ophioglossaceae (e.g. Copeland,
1947; Tryon and Tryon, 1982) is mainly that of Clausen (1938) who recognized
the genera Botrychium, Helminthostachys and Ophioglossum in the family. He
classified Botrychium in three subgenera (Osmundopteris, Sceptridium and
Botrychium) with seven sections, and Ophioglossum into the four subgenera
Ophioglossum, Rhizoglossum, Cheiroglossa and Ophioderma. In his revised sys-
tem, Nishida (1952) divided the Ophioglossales into two suborders, Ophioglos-
sineae and Botrychiineae. The Ophioglossineae consisted of four genera, Ophio-
glossum, Ophioderma, Cheiroglossa and Rhizoglossum, belonging to the one family
Ophioglossaceae, while the Botrychiineae comprised two families, Botrychiaceae
with the genera Botrychium, Sceptridium and Osmundopteris (= Botrypus), and
Helminthostachyaceae with the sole genus Helminthostachys. Following Clausen
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Classification of Ophioglossaceae S)
(1938) in general scheme, Kato and Sahashi (1977) recognized a fourth subgenus
Japanobotrychium in genus Botrychium, which had been treated as a section of
subgenus Osmundopteris by Clausen (1938).
The classification of Ophioglossaceae is still controversial, however. Many au-
thors consider Sceptridium the most primitive, and Botrypus (= Osmundopteris)
and Botrychium sensu stricto advanced, mainly from morphological characters
(Bower, 1926; Clausen, 1938, 1954; Nishida, 1957; Nozu, 1955). Based on the
vascular anatomy of leaves, Chrysler (1945) and Nishida (1952) presumed
Botrychium sens. str. to be the most primitive, and Botrypus (as Osmundopteris)
and Sceptridium advanced. Another different opinion on the relationship was
presented by Kato (1978a) utilizing sporangia as well as such foliar characters as
basal sheath, and size and cutting of leaves. Botrypus (as subgenus Osmundopteris)
was regarded as the most ancestral while Japanobotrychium, Sceptridium and
Botrychium sens. str. as successively specialized.
The systematic position of Helminthostachys is also debatable. It is generally
placed as an intermediate between Botrychium sensu lato and Ophioglossum
(Clausen, 1938; Nishida, 1957). Nishida (1952) recognized a monotypic Helmin-
thostachyaceae and a family Botrychiaceae in Botrychiineae. Kato (1982) sug-
gested a closer affinity of Helminthostachys to Japanobotrychium and Sceptridium
among the then recognized subgenera of Botrychium.
Since previous schemes of classification of Ophioglossaceae differ from each
other, and none is widely accepted, the family needs re-examination from the
viewpoint of phylogenetic classification. Because no closely related fossils have
been found, such a systematic study must be based on comparative analysis of living
genera and general paleobotanical information.
The best method of constructing a phylogenetic classification seems to me to be
cladistics (e.g. Bremer and Wanntorp, 1978; Cracraft, 1974; Estabrook, 1978;
Hennig, 1965, 1966; Nelson, 1974; Wagner, 1969; Wiley, 1980). In cladistics a
phylogenetic classification is constructed by recognizing levels of monophyletic
groups, which have synapomorphic characters derived from those of the nearest
common ancestors. Monophyletic groups should be distinguished from paraphyle-
tic (symplesiomorphic) and polyphyletic groups. Cladistics has since Hennig elabo-
rated several different methods, each suitable for certain groups or subjects (Dun-
can and Stuessy, 1985; Funk and Stuessy, 1978). By adopting the basic method of
cladistics (Hennig, 1965, 1966; Estabrook, 1978), an attempt is here made to
examine the genealogical relationship in the Ophioglossaceae and arrange them in
a phylogenetic classification. Basic monophyletic groups treated here are Bot-
rychium sens. str., Botrypus (= Osmundopteris), Helminthostachys, Japanobot-
rychium, Ophioglossum and Sceptridium, although most of the genera need more
detailed revisionary work.
Phylogenetic Estimation of Characters
Almost all data on the characters analyzed here are cited from previous works.
The basic source for this study is Kato and Sahashi (1977) and Kato (1978a, 1978b,
1982, 1983). The characters of each genus are presented in Table 1, and character
State, primitive or plesiomorphic, vs. advanced or apomorphic, is estimated in each
character.
Rhizome and stele. — The rhizome is radial in construction in Botrychium,
Botrypus, Japanobotrychium, Ophioglossum and Sceptridium. It has been reported
that Sceptridium (as Botrychium sens. lat.) multifidum has a spiral phyllotaxy of
(1/3)2/5(3/8) (Stevenson, 1980), Ophioglossum petiolatum 2/5, O. crotalophoroides
1/2 (Webb, 1975, 1981), and O. pendulum 1/3 and 3/5 (Petry, 1914). The creeping
f
“
- CAC tie — Pi. . 2 ee
Botrychium lunaria. P|. 1: Transection of rhizome. C, cambium; E, endodermis; PH,
primary phloem; PX, primary xylem; SX, secondary xylem. PI. 2: Longisection of shoot,
showing pore closing at leaf sheath. A, shoot apex; FP with arrowheads, closed pore; P,
pore; SH1/, leaf sheath of first leaf primordium; SH2, leaf sheath of second leaf primor-
dium. Scales 100 pw.
rhizome of Helminthostachys 1s dorsiventral with two rows of leaves on the dorsal
side (Farmer and Freeman, 1899). As is generally the case in vascular plants, the
radial construction of rhizome in most genera of Ophioglossaceae is primitive while
the dorsiventral rhizome of Helminthostachys is advanced.
The rhizomes of Botrypus, Japanobotrychium and Sceptridium have a cambium
producing secondary vascular tissue. Although Botrychium lunaria has been de-
scribed as having no vascular cambium (Nozu, 1956) or as having a cambium
derived from the pericycle (Lang, 1913), it has the same organization as the others,
excepting lower cambial activity (Takahashi, pers. commun.; Plate 1). There is no
cambium in Helminthostachys and Ophioglossum, and Farmer and Freeman (1899)
described the xylem as differentiating very slowly from the procambial strand in
Helminthostachys. However, addition of xylem occurs both in Helminthostachys
(Farmer and Freeman, 1899; Takahashi, pers. commun.) and Ophioglossum
(Boodle, 1899). It is suggested that cambial activity has weakened and eventually
disappeared, as in the evolutionary trend from a woody to herbaceous habit in
Isoetales and related fossils (Rothwell and Ervin, 1985) and the flowering plants
(Eames, 1961). The presence of a vascular cambium may be primitive and its
absence advanced. Helminthostachys has a dorsiventral solenostele and Ophioglos-
sum has a radial dictyostele, and this suggests a different or phylogenetically
remote origin. It is interesting that Ophioglossaceae are similar to Isoetales (Lyco-
podiopsida) in possessing secondary tissue in very small (probably reduced) rhi-
zomes.
Leaf sheath. — The stem apex and leaf primordia are covered by leaf sheaths to a
varying extent depending on the genus (Clausen, 1938; Kato 1978a). The covering
is incomplete in Botrypus in which sheath margins overlap along the entire length
except at the base. In Botrychium, Helminthostachys, Japanobotrychium and Scep-
tridium, leaf sheaths cover more completely, and have a transversely elongate,
small pore on the side opposite the leaf base, through which the inner cavity
surrounding younger leaf primordia opens to the exterior. A further complete
covering is seen in Botrychium. In B. lunaria the pore that exists in young leaf
primordia disappears during leaf development (Plate 2; Kato, 1982). In Ophioglos-
sum with a sunken stem apex, the pore is a narrow channel opening more or less
obliquely (Holle, 1875; McAlpin, 1971). Closing of the pore takes place also in O.
termale var. nipponicum (Imaichi, pers. commun. ). In the light of the morphologic-
al basis that more appendicular structures are generalized, it is suggested that the
‘a
a
:
Pn
s
D
y+
= .
a
Plate 3. Silhouettes of Ophioglossaceae leaves. A, Botrypus strictus (Ohba 411, Honshu, Japan; TI);
B, Japanobotrychium lanuginosum (Hara et al. 6305423, E. Nepal; TI); C, Sceptridium
daucifolium (Hara et al. 2458, Darjeeling, Sikkim; TI); D, Helminthostachys zeylanica (Taga-
wa & Iwatsuki 2170, Ryukyus, Japan; TI); E, Botrychium lunaria (Makino s.n. 25 Jul. 1881,
Mt. Fuji, Japan; TI); F, Ophioglossum petiolatum (Yoshinaga s.n. 19 Jun. 1938, Shikoku,
Japan; TI); S, sporophyll.
incomplete covering of the leaf sheath in Botrypus is primitive and that of the other
genera advanced. Among the latter, Japanobotrychium, Sceptridium and Helmin-
thostachys are relatively primitive in this character, as compared with Botrychium,
which has a further advanced leaf sheath.
Trophophyll morphology. — The trophophyll is variable in size and cutting in diffe-
rent genera, and variation in the two characters is more or less correlated (Plate 3;
Clausen, 1938). Botrypus and Japanobotrychium have the largest and most
6 Gard. Bull. Sing. 40(1) (1987)
dissected trophophylls in the Ophioglossaceae, and Sceptridium has smaller and
less dissected trophophylls. The trophophyll of Helminthostachys is of almost the
same size as that of Sceptridium but is much less dissected. The trophophylls of
most Botrychium and Ophioglossum are the smallest, although in Ophioglossum
pendulum they are exceptionally large (to 1 m long). They are simply pinnate in
Botrychium while in Ophioglossum they are in general simple and entire, but often
forked in O. pendulum and lobed in O. palmatum.
Except for Ophioglossum pendulum which might show some secondary leaf
enlargement, Ophioglossaceae exhibit a continuous variation in leaf size and cut-
ting between the larger and highly dissected trophophylls and the smaller and less
dissected. Reduction and simplification, which is a general tendency in the lower
vascular plants, is also conceivable in the Ophioglossaceae. This assumption 1s
supported by an ontogenetic study by Imaichi and Nishida (1986) who showed
apparently three-dimensional organization of trophophylls at early ontogenetic
stages and the persistence of a tetrahedral apical cell at the apex of the primary
through tertiary leaf axes in Sceptridium ternatum. It is also supported by an
out-group criterion that the larger and finely dissected trophophylls are shared by
Japanobotrychium and Botrypus which is an out-group for another group including
the former genus, as indicated by sporangial morphology and other characters (see
below). Therefore, Sceptridium, Helminthostachys, Botrychium and Ophioglossum
have advanced trophophylls, compared to primitive ones in Botrypus and Japano-
botrychium. Among the former, Botrychium has more advanced trophophylls and
Ophioglossum has the most specialized.
Helminthostachys has a characteristic trophophyll: it is imparipinnate with
oblong entire terminal and lateral pinnae, the basal and usually also the subbasal
pinnae with conforming basiscopic pinnules. The presence of such basiscopic
pinnules is shared with Japanobotrychium, Sceptridium and part of Botrychium (B.
lanceolatum). Moreover, Helminthostachys and Sceptridium exhibit a strong simi-
larity in leaf shape at the young ontogenetic stages: juvenile leaves of Helminthos-
tachys with three deltoid-ovate leaflets (Fig. 1) are quite similar to those of Scepztri-
dium japonicum (Nishida, 1955). It is presumed that the trophophyll of Helmin-
thostachys has been derived from, or from a common ancestral form of, that of
Sceptridium which at maturity is the most similar to Helminthostachys in leaf size
and dissection.
Vascular branching in leaf axes. — Bower (1923) recognized two types of
vascular branching in fern leaves; of those he regarded marginal branching as
Fig. 1. Leaves of very young (left) and young plants of Helminthostachys zeylanica (Kato et al. C-8043,
Ceram. Indonesia; TI).
Classification of Ophioglossaceae 7
primitive and extra-marginal as advanced. The same evolutionary trend was ap-
plied to the Ophioglossaceae by Chrysler (1945), Nozu (1955) and Nishida (1957).
Their arguments were based on an assumption that a simpler branching type is
primitive and the more elaborate, advanced, and the three-dimensional archaic
branching is maintained in a simpler form in vasculature in Ophioglossaceae leaves.
Kato (1978b) made a detailed re-examination of leaves of the family and criticized
the validity of such an assumption. He concluded that in the Ophioglossaceae
simplification from extra-marginal branching to marginal might have taken place
parallel to the leaf reduction and simplification discussed above. It is estimated that
in vascular branching, Botrychium and Ophioglossum are derived and other genera
primitive.
It is of little doubt that marginal and hetero-marginal branchings of the sporo-
phyll vascular bundles are each related to extra-marginal, and the first two have no
direct relationship to one another (Kato, 1978b). Based on the evolutionary trend
in vasculature noted above and also the phylogenetic relationship discussed below
that Botrypus and a group of the other genera are sister-groups, the hetero-
marginal branching of Helminthostachys and the marginal one of Botrychium have
been derived in parallel from extra-marginal by specialization, respectively.
Venation. — Veins are free in Botrychium, Botrypus, Helminthostachys, Japano-
botrychium and Sceptridium, and anastomosing in Ophioglossum. Elaboration
from free to anastomosing venation is a general evolutionary trend in megaphyllous
leaves, although the reverse tendency seems to have taken place in occasional
cases, e.g. Polypodiaceae, in relation to reduction in leaf size. In Ophioglossaceae,
as in general in megaphyllous plants, the anastomosing venation in Ophioglossum
with specialized simple leaves is believed advanced, and free venation in the other
genera as primitive.
Among the free-veined genera, Botrypus, Japanobotrychium and Sceptridium
have pinnate and forked venation in leaf segments, while veins in Botrychium are
parallel, and lateral veins are parallel and occasionally anastomosing in Helmin-
thostachys (Bhambie and Madan, 1982). Venation of the first three genera is
primitive compared with that of the last two, being parallel to the above suggested
evolutionary trend in trophophyll morphology.
Position of sporophyll on trophophyll. — The sporophyll is attached at or near
the top of a stipe of the trophophyll in most genera, i.e. Botrychium, Botrypus,
Helminthostachys and most species of Ophioglossum. In O. pendulum a sporophyll
occurs on the lower part of the blade which is attenuated toward the short stipe,
and in O. palmatum several sporophylls are attached near the blade margin.
Ophioglossum bergianum also is aberrant in having a sporophyll at the stipe base.
In Japanobotrychium the sporophyll occurs on the lower part of the trophophyll
rachis. In most species of Sceptridium the sporophyll is attached on the lower part
of the trophophyll stipe, and in S. daucifolium it is at the upper part of the stipe.
Thus, that sporophylls occur at or near the top of the trophophyll stipe is a general
and so probably primitive character state which exists in different genera with
various leaf morphology. Comparatively, Japanobotrychium, some species of
Ophioglossum and Sceptridium are apparently advanced. They are not considered
a synapomorphic group of common origin, because the position of sporophylls
differs in different genera, and each primitive stage is seen in most species in
Ophioglossum, and in S. daucifolium in Sceptridium. In Japanobotrychium the
vascular supply to the sporophyll actually branches below the branching point of
vascular supplies to the trophophyll pinnae, thus indicating a close anatomical
similarity to Botrypus with likewise extra-marginal vascular branching (Kato,
1978b).
Form and construction of sporophyll. — The Ophioglossaceae exhibit strong leaf
dimorphism, sporophylls generally consisting of lamina-lacking leaf axes and spor-
8 Gard. Bull. Sing. 40(1) (1987)
angia, although abnormally, a lamina develops on sporophylls to some extent, or
some sporangia are produced on trophophylls. The most extraordinary are the
leaves with no trophophylls in Botrychium paradoxum (Wagner and Wagner
1981), Ophioglossum kawamurae (Tagawa, 1939), and O. lineare (Brause, 1912).
Sporophylls are foliar, variously pinnate and more or less similar to trophophylls in
Botrychium, Botrypus, Japanobotrychium and Sceptridium, while they are spikes in
Helminthostachys and Ophioglossum. The former is a primitive character and the
latter advanced. The last two are not considered synapomorphic, because they
differ considerably in form and construction. In Helminthostachys the sporophyll is
a spike consisting of a stipe and a rachis with sporangiophores along both sides. In
Ophioglossum it is a simple spike which is considered to consist of a stipe and a
narrow blade (not a rachis), because the sporophyll has an entire margin and
anastomosing veins as the trophophyll, but with areoles in few rows, necessitated
by its narrowness (Fig. 2).
The sporophylls are flattened, as are leaves in general, in all the genera of
Ophioglossaceae but Helminthostachys in which they are cylindrical: the sporan-
giophores are arranged in two or three irregular rows along each side of the rachis
(Kato, 1978a). In the family flattened leaves are primitive and cylindrical adv-
anced. The sporangiophores are modified branches of leaf axes and vascular
branching to the sporangiophores is extra-marginal, a character common to all
genera except for Botrychium and Ophioglossum (Kato, 1978a).
Position and dehiscence of sporangia. — Phylogenetic comparisons among spor-
angia of the vascular plants and those of Ophioglossaceae have been made by Kato
(1978a, 1982, 1983). This character is the most basic for reconstructing the phy-
logeny in this study, and the assumed evolutionary trend is in good accordance with
paleobotanical evidence and comparative morphology of the extant groups. There
are two types of sporangia, each in one of two phyletic lines in the early vascular
plants and their descendants. In the megaphyllous group derived from the Rhy-
niopsida and Trimerophytopsida, sporangia are terminal at the end of axes or veins
that have been derived from axes and dehisce longitudinally from the top to the
base. In the microphyllous group including the Zosterophyllopsida, sporangia are
attached laterally on axes or leaves (on the adaxial side) and dehisce distally along a
transverse line.
In Ophioglossaceae, belonging to the megaphyllous group, sporangia terminal at
the endings of leaf axes and with longitudinal dehiscence are primitive, while those
apparently lateral on the side of leaf axes (but terminal at vein-endings) and with
apparently distal dehiscence are advanced (Fig. 2). The former are present in
Botrypus (particularly B. strictus) and the latter in the other genera. The sporangia
of Ophioglossum, although quite the same in morphology as those of the latter
genera, are unique in being sunken and embedded in lamina tissue along the
margin of the spike which is a narrow blade, and not of an axial nature, as noted
above (Fig. 2). Therefore Ophioglossum is in a further advanced character state in
the position of sporangia. It is noteworthy that a similar evolutionary trend in
sporangium morphology with reference to dehiscence is seen in the leptosporangi-
ate ferns (e.g. Bower, 1935).
Suspensor. — It has been reported that the embryo of Helminthostachys and
Sceptridium has a suspensor (Bierhorst, 1971; Lang, 1914; Lyon, 1905; Nishida,
1955: Nishida and Imaichi, 1971) while no suspensor is known in the other
genera. A suspensor-bearing embryo is known in various vascular plant groups
such as Lycopodiales, Selaginellales, Marattiales, gymnosperms and angiosperms
which have a massive or compact gametophyte (Bower, 1908; Land, 1923; Lang,
1914). Such an embryo is generally regarded as primitive. Although there is no
decisive evidence for the evolutionary trend in the suspensor of Ophioglossaceae,
ap 2 ___SIMPLELEAF
~ANASTOMOSING VEIN
E | 3 |
Fig. 2. Fructifications of Ophioglossaceae. A, presumed ancestral form; B, Botrypus; C & D, Bot-
rychium, Japanobotrychium and Sceptridium; E, presumed ancestral form with distal sterile
segments, equivalent to B; F, Helminthostachys; G, distal part of trophophyll of Ophioglos-
sum; H, distal part of sporophyll of Ophioglossum (D, dehiscence line; S, sporangium; V,
vein); /, J, sporangia; J, Botrypus strictus; J, Botrychium, Japanobotrychium and Sceptridium.
(A-C, E, F, 1 & J after Kato (1978a)).
the cladogram presented below suggests the existence of a suspensor to be ad-
vanced, as far as information available at present is concerned. Another possibility
is that the presence of a suspensor is ancestral in the family and this has disappeared
in several genera, or that a suspensor-bearing embryo may be more widespread in
the family than we know.
Phylogenetic Relationship and Classification
The evolutionary trends of all the characters examined and a deduced phylogene-
tic relationship between genera of Ophioglossaceae are presented in Figure 3. All
character phylogenies are compatible if only we assume that the presence of a
Suspensor in the embryos of Helminthostachys and Sceptridium is an advanced
character state, based on its absence from Botrypus and Japanobotrychium both of
which are out-groups of Helminthostachys and Sceptridium, and on the embryos of
Botrychium and Ophioglossum lacking a suspensor. Botrychium and Ophioglos-
sum, and Helminthostachys and Sceptridium are each synapomorphic groups which
in turn constitute a higher level synapomorphic group with Japanobotrychium.
Among the genera, Botrypus is the most primitive and Ophioglossum is the most
advanced. Japanobotrychium, Sceptridium and Helminthostachys, and Botrychium
are successively more advanced genera between the two extremes.
Each genus possesses its own unique advanced character(s), showing that it is an
elaboration from its nearest ancestor. Exceptionally, Botrypus consistently shows
the primitive state of all characters examined. However, B. cicutaria and B.
virginianus are derived in their sporangial morphology and B. strictus in its spike-
like narrow sporophyll. It is, therefore, obvious that Borrypus is also an elaboration
>
|
gfemese sg
{ ———
ae
an
|
to)
Fig. 3. Cladogram showing character state and phylogenetic relationship in Ophioglossaceae. Num-
bers at left side are character codes shown in Table 1. White and black squares indicate
respectively primitive and advanced character states. BP, Botrypus; J, Japanobotrychium; S,
Sceptridium,; H, Helminthostachys; BC, Botrychium; O, Ophioglossum.
from the common ancestor, with its sister group being all the other extant genera of
Ophioglossaceae.
The phylogenetic scheme presented in Figure 3 remarkably contradicts rela-
tionships postulated in currently accepted classifications. In these (e.g. Clausen,
1938; Nishida, 1952), Botrychium, Botrypus, Japanobotrychium and Sceptridium
are united in a single group, i.e., genus Botrychium sensu lato or family
Botrychiaceae. In the relationship obtained in this study it is not a monophyletic
group and at most a paraphyletic or polyphyletic group.
For a comparison between the phylogenetic relationship of this study and that of
Clausen (1938) and Nishida (1952) which differs from Clausen principally in rank of
taxa, the divergence degree of the genera is estimated (Table 2). For each charac-
ter, the primitive state is coded as 0 and the derived state as 0.3, 0.5, 0.7 or 1
according to the degree of deviation from the primitive state. All of the values are
summed up as divergence degree. The phylogenetic relationship and divergence of
10
Classification of Ophioglossaceae 11
Table 2. Divergence of characters in Ophioglossaceae
Character code* Divergence
Genus degree
£2 3 4 5 6 7 8 9 et 12
Botrypus 00 0 0 0 0 0 0 0 O O40.” OG
Japanobotrychium 0 0 0.5 0 0 0 O05 PCL of! een Seay
Sceptridium eeo.s O55," 0.3 0 0: 4:0 ee O.8
Mepmamostachys ©1105 05° .:0.7)'-05 0.3 erin 050 OL 17D
Botrychium COO TOF LOY 03 0 GEN OS EA ISS
Ophioglossum eA ae ROA OLOY © ROY STD OLN, 706 OS 2i'8.5
* Following Table 1 (13 excluded).
time
divergence
Fig. 4. Phylogenetic relationship and divergence of characters in Ophioglossaceae. BP, Botrypus; J,
Japanobotrychium; S, Sceptridium; BC, Botrychium; H, Helminthostachys; O, Ophioglossum.
characters are shown in Figure 4. From this estimation of divergence, it is clear that
Helminthostachys and Ophioglossum are the most specialized genera while Bot-
rychium, Botrypus, Japanobotrychium and Sceptridium maintain relatively general-
ized characters. In the previous classifications, the former highly specialized groups
have been treated as independent genera, while the latter relatively primitive
genera have been classified into a single, artificial, non-monophyletic group (Bot-
rychium sensu lato or Botrychiaceae).
As regards the phylogenetic classification of Ophioglossaceae, I admit that the
pair of the most closely related genera, Botrychium and Ophioglossum, and Hel-
minthostachys and Sceptridium, are distinct enough to warrant generic status, and
therefore the others also deserve generic or higher rank. Although some authors
(e.g. Nishida, 1952) recognize as genera Cheiroglossa, Ophioderma and Rhizoglos-
sum as well as Ophioglossum, these share advanced characters as shown in Table 1
and Figure 3, so that they are best treated as infrageneric taxa. It is not feasible to
classify the family into subgroups in any way although the most meaningful two
12 Gard. Bull. Sing. 40(1) (1987)
groups would be Botrypus and then another group consisting of all the other
genera. In conclusion, I propose the following classification:
Family Ophioglossaceae
1. Genus Botrypus 3-2. Genus Helminthostachys
2. Genus Japanobotrychium 4. Genus Botrychium
3-1. Genus Sceptridium 5. Genus Ophioglossum
In this scheme the genera are arranged in sequence as to reflect the phylogeny
shown in Figures 3 and 4, using Cracraft’s (1974) sequencing classification method.
In this classification the following relationship is indicated: 1—2—3 (1—2)—4—5,
and genus 1 is the most primitive and 5 the most advanced.
Acknowledgements
I am thankful to Dr H. Ohba, University of Tokyo, for discussion and literature
on cladistics, and Dr H. Keng, National University of Singapore, for reading the
manuscript. I am also indebted to Mr A. Takahashi, Osaka University, for col-
laborative work and Plate 1. Part of this study was done at Department of Botany,
National University of Singapore, supported by JSPS (Japan Society for the Prom-
otion of Science) grant for JSPS-NUS (National University of Singapore) Scientific
Cooperation Programme, both of which are acknowledged for providing facilities
and financial support, respectively.
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The Herbarium and Arboretum of the Forest Research Institute of
Malaysia at Kepong — a Historical Perspective
K.M. WONG
Forest Research Institute of Malaysia, Kepong, Selangor, Malaysia.
Abstract
The beginnings of the Herbarium and Arboretum of the Forest Research Institute of Malaysia at
Kepong are documented, as are the roles of the people associated with these events and developments
from British colonial times until Malayanisation in the 1960s. The Herbarium was founded by Alfred M.
Burn-Murdoch, the first Chief Forest Officer of the Straits Settlements and Federated Malay States, in
1908. The Arboretum was begun in 1928-29, with James Watson (Forest Economist) and Frederick
Foxworthy (Forest Research Officer) playing significant roles in its establishment. The influence of the
government of the Straits Settkements and Federated Malay States and the roles of Burn-Murdoch,
Foxworthy, Watson, Symington and Wyatt-Smith, likewise of their colleagues and associates, in the
development of botanical studies in Malaya are traced from the beginning of the 20th Century, through
military occupation during the Second World War, until the 1960s.
Prologue
Early notions of placing the guardianship of the forests of the Malay Peninsula
under a more systematic arrangement may be traced to the time of the Straits
Settlements, a British Crown Colony consisting of Malacca, Penang, Province
Wellesley, Singapore, Pangkor and the Sembilan Islands. It was in 1879 that the
Colonial Engineer of the Straits Settlements, Major J.F.A. McNair, provided the
earliest record of forestry in a report that supplied descriptions of the main timber
trees and which recommended the formation of a forest department. It was not
until 1883 that a small department of forests in Singapore was created under the
Superintendent of Gardens, Straits Settlements, which position was held by Natha-
niel Cantley from 1880 until his death in 1888.
In 1895, when the states of Perak, Selangor, Negri Sembilan and Pahang formed
the Federated Malay States (each of which had a British Resident), the control of
forests in the Straits Settlements was transferred from the Director of Gardens to
the State Land Officers, as was the practice in the Federated Malay States. Henry
Nicholas Ridley (who succeeded Cantley), pointed out that this was not a satisfac-
tory arrangement for looking after all matters pertaining to forests. In a report on
the forests of Selangor in 1896 he thus recommended the establishment of a forest
department for the Federated Malay States (Mead 1936; Watson 1950). H.C. Hill,
commissioned in 1900 from the Indian Forest Service to report on forest adminis-
tration in the Straits Settlements and Federated Malay States, recommended the
appointment of a Chief Forest Officer to take charge of forests (Hill 1900a, 1900b).
This resulted in the transfer of Alfred M. Burn-Murdoch from the Burma Forest
Service in 1901 to the post of Chief Forest Officer of the Straits Settlements and
Federated Malay States.
The Beginnings of the Herbarium
Burn-Murdoch had, from 1901 (the year he arrived in Malaya) until his death in
Klang in 1914 (Anon. 1914), recognised the relevance of producing an account of
15
16 Gard. Bull. Sing. 40(1) (1987)
commercially important timber tree species of the Malay Peninsula. It was,
therefore, of direct concern to him to obtain first-hand knowledge of forest trees,
and so he developed an interest in collecting plants as reference material. His
earliest documented plant collections in Malaya date from 1903 (van Steenis-
Kruseman 1950). From 1904, Burn-Murdoch’s office was known as the Conserva-
tor of Forests, Straits Settlements and Federated Malay States. Burn-Murdoch
(1909) recorded in the 1908 Annual Report of the Forest Department that
‘during the year, a collector of specimens was employed by the Conservator, who
collected many herbarium specimens of forest trees ... submitted to Mr. Ridley ...
for identification ...”” and that ‘... a collection of the named specimens was
commenced and ... kept in the office of the Conservator for ... reference.”
In 1908, therefore, the Forest Department started a managed collection of
pressed plant specimens that was to develop into the present herbarium of the
Forest Research Institute of Malaysia at Kepong. Burn-Murdoch, the man who
founded this herbarium, had himself travelled extensively in the field collecting
plants in all states of present-day Peninsular Malaysia except Perlis, Kedah, Treng-
ganu and Kelantan (Burkill 1915; van Steenis-Kruseman 1950). He distributed
some of his collections to the herbaria of the Singapore Botanic Gardens and the
Royal Botanic Gardens, Kew.
Burn-Murdoch was a person of tremendous energy; it is to his credit that the first
reserved mangrove swamp forests were established for Perak and Selangor. His
book, Trees and Timbers of the Malay Peninsula (Burn-Murdoch 1911, 1912), is the
forerunner of forestry publications that form the series of Malayan Forest Records.
Despite his primary interest in forest trees, Burn-Murdoch’s collections encompas-
sed other groups of plants, including shrubs, lianes, palms, parasites, saprophytes,
ferns and aquatic herbs (Burkill 1915), a testimony to his wide interest in natural
history.
At the beginning of 1915, the year following that of his death, the Forest
Department herbarium consisted of 481 numbers, representing about 370 species
(Foxworthy 1918), collected by both British serving officers as well as local forestry
staff (many of these collectors were forest rangers) and accommodated in a single
small cupboard at the Forest Department premises in the Supreme Court building
on Court Hill in Kuala Lumpur. Under Burn-Murdoch’s successor, G.E.S. Cubitt,
accessions to this collection (often referred to as the ‘“‘local herbarium” in depart-
mental reports) continued, albeit slowly, and by early 1916 there were 500 numbers
representing about 385 species (Foxworthy 1918). In 1916, the Wray herbarium of
the Agriculture Department was transferred to the Forest Department in Kuala
Lumpur (Cubitt 1919).
With the potential of forest resources and a rubber-dominated agriculture in
Malaya becoming increasingly recognised, the scenario of the time saw a gradual
de-emphasis of agri-horticultural and forestry organization in Singapore, where the
Botanic Gardens had served the agro-economic development of the Straits Settle-
ments since 1875, and its centralization in Kuala Lumpur, the administrative centre
of the Federated Malay States (Furtado & Holttum 1960; Burkill 1983). In 1918, a
desire was already crystallizing to intensify botanical research based in Kuala
Lumpur where was housed the headquarters of the Department of Forests, Straits
Settlements and Federated Malay States, and the Department of Agriculture,
Federated Malay States. This was the year in which Cubitt secured the services of
Frederick William Foxworthy as the first Forest Research Officer of the Federated
Malay States and Straits Setthements, after which the size of the Forest Depart-
ment’s “‘local herbarium’ was substantially increased. Very much a botanist,
Foxworthy, upon his arrival in Malaya, spent a great deal of time acquainting
himself with the local flora and vegetation, and rearranging both the Wray herbar-
Herbarium and Arboretum at Kepong 17
ium (maintained separately) and the “local herbarium,” identifying, labelling and
recording a large amount of old and newly collected material (Cubitt 1919).
Foxworthy had the assistance of I.H. Burkill in identification, and both Burkill and
C.F. Baker, of the Singapore Botanic Gardens, together with Foxworthy and
several Forest Department staff members, became the main contributors to this
collection (Cubitt 1919). Foxworthy also distributed duplicates of plant collections
to Singapore and Kew (Cubitt 1919), from what was now called the Forest Herbar-
ium, housed in the same premises at Kuala Lumpur’s Court Hill.
Also in 1918, the Governor of the Straits Settlements (S.S.), Sir Arthur Young,
who was also the High Commissioner to the Federated Malay States (F.M.S.),
expressed to the Secretary of State for the Colonies the desirability of maintaining
work in systematic and economic botany outside the proposed joint S.S. & F.M.S.
Department of Agriculture. By December, 1923, a forestry committee formed by
Cubitt had met with A.S. Haynes, Secretary for Agriculture, S.S. & F.M.S., and
I.H. Burkill, Director of Botanic Gardens, S.S., to discuss the centralization of
botanical work in Malaya. They decided that the Singapore Herbarium, as well as
work in systematic and economic botany carried out by the Singapore Botanic
Gardens and the F.M.S. Museums Department, should be transferred to the
premises of a new Botanical Department to be established in Kuala Lumpur.
Furthermore, the Public Gardens in Kuala Lumpur was to be developed as a
botanic garden, and the Penang and Singapore gardens were to function as bran-
ches. These plans, as explained by Burkill (1983) in some detail, were not trans-
lated into reality. The rubber slump of 1924 affected revenue adversely and Sir
Lawrence Guillemard, then Governor/High Commissioner, did not overrule strong
opposition from Sir George Maxwell, Chief Secretary to the F.M.S., against
effecting the centralization of botanical research. Although Maxwell finally agreed,
just before his retirement in 1926, to the establishment of a new department (to be
known as Botanical Survey, Malaya) the S.S.-F.M.S. Government appears to have
lacked any further impetus towards realizing these objectives. In 1926, the Wray
herbarium, hitherto curated in the Forest Herbarium at Kuala Lumpur, was
incorporated into the Singapore Herbarium (I.H. Burkill 1927; van Steenis-
Kruseman 1950). H.M. Burkill (1983) records —
“In 1929. the world was perceptively sliding into another slump ... the Singapore Botanic Gardens
continued to provide the botanical service Malaya needed. The Forest Department was particularly
affected since a major aspect of tropical forestry is to be able to make reliable inventories of forest
species. Failure to bring the resources of the Singapore Herbarium to Kuala Lumpur led to the setting
up of specialist facilities within the Forest Research Institute, Kepong ...”
Under Foxworthy, the Forest Herbarium developed rapidly. By the end of 1919,
the herbarium (excluding the Wray collection) contained some 5000 numbers
representing 1700 species, of which about 1200 were trees (Cubitt 1920), and at the
close of 1920, this had increased to 6000 numbers representing about 2100 species
of which about 1600 were trees (Barnard 1921). Although Foxworthy is primarily
remembered for his role in organizing the beginnings of forestry research in
Malaya, his account of The Commercial Woods of the Malay Peninsula (Foxworthy
1921) is the first comprehensive work summarizing the botany of Malayan dipter-
ocarps in a form suited to foresters. This publication began the Malayan Forest
Records as its first number; his Dipterocarpaceae of the Malay Peninsula, issued as
No. 10 of the same series (Foxworthy 1932), was the first plant systematic mono-
graph produced at the newly established Forest Research Institute. It incorporated
the results of contemporary revisionary work and set the stage for a more compre-
hensive treatment of this important timber family. Foxworthy had been a botanist
with the Bureau of Science in Manila and held the same breadth of vision as
Burn-Murdoch in the attention he gave to plant groups of lesser commercial
Plate 1. F.W. Foxworthy (1877-1950), the first Forest Research Officer of the Federated Malay States
and Straits Settlements. His Dipterocarpaceae of the Malay Peninsula, published in 1932, was
the first plant systematic monograph produced at the then newly established Forest Research
Institute at Kepong.
importance. His account of Minor Forest Products of the Malay Peninsula (Foxwor-
thy 1922) testifies to this, and in his annual report for 1923 (Foxworthy 1924) he
noted: ‘“‘The most serious lack about our herbarium at present is room ,.. the
condition has become very acute with the beginning of our collection of rattans.”’ In
1926, a number of bamboo specimens were identified by the specialist J.S. Gamble
at Kew (Foxworthy 1927).
18
Herbarium and Arboretum at Kepong 19
With Foxworthy as Forest Research Officer, the Forest Research Institute (FRI)
was first functionally established at its present site at Kepong in 1926, when a forest
nursery and several experimental plantations were begun. Although the decision to
set up the FRI was first taken in 1921 it was not until August 1925 that the Regent
of Selangor was approached to approve 800 acres at Kepong for the purpose, and
only in 1929 was the main office building constructed (Anon. 1950b; Watson 1950;
Menon 1969), and the Forest Herbarium moved from its original site to a room on
the upper floor of the east wing of this building (Foxworthy 1930). The following
year, Foxworthy noted (Foxworthy 1931) that *... the herbarium now contains
material of about 70 percent of the known tree species of the Peninsula; ...” this
was based on his earlier estimate (Foxworthy 1918) of about 2200 species of trees
recorded from the Peninsula. During this formative period of the herbarium, in the
latter years of the 1920s, there was a continuing spirit of cooperation between the
Singapore Botanic Gardens and the F.M.S. Forest Department. Apart from the
assistance of botanists based at Singapore, the Forest Department had also en-
listed, in several assignments, the help of Mohamed Nur bin Mohamed Ghous, the
Herbarium and Museum Assistant of the Singapore Botanic Gardens. Mohamed
Nur was prominent among collectors who had accompanied Foxworthy in the field
(Burkill 1958). Foxworthy retired in 1932, after which the post of Forest Research
Officer was abolished and replaced by a conservatorship.
Watson and the Kepong Arboretum
The last years of the 1920s were significant in another respect. As the Forest
Research Institute settled down in its new premises at Kepong, it was recognized
that botanical and forestry research required the support of carefully planned living
plant collections. Even though experimental tree plantations at Kepong were
initiated in 1926, Foxworthy looked forward to the eventual specialised collections.
In a memorandum to the Forest Economist, James Gilbert Watson (No. 3 in
F.R.O. 189/27 dated 20th October 1927), Foxworthy wrote of palms: ‘‘A special
place might be set aside for a collection of the tall growing palms. Fairly good soil
will be needed and the plants should be set far enough apart to give them a chance
to spread ...”’ In effect this must have been carried out, although there is no
traceable record on where the “‘palm site’’ was; the oldest palm specimens to be
found in the arboretum records are a Bayas (Oncosperma horridum) and a Nibong
(O. tigillarium) planted much later in 1953, and today still standing side by side at
the intersection of Arboretum Road with the Sungei (River) Kroh.
James Watson (son of William Watson, once Curator of the Royal Botanic
Gardens, Kew) had had some practical experience at Kew as well as at Berlin when
he studied forestry at Eberswalde in Germany. He joined the Forest Department in
Malaya in 1913 as a Forest Officer and eventually succeeded Foxworthy to lead the
Forest Research Institute in 1932. His Malayan Plant Names (Watson 1928a) and
Mangrove Forests of the Malay Peninsula (Watson 1928b) had just appeared as
Malayan Forest Records No. 5 and 6, respectively. In a note (No. 3 in F.E. 93/27
dated 23rd September, 1928) to Foxworthy, Watson volunteered to begin establish-
ment of an arboretum:
“T should very much like the formation and care of this part of the Kepong scheme to be included in
my duties on my return from leave, more particularly in view of my past experience in this class of work,
and from the fact that I should be living in the middle of the area that it is proposed to dedicate to the
purpose. I discussed the matter on several occasions with the Conservator (Mr. Cubitt) who is, I believe,
substantially in favour of the principle on which I wish to work, and who pointed out the desirability of a
representative collection in view of the impending transfer of botanical research from Singapore to K.
Lumpur. When we discussed the subject a short time ago, I do not think that we were regarding it from
the same angle. I should like, therefore, to present the case as it appears to me ...
Plate 2. J.G. Watson (1889-1950), Conservator in charge of the Research Branch of the Malayan
Forest Service during 1932-1936. He was responsible for the establishment of the Arboretum
of the Forest Research Institute at Kepong in 1929, when he was Forest Economist.
“My idea of an arboretum is essentially a well-grown and representative collection of trees
maintained for purposes of reference and as a convenient source of seeds and herbarium material for
exchange ... In other words, it is primarily a living herbarium and, as such, it should be arranged as
systematically as is practicable.
‘It is clearly impossible to provide the trees in such a collection with the conditions that they will meet
with in nature, for it may be assumed that each has its peculiar optimum which is not likely to be found
in Open country covered with lalang //mperata cylindrica], following exhaustion of the soil from
root-crops, exposure, and the cumulative effects of Chinese vegetable gardening. Further, the wide
spacing that must be adopted to-allow the trees to develop, precludes any mutual shelter or natural
enrichment of the soil for many years to come. It will be necessary ... to give them more attention than it
is practicable or desirable to give to forest plantations.
20
Herbarium and Arboretum at Kepong 21
“The growth of individual trees in an arboretum cannot be expected to provide data of much
silvicultural value, though (particularly in the case of exotics) it may provide useful preliminary
indications and forestall needless expenditure on ... species that are foredoomed to failure through
unsuitability of climate ...
‘Apart from its scientific values, an arboretum at Kepong will serve a very useful purpose in the way
of advertisement and propaganda, for with it we can demonstrate our ability to grow trees at a price, and
thereby emphasize the difficulties with which we are faced in the field. And, finally, there is the aesthetic
aspect to be considered, and the desirability of ... encouraging the rapid and healthy growth of trees in
the neighbourhood of the office and residential portions of the reserve ...
“Tt will not be possible to adopt a systematic grouping according to families (as I originally suggested)
as supplies are likely to be too irregular ... but I suggest that one third of the area be definitely dedicated
to the dipterocarps, and a small portion to exotics ... Belukar [secondary forest] forms, and trees that
are commonly found and are easily accessible elsewhere at Kepong, will not ordinarily be planted,
though they may be retained if they happen to be growing in the arboretum area. Generally speaking ...
the commercial trees will take precedence if space is at all cramped ...
“But as ... it is not likely that funds will be available for planting this year ... I suggest, therefore, that
the available seedlings in the nursery should be earmarked and listed, and that we content ourselves this
year with marking out their positions in the field in order that planting may start at once in 1929.”
In the ensuing years following establishment of the dipterocarp and non-
dipterocarp sections of the Arboretum, departmental correspondence indicates an
active pursuit of this goal by both Watson and Foxworthy. These two sections were
to mature, many years later, into the two largest collections at Kepong, the
dipterocarp collection being the finest in the world.
Symington
In 1929 Colin Fraser Symington joined the FRI (Forest Research Institute) as
Assistant Conservator of Forests and began to assist in the running of the herbar-
ium; in 1934 he was designated the first Forest Botanist. Symington had envisioned,
in 1936, producing a foresters’ tree manual comprising all the Malayan timber-
producing families. However, it was obvious that much research in systematic
botany was still required and that a great amount of instability still existed in the
botanical nomenclature. He concentrated on, and became the authority on the
Dipterocarpaceae, the most important timber-tree family in Malaya and SE. Asia.
Symington completed his Foresters’ Manual of Dipterocarps (No. 16 of the
Malayan Forest Records) at Kepong in 1940. In November, 1941, typesetting of the
book had begun at the Caxton Press in Kuala Lumpur but by the following month,
both Symington and the manager of the press were forced to retreat to Singapore in
the face of the Japanese invasion. In January 1942, Symington with his family
boarded a ship for Australia (Corner 1981). At Singapore, E.J.H. Corner, Assis-
tant Director of the Singapore Botanic Gardens, learnt from Mrs. Symington that
the typescript had been left with the printer in Kuala Lumpur, but he was not
successful in persuading Symington to leave a second copy at the Singapore
Herbarium. Symington’s important work on the Dipterocarpaceae would have
been lost during the War, were it not for the timely initiative of Corner (then
confined with other staff members to the Gardens), who informed Hidezo Tanaka-
date, then acting Director of Raffles Museum in occupied Singapore, of the
typescript left in Kuala Lumpur. Tanakadate, a professor at the Tohoku Imperial
University of Sendai, understood the value of Symington’s work and intervened in
a similarly timely manner. He obtained the release of H.E. Desch (who had
worked closely with Symington at Kepong, and was author of The Timbers of the
Dipterocarpaceae, No. 14 of the Malayan Forest Records) from the Changi Military
Camp. Tanakadate and Desch travelled to Kuala Lumpur to find the galley proofs
of the manual kept partly at the Caxton Press by L.E. Labrooy (Desch 1962) and
partly at the FRI, Kepong (Tanakadate 1943). The proofs were corrected by Desch
Plate 3. C.F. Symington (1905-1943), designated the first Forest Botanist in the Forest Research
Institute at Kepong in 1934. He was forced to flee Malaya in 1942 in the advent of war, and
died in Nigeria the following year.
in Singapore at the Museum and the Changi PoW Camp, and the botanical names
were scru‘inised by Corner in the Gardens. The cost of printing 500 copies of
Symington’s manual (Symington 1943), at the Caxton Press in Kuala Lumpur, was
met personally by Tanakadate and the Marquis Yositika Tokugawa, then acting
President of the Raffles Museum and Library (Corner 1946). The book appeared
under the Japanese title of Malai Hanto no “‘Dipterocarpaceae’’ Mokuzai no
Hokoku, and was priced at 10 yen.
be
MALAI HANTO NO “DIPTEROCARPACEAE™
MOKUZAI NO HOKOKU
MALAYAN FOREST RECORDS
No. 16
FORESTERS MANUAL OF DIPTEROCARPS
by
C. F. SYMINGTON
Plate 4. The cover of the original issue of Symington’s Foresters’ Manual of Dipterocarps, issued by the
Japanese Administration in Singapore in 1943.
24 Gard. Bull. Sing. 40(1) (1987)
It is a credit to the scientific community that men like Tanakadate could in time
of war and hostility write: “‘scientific literature should not be obstructed by war”
(Tanakadate 1943). In spite of Symington’s departure from occupied Malaya, and
the events that led to the publication of his book, the FRI was never to have back
the botanist whose monograph of the Dipterocarpaceae was to become one of the
best known works in Malayan botany. Symington returned to England and was
posted to the forestry service in Nigeria where, in desperation, he took his own life
in 1943, unaware of the fate of his book.
The War Years at the Ringyo Shikenjyo
After war broke out on 8th December, 1941, there were no resident British
officers in charge at the Forest Research Institute, which the Japanese called the
‘“Ringyo Shikenjyo” (Forestry Experimental Station). V.L. Bain, by virtue of
being Eurasian, was exempted from detention and appointed the acting State
Forest Officer for Selangor by the Japanese Military Administration. Bain was able
to reappoint several local staff members at Kepong in 1942. Aziz bin Budin,
Technical Assistant to the Herbarium, reappointed in April 1942, was put in charge
of the Botanical Division of the Institute. Aziz reported for 1942 —
‘During the former regime 78 herbarium cases were filled with herbarium sheets ... The looters have
done considerable damage to this section. They removed almost all the cases, throwing all the valuable
specimens on the floor. Some sheets were also taken away ... The herbarium, a room of 33 ft by 50 ft,
was completely filled with mounted herbarium sheets filled to a foot depth. In addition to this the looters
removed four copper heads of the laboratory taps, thus letting water to overflow the room. The result
was that most of the herbarium sheets were soaked completely ... Those sheets which were beyond
salvage, were discarded ...
“It is estimated that about * of the original specimens of 43,000 sheets will be saved ...”
The plundering of the herbarium by looters was a grim prologue to the uncon-
trolled felling of forests for timber and food-growing that was beginning during
those war years.
In September 1942, Yukio Tsuji was despatched by the Japanese Military Admi-
nistration in Malaya to serve as the Chief Research Officer of the Forest Research
Institute at Kepong. Tsuji graduated from the Department of Forestry of Tokyo
University in 1919 and had been a research officer in the Japanese Government
Forest Experiment Station (known as the Forestry and Forest Products Research
Institute since 1958) before his appointment in Malaya. He remained in charge of
the Forest Research Institute at Kepong until the end of the War.
In his programme of reorganization of the herbarium, Aziz wrote —
‘All strewn herbarium sheets comprising nearly 30,000 in all to be critically examined, identification
supplied from herbarium registers and finally to be distributed to their various family group. Each family
is further to be ... rearranged into genus and species alphabetically ... kept in the herbarium cases, at
present 24 cases in all ... all salvaged sheets to be dried individually ... Naphthalene flakes or
Paradichlor benzene to be kept in the cases ... If index cards are available each herbarium sheet is to be
indexed.”
Aziz noted that ‘“‘missing sheets [were] to be replaced either by duplicates or by
new collection” and in fact during the year made collecting trips to Sungei Buloh,
Bukit Bruang, Bukit Lanjan and the Central Experimental Station at Serdang, with
the assistance of Forest Guards Sow bin Tandang and Tachun bin Baba, who were
reengaged in August. Sow had worked the previous twelve years together with
Symington. Aziz’s report for 1943 seemed less desperate:
‘Identifications were supplied to the Forest Department, majority of the requests were to identify
some of the Commercial Timber-trees of Malai [Malaya] ... Routine identifications ... has taken up a
disproportionate portion of the Technical Assistant’s time ...
Herbarium and Arboretum at Kepong 25
‘Some 46 mounted duplicates were sent to the State Forest Officer, Kelantan, as a guide to the
identification of trees ... Some specimens were also ... forwarded to the Navy Department, Syonan-to
[Singapore] ...
“The Technical Assistant together with one of the collectors accompanied the Director of Forestry,
on his inspection tour of Klang, where swamp jungle was studied.
“Acquisition of herbarium material during the year totalled 312 ... The figure seemed very small
indeed as compared to the past years ...””
‘Several Nippon Officers visited the Herbarium during the year — and some have spent good time in
studying plants of commercial utility.”
The visiting Nippon officers referred to in Aziz’s report included Professor
Ryujiro Ishida, who was despatched to survey research organisations in Malaya in
1943. Together with another officer named Utsuki, Ishida visited the Forest Re-
search Institute at Kepong on 17 April 1943. He subsequently wrote a report on 14
organisations he visited, including the Wood Technology Laboratory in Kuala
Lumpur and the Rubber Research Institute.
Watson, who succeeded Foxworthy as Conservator in charge of the Research
Branch of the Malayan Forest Service in 1932, and later, J.P. Mead as Adviser on
Forestry in October 1940 had retreated to Singapore with the advent of war and
was interned when Singapore fell on February 15th, 1942 to Japanese hands. He
was to spend three and a half years as internee, first at Changi Prison and then at
Sime Road (Anon. 1950a). Most of the remaining officers of the Forest Service
who had not left Singapore were also interned or were prisoners-of-war, until the
surrender of the Japanese on August 15th, 1945. Bain, as acting State Forest
Officer for Selangor, had appointed J.S. Wijasuriya as the Chief Clerk of the Forest
Research Institute, who took charge of the Institute during this dismal period since
28th July, 1942. Thus, Wijasuriya, together with Technical Assistants P.K. Balan
Menon and Aziz Budin, both reengaged along with 3 clerks and a skeleton team of
subordinate field staff and labourers, went about the business of reorganizing and
maintaining the facilities and work as best as possible under the clouds of war.
By the end of the War, the area under experimental plantations on the Institute’s
premises was reduced from 494 to 377 acres. Of a total of 363 trees in the
arboretum, 97 were felled or damaged irreparably by fire.
From War to Malayanisation
There was no hiatus following the cessation of war in Malaya in the August of
1945. Reassessment and rebuilding had to begin. Watson returned to serve as
Forestry Adviser to the Colonial Office during 1946-47 before retiring (Anon.
1950a) and F.H. Landon was placed in charge of the Institute from 1946 to 1948
(Menon 1969). The herbarium came under the charge of John Wyatt-Smith, who
served as Forest Botanist between November 1946 and May 1955. The Forest
Herbarium at Oxford University returned to Kepong herbarium, as a gift, its
duplicate material of specimens from the Kepong collection that were lost in the
war. Field collection of specimens regained momentum. In 1949 the herbarium was
rearranged (Anon. 1950c) according to the classification of Bentham & Hooker’s
Genera Plantarum; this system has been retained as the basic framework of classi-
fication since then. With the publication of Index Herbariorum in 1952 (Lanjouw &
Stafleu 1952), the FRI herbarium, Kepong, adopted KEP as its acronym.
Although in full realization that a companion volume to Symington’s Dipter-
ocarp Manual was essential, in that similar information on timber trees of other
families were needed, Wyatt-Smith was aware that the state of knowledge and
systematy of these other plant families could only be advanced gradually. He thus
produced a series of illustrated notes on the Burseraceae, Leguminosae, Myristi-
caceae, Sapotaceae, Lauraceae and Sapindaceae (Wyatt-Smith, 1953-54) that could
Plate 5. J. Wyatt-Smith (b. 1917), Forest Botanist at the Forest Research Institute at Kepong during
1946-1955, was later Forest Ecologist and Forest Silviculturist for some years.
serve as preliminary accounts and simultaneously would be of practical use to
foresters. Wyatt-Smith also worked with M.R. Henderson, then Director of the
Singapore Botanic Gardens, on revising the difficult genus Calophyllum for Malaya
(Henderson & Wyatt-Smith 1956). Such efforts, however, were interrupted.
Wyatt-Smith’s tenure as a forest botanist in the widest sense of the term saw him
first as Forest Botanist (November 1946 — May 1955, since 1951 as Forest Botanist
and Ecologist), then after a period of leave as Forest Ecologist (Botany) (Septem-
26
Herbarium and Arboretum at Kepong 27
ber 1958 — 1959) and subsequently as Forest Silviculturist (1959 — April 1963).
The nature of these various posts reflected a renewed emphasis on silvicultural
research and practices. Between 1950 and 1954, Wyatt-Smith and R.C. Barnard
(the latter then Silviculturist) were put mainly in charge of producing a contempor-
ary manual of silviculture which was the first priority of the Silvicultural Division.
Although Barnard had published a preliminary treatment of the main scope of
silvicultural work (Barnard 1954), he retired in 1956 by which time Wyatt-Smith
had been transferred on field duty to Kedah as its State Forest Officer. In 1957,
G.G.K. Setten, then Chief Research Officer, petitioned for the return of Wyatt-
Smith to complete the manual of non-dipterocarps and of J.E. Cousens, formerly
Instructor of the Forest School at Kepong, to complete the manual of silviculture.
But Cousens left the Forestry Service in 1958 and circumstances dictated the
transfer of Wyatt-Smith from botany to silviculture (Mohammed Alwy, foreword in
Wyatt-Smith 1963).
If botanical work towards the non-dipterocarp manual of timber trees was
interrupted by these circumstances, it also amply displayed the relevant association
between forest botany, ecology and silviculture. With broad perceptions accruing
from much field experience in these areas, Wyatt-Smith published the bulk of the
work in his manual of silviculture (Wyatt-Smith 1963). His ecological research
plots at Bukit Lagong and Sungei Menyala, set up in 1947, were to become,
decades later, among the oldest study plots in the tropical world.
With the approach of the formation of the Federation of Malaya and thereafter
the Federation of Malaysia in 1962, the transition towards Malayanisation came
into being. Kizhakkedathu Mathai Kochummen, working with Wyatt-Smith as
Research Assistant from January 1953 to May 1955, and Assistant Botanist from
June 1955 to June 1971, took sole charge of the herbarium during 1960-63. The
targetted deadline for Malayanisation of the Research Branch was 1965. In 1963,
G.G.K. Setten, the last expatriate Chief Research Officer, left and was succeeded
by Abdul Rahman bin Mohamad Ali. The last foreigners to leave the Forest
Research Institute were A.J. Vincent, Deputy Chief Research Officer, and B.A.
Mitchell, Afforestation Officer. Francis $.P. Ng was recruited in June 1964 as
Forest Botanist.
When an extension to the main office building was completed in 1965, the
herbarium moved, in May, to the top floor of the new block. By 1965, the
collection numbered 74,694 specimens. With this new scheme of things, and
Kochummen and Ng attached to Forest Botany, the non-dipterocarp manual was
again reemphasized and Timothy Charles Whitmore was engaged under the Col-
ombo Plan Aid to reorganize the project to produce a Tree Flora of Malaya.
Whitmore was with the FRI between September, 1965 and April, 1972. Kochum-
men became Forest Botanist in 1971 and Senior Forest Botanist in 1980, and Ng
was Senior Forest Botanist from 1974 until 1978, subsequently assuming the post of
Assistant Director of the Institute. The first volume of the Tree Flora of Malaya
(Ed. Whitmore 1972), largely written by Kochummen, Whitmore and Ng, also
received contributions of family accounts from other botanists in the region, viz.
Peter F. Cockburn, Hsuan Keng and Benjamin C. Stone.
The development of the Herbarium and Arboretum subsequent to Malayanisa-
tion, and the individual achievements of Kochummen and Ng, are not here dealt
with. The present account chronicles the beginnings of the Herbarium and
Arboretum of the present Forest Research Institute of Malaysia at Kepong, and
traces the roles of the people associated with these events and developments until
Malayanisation in the late sixties.
25 Gard. Bull. Sing. 40(1) (1987)
Acknowledgements
I thank the following for providing helpful comments and information: Mr. H.M.
Burkill, Dr. K.L. Chang, Professor E.J.H. Corner, Dr. J. Dransfield, Professor
R.E. Holttum, Mr. K.M. Kochummen, Mr. T. Matsumoto, Dr. F.S.P. Ng and Dr.
T.C. Whitmore. I am grateful to Khoon Cheong for assistance during the prepara-
tion of the manuscript.
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Tree Species Enumeration of 0.5 Hectare on Halmahera
T.C. WHITMORE
Department of Plant Sciences, Oxford University, England,
K. SIDIYASA
Forest Research & Development Centre, Bogor, Indonesia,
T.J. WHITMORE
St John’s College, Cambridge, England.
Abstract
A plot of 0.5 ha in east central Halmahera in the Moluccas had 76 species, 31 families and 371 stems of
trees > 10 cm dbh. It differed substantially from plots enumerated previously in Seram.
It is well known that within Malesia once one crosses Wallace’s Line eastwards
the tree flora becomes poorer. Many rain forests in fact show greater richness in the
western (Sumatra, Malaya, Borneo) and eastern (New Guinea) regions of the
archipelago than in the centre (Sulawesi and the Moluccas), and some groups are
very rich in species at either west or east. These aspects of floristics are reflected by
tree-species numbers on small plots.
There are a few counts which show some very rich forests in western Malesia; the
largest counts ever recorded for rain forest come from Sarawak (223, 214 species/
hectare of trees => 10 cm dbh; Proctor et al. 1983). For Kalimantan there are
records of 149, 129 and 128 spp./ha (Kartawinata ef al., 1981), while for Papua
c. 184, c. 181, c. 152, c. 145 spp./ha have been recorded (extrapolated from plots of
0.8 ha area; Paijmans, 1970). There are very few counts from central Malesia.
The exact location and shape of a small plot is likely to affect the number of
species counted. Shortage of time has invariably prevented workers from simply
adding more subplots to reach the asymptote of the species-area curve. This
measure of abundance in species is thus not entirely satisfactory, but an accumula-
tion of plot counts from different places will eventually give some fairly objective
data on forest composition.
Thus, we offer here the first enumeration made on Halmahera, an island in north
Moluccas, as a modest contribution to forest quantification.
The site lay on a broad ridge crest at 630 m over shale at 0°30°N, 125°30°E, near
the logging camp at Tapayo, 20 km southeast of Dodinga on Telok Kau, on the
central mountainous spine of the northeastern peninsula. The soil was a shallow
reddish clay only c. 40 cm thick. The surface had a continuous thin leaf litter. The
tallest trees were c. 40 m. Big woody climbers were rare. Most boles were clean of
epiphytes and of adherent climbers (of which the most abundant were two
Freycinetia spp. both only ‘occasional’). There was a dense seedling carpet. The
forest was mostly in the mature phase of the growth cycle, with a few small gaps
formed by one or two fallen trees (Fig. 1). Otherwise big trees were ‘dying on their
feet’, disintegrating in situ, and forming virtually no gaps.
The whole region had been recently heavily logged and we were only able to find
a homogeneous intact patch of less than one hectare. Within this patch we enumer-
ated all trees > 10 cm dbh on a single half-hectare plot, made up of 11 subplots of
0.04 ha plus 3 of 0.02 ha (Fig. 1).
20M
Fig. 1. Plot layout and forest phase diagram. Gap-phase forest is left unshaded. Rest is mature phase.
In Fig. 2 we present the species-area curve, which shows signs of flattening out.
In Table 1 we present a summary of floristics. Myrtaceae, Guttiferae and
Lauraceae dominate the forest both in terms of species and stem numbers. We had
a total of 76 species in 31 families on the half hectare, and 371 stems.
The only other published plots we know of for the Moluccas were enumerated on
Seram island. There, on an area of 8 ha, comprising four transects each 20 x 1000m,
only 54 species in 27 families were encountered (Sidiyasa & Tantra 1984). Such
long belt transects are likely to maximise species recorded because of their length,
Spp.
20
40
30
20
0:1 0:2 0:3 0:4 0:5 ha
Fig. 2. Species-area curve.
32
Tree Enumeration on Halmahera
Table 1. Families in descending sequence of abundance on the 0.5 ha plot.
(For species authorities see Whitmore & Tantra in press)
Myrtaceae
Guttiferae
Lauraceae
Burseraceae
Myristicaceae
Euphorbiaceae
Theaceae
Anacardiaceae
Rubiaceae
Fagaceae
Sapindaceae
Dilleniaceae
Leguminosae
Meliaceae
Araucariaceae
Dipterocarpaceae
Melastomataceae
Celastraceae
Ebenaceae
Juglandaceae
Sapotaceae
Cunoniaceae
Flacourtiaceae
Palmae
Ulmaceae
Annonaceae
Elaeocarpaceae
Lecythidaceae
Moraceae
Rhizophoraceae
Rosaceae
TOTALS
Voucher specimens either deposited at BZF (most are solely fallen leaves) or, for TCW collection
11 Eugenia spp. (30+ 7+9 x 1)
Calophyllum soulattri (23 stems)
3 Calophyllum spp. (35 + 3 + 1)
Garcinia celebica (31)
SGorcmaspy. (6 73-F lt)
Cryptocarya sp. (1)
Endiandra rubescens (11)
2 Litsea spp. (9 + 7)
3 Lauraceae indet. (1 '+-1 +-1)
Canarium vulgare (1)
2 Canarium spp. (23 + 1)
Santiria sp. (1)
2 Burseraceae indet. (2 + 1)
Gymnacranthera forbes (3)
Gymnacranthera sp. (1)
Horsfieldia sp. (5)
Myristica sp. (7)
Myristicaceae indet. (2)
Blumeodendron sp. (1)
Macaranga sp. TCW 3621 (28)
Macaranga sp. TCW 3680 (2)
Adinandra sp. (1)
Laplacea sp. (3)
Ternstroemia sp. TCW 3669 (4)
Dracontomelon dao (1)
Gluta sp. (1)
Semecarpus sp. (1)
Gardenia sp. (1)
Nauclea mitragyna (1)
Nauclea sp. (2)
Castanopsis buruana (6)
Lithocarpus celebicus (17)
Pometia pinnata (4)
Sapindaceae indet. (1)
2 Dillenia spp. (3 + 1)
Archidendron sp. (2)
Cynometra ramiflora (1)
Aphanamixis polystachya (1)
Aphanamixis sp. (2)
Agathis dammara (11)
Anisoptera thurifera (10)
Memecylon sp. (9)
Lophopetalum sp. (4)
Diospyros lolin (4)
Engelhardtia serrata (4)
Planchonella moluccana (3)
Schizomeria serrata (2)
Flacourtiaceae indet. (2)
Pinanga ternatensis Scheff. (2)
Gironniera subaequalis (2)
Popowia sp. TCW 3605 (1)
Elaeocarpus sp. (1)
Barringtonia sp.-TCW 3639 (1)
Ficus sp. TCW 3642 (1)
Gynotroches axillaris (13)
Maranthes corymbosa (1)
31 families
series, at BO, BZF, K and L.
Total
species
11
GK
th
i) NM bh
in)
ne ee ee ee ee ee _ _s
76 species
Total
stems
46
104
31
31
= Se eS ON Nw BB DP
371 stems
34 Gard. Bull. Sing. 40(1) (1987)
in contrast to compact plots. On the Seram plots no family was well repre-
sented. Myrtaceae (6 species), Euphorbiaceae (5 species) and Ebenaceae, Gut-
tiferae, Moraceae, Sapotaceae and Sterculiaceae (3 species each) were richest.
Both sites lie in rain forest climates, types A (Halmahera) and B (Seram) in the
Schmidt & Ferguson (1951) scheme (see map Fig. 4.1 in Whitmore 1984). Neither
forest had any noticeable seasonal-climate floristic element.
Stem number on our plot at 742/ha is high in comparison with most rain forests.
The Seram plot at 390 stems/ha is about average (see Table 1 in Whitmore &
Sidiyasa 1986).
The Halmahera and Seram samples are very different from each other. In the
long term, interpretation of variation in forests between different parts of Malesia
needs some kind of quantitative data base, towards which we publish this enumera-
tion.
References
Kartawinata, K., Abdulhadi, R. & Partomihardjo, T. (1981). Composition and
structure of a lowland dipterocarp forest at Wanariset, East Kalimantan, Malay.
Forester 44, 397-406.
Paijmans, K. (1970). An analysis of four tropical rain forest sites in New Guinea. J.
Eeol58 71-11.
Proctor, J., Anderson, J.M., Chai, P. and Vallack, H.W. (1983). Ecological studies
in four contrasting rain forests in Gunung Mulu National Park, Sarawak. I.
Forest environment. J. Ecol. 71, 237-60.
Schmidt, F.H. & Ferguson, J.H.A. (1951). Rainfall types based on wet and dry
period ratios for Indonesia with western New Guinea. Verh. Djawatan Met. dan
Geofisik., Djakarta 42.
Sidiyasa, K. & Tantra, I.G.M. (1984). Analisis flora pohon hutan dataran rendah
Wae Mual, Taman Nasional, Seram-Muluku. Buletin Penelitian Hutan 462,
19-34.
Whitmore, T.C. (1984). Tropical rain forests of the Far East. Clarendon, Oxford.
—____ . & Sidiyasa, K. (1986). Composition and structure of a lowland
rain forest at Toraut, northern Sulawesi. Kew Bull. 41, 747-755.
. & Tantra, I.G.M. (in press) Tree Flora of Indonesia Check List for
Maluku. Forest Research and Development Centre, Bogor.
Additions to ‘The Use of Tifgreen and Tifdwarf Bermuda Grasses in
Two Singapore Golf Courses’
WONG Yew Kwan*
89 Soo Chow Garden Road, Singapore 2057
Further to my recent publication of the report (Wong, 1986) on the performance
of the two tifgrasses, Tifgreen and Tifdwarf, in the golf courses at the Tanah Merah
Country Club (TMCC) and the Sentosa Golf Club (SGC), I was fortunate to have
the opportunity of revisiting the Jagorawi Golf and Country Club near Bogor,
Indonesia, on 7 January 1987. My first visit was six years ago when the first 9 holes
were just ready for play. At the time the tees were covered with pure Santa Ana
grass while the fairways and greens were respectively turfed with Tifgreen and
Tifdwarf. It was already noticed that broad-bladed local grasses such as Axonopus
compressus (Cow Grass) and Paspalam conjugatum were beginning to invade the
fairways and workers were seen busy weeding these out. The resident superinten-
dent at the time, however, assured me that with minimum weeding the turf could
be kept pure. This has turned out to be otherwise.
As Jagorawi was our source of the grasses for TMCC and SGC, I made it a point
to look very closely at the turf as we played along so as to establish a visual
comparison between the situation in Singapore and that at Jagorawi vis-a-vis the
fate of the introduced grasses. Jagorawi, just like TMCC and SGC, used Santa Ana
for the tees, Tifgreen for the fairways and Tifdwarf for the greens, and just like our
two courses in Singapore the intention was to keep the turf as pure cultures. After
having seen the whole course during the round of golf, I have to say that the present
situation at Jagorawi is very similar to that at TMCC and SGC — except for the
greens, the Tifgreen in the fairways and the Santa Ana on the tees have almost
been totally replaced by other grasses and sedges.
The whole range of species of the grasses and sedges occurring in TMCC and
SGC could be found in Jagorawi but the density patterns are very different in the
two places. In TMCC and SGC there are more of the coarser species, such as
Brachiara distachya, Axonopus compressus, and Cyperus radians whereas in Jagor-
awi, species with finer habits have come into dominance. These are the Serangoon
Grass, Digitaria didactyla, a species of Eragrostis and patches of Polytrias amaura.
The rather fine sedge, Cyperus kyllingia, is also common but not as dominant as in
the two courses in Singapore. With the finer grasses dominating the scene, the
Jagorawi course has a much more uniform look than our two courses.
Cyperus radians, the coarse sedge with a tufted rosette habit, found so commonly
in the two courses in Singapore, is hardly seen at Jagorawi. This may be due to the
better drainage of the volcanic soil in the latter place. It may be recalled that this
sedge comes into prominence when there is locally impeded drainage. That this
should happen on a course constructed with a sand overburden on clayey fills, such
as is done in Singapore, may seem surprising. Sand is supposed to be rather free
draining. Unfortunately the sand overburden that was laid is not deep enough in
certain spots, and since the underlying clay is rather impervious, water tends to be
retained at the distinct interface, particularly if the topography is flat at the spots in
“Mr. Wong is a founder member of TMCC and was at various times serving on its management
committee and the Sentosa Golf Board.
35
36 Gard. Bull. Sing. 40(1) (1987)
question. This would cause local ponding during wet spells or when the place is
over watered. Volcanic soil on the other hand is well known for its better structure
and this accounts for better drainage and hence the absence of this hydrophilic
sedge.
Reference
Wong, Yew Kwan (1986). The Use of Tifgreen and Tifdwarf Bermuda Grasses in
Two Singapore Golf Courses. Gard. Bull. Sing. 39: 203-214.
Morphological Adaptations for Bird Pollination in Nicolaia elatior*
(Jack) Horan (Zingiberaceae)
REGINE CLASSEN
Institute of Botany, Department of Plant Morphology
R.W.T.H. Aachen, West Germany
Abstract
A report is given in detail on the behaviour of Anthreptes malacensis when visiting the inflorescence of
Nicolaia elatior, with special reference to the Singapore Botanic Gardens. It is shown how in Nicolaia
ornithophily became a realistic alternative to psychophily by the formation of a dense inflorescence unit,
the main steps of which being the aggregation of flowers and the expansion of (sterile) bracts.
Nicolaia elatior is often cultivated in tropical gardens. Its popularity is attributed
to the spice which the flower buds provide and the ornamental value of the
intensively red and glossy blossom? (plate 1) attracting nectar-searching animals,
especially birds. Knuth (1904, 183) had observed sunbirds on the flowering heads of
Nicolaia elatior, but he took them for ‘more harmful than useful’ visitors. His
Plate 1. The flower head of Nicolaia eliator surrounded by showy, extrafloral bracts.
Within the inflorescence, the open flowers form a whorl and present their yellow signals
(arrow); below, ripening fruits; above, flowers still in bud.
* More familiar synonyms are Elettaria speciosa Bl. and Phaeomeria magnifica (Roscoe) Schum. For a
more complete list see Valeton (1921: 138), Burtt and Smith (1972: 210-211), and Weber (1980: 152).
Presently, Etlingera elatior (Jack) R.M. Smith is the correct name. This new combination (Smith,
1986) has reduced Nicolaia elatior to a synonym.
+ The term ‘blossom’ is used in the sense of Faegri & v.d. Pijl (1979: 21), that is as an ecological term. In
the morphological point of view, a blossom may be an inflorescence (pseudanthium), a flower
(euanthium) or a part of a flower (meranthium).
37
38 Gard. Bull. Sing. 40(1) (1987)
opinion was that the birds visited the blossoms only for catching insects there, and if
they happened to insert their bill into the blossoms it was only incidental and
pollination was not effected. My observations at the Botanic Gardens in Singapore
strongly supports the view that Anthreptes malacensis is a pollinator.
Nicolaia elatior (Jack) Horan is a perennial herb which grows naturally in
primary and secondary forests in SE. Asia. Frondose leaves on the vegetative stems
measure several meters long and loosely shade over the reproductive, shaft-like
stems, which are 1-2 m long. On these are exposed a terminal flower head. Apart
from the large showy leaves which are integrated into the habit of the blossom these
reproductive stems bear only small bracts.
The inflorescence looks like a simple cephalium with about 300 flowers spirally
set on an undetermined apex. But recently, Weber (1980: 158) and Kunze (1985:
127) have shown that this flower head is a simple, highly reduced polytelic thyrsus.
The base of the flower mass is surrounded by large showy bracts, which form a
pseudocorolla of 20-cm diameter (pl. 1) and are the most conspicuous element of
the blossom. Except for a narrow pale margin, both sides are intensively carmine
and show a fine brightness which is caused by the waxy surface. The proximal
leaves are sterile and follow on abruptly the inconspicuous bracts of the shaft while
the distal, flower-subtending bracts gradually reduce in size. Before flowering, the
showy leaves completely envelop the flower buds; then, during anthesis they stick
out horizontally and finally, in the postfloral phase, they hang down retrorsely.
Anthesis of a head is 3 weeks. Every day, the flowers of a whorl would open
simultaneously and fade a few hours later. Thus the flowering zone shifts upwards
along the inflorescence axis which elongates from 5 to 15 cm. While the flowers at
the top are still in bud, young fruits would have developed at the base.
The flower is sessile and zygomorphic (pl. 1, figs. 1 & 2). Its characteristic
boat-shape is brought about by the labellum* which forms a staminodial tube
together with the fertile stamen (fig. 2). The style is — in the typically zingiber-
aceous manner — fixed in its position by the anther (fig. 4). Pollen-sacs and stigma
are oriented towards the labellum (fig. 3). By the unique arrangement of the
labellum, anther and stigma, pollination occurs whenever a sucking organ is dipped
into the tube (fig. 5) — as Knuth (1904) has already described in some detail.
* Weber (1980) interprets the labellum as a homologon to the two inner staminodes of the zingiber-
aceous flower. Kunze (pers. comm.) however is convinced that the labellum is represented by all the
five staminodes together.
Facing Page
Fig. 1 Habit of a flower of Nicolaia elatior.
pr, prophyll; ca, calyx; co, corolla lobe; /ab, labellum.
Fig. 2. Diagram of a flower of Nicolaia elatior (empirical, see note 3 on p. 41).
sb, subtending bract; ax, axillary bud; sta, fertile stamen; ep, epigynous gland; gyn, gynoecium;
solid: staminode tube.
Fig. 3. Optical longitudinal section showing the position of the anther-stigma-complex in relation to
the labellum.
sty, style; th, theca; ko, connective; fil, filament; hb, hairy belt.
Fig. 4. A flower of Nicolaia elatior, \ongitudinally opened.
sti, stigma; dl, dehiscence line of pollen sac; hg, hairy groove of the filament.
Fig. 5. Flower of Nicolaia elatior as pollination apparatus.
to, tongue; bi, bill; — pollen transference; nr, nectar reservoir.
40 Gard. Bull. Sing. 40(1) (1987)
Nearly all parts of the flower are more or less red coloured. Exceptions are the
pale, hardly visible anther and the very conspicuous, intensively yellow margin of
the labellum. The open flowers present these yellow parts like signals pointing to
the nectar-store. This may be of great importance, because the flowers lack — at
least not detectable — any alluring odour. Nectar is produced in great amount. It is
secreted by epigynous glands at the bottom of the flower and is stored in the
staminode tube, which functions as a nectar-reservoir (fig. 5). As is shown in fig. 4,
the inner surface of the tube is partly covered with long hairs. The filament is
densely hirsute and, approximately in the middle of the tube, there is a thick hairy
belt. Here, the hairs are arranged around two cuticulous protuberances running
diagonally from the filament border to the median of the labellum. It is quite
reasonable to assume that these hairs let the nectar move up by capillary absorp-
tion.
Upper right
Plate 2. A male Anthreptes malacensis (scale: 2 cm). The skin was kindly made available by the
Zoological Museum in Bogor, Indonesia.
Left
Plate 3. A male Anthreptes malacensis visiting the flower head of Nicolaia elatior in the Botanic
Gardens, Singapore.
Lower right
Plate 4. Askin of Anthreptes malacensis with the head and bill artificially inserted into a flower tube to
simulate the natural position when pollinating (nat. size).
I had the opportunity to observe for three weeks, flowering plants of Nicolaia
elatior and to record the blossom visitors. The most frequent and conspicuous was a
sunbird with a metallic sheen, the male Anthreptes malacensis (pl. 3). At intervals,
about 2-3 times hourly, it flew from the nearby Pandanus-thicket directly to one of
the flowering heads and landed on it.
There was no insect-picking at all though a lot of small ants, spiders and others
were available. After a short look around — probably to check on the safety of the
place — the bird inserted its bill into an open flower. It remained in that position
Pollination of Nicolaia elatior 4]
for up to 5 seconds. After that, the bird dipped its bill systematically into a second,
third and fourth flower. Its movement was restricted to a mere turnabout, step by
step, such that the bill was always exactly above the next open flower. Standing on
the top of the blossom, the bird only had to bend the front part of its body
downwards to reach the flower tube (pl. 3). As long as the bird was undisturbed,
it exploited up to 7 flowers per inflorescence, thus, less than half of the open
flowers in a single turn. Generally, before flying back into the thicket, it would look
round for more, then visit a second and sometimes even a third inflorescence. In its
resting place, the bird was mostly engaged in cleaning its bill and its plumage. It
rested for 10 minutes or as much as an hour before it returned to visit the blossoms
in the described manner.
Obviously, the blossoms of Nicolaia elatior attract Anthreptes malacensis Scop.
Though this observation was made in an artificial setting, I am convinced that
Anthreptes malacensis is one of the natural pollinators of the plant. This is sup-
ported by the remarks of Werth (1915) and Porsch (1924) concerning bird pollina-
tion in SE. Asia. Besides that observation, I have seen another male Anthreptes
visiting Nicolaia elatior, this time in a natural site near Bogor, W. Java. In general
Anthreptes malacensis is very common in SE. Asia and shares the same biotope
with Nicolaia elatior (King et al. 1984: 413). As with all Nectarinidae, it is characte-
rized by a specialized tongue which is well constructed for sucking honey: the tip is
forked and able to lap up drops of nectar while the tongue itself is folded over its
whole length forming, together with the bill, a perfect sucking tube (Werth 1900:
256-7).
With regard to bird pollination,
1. it is clear that Nicolaia elatior first of all attracts the bird optically by the
intensive red colour of the blossom — a colour which is known to be quite common
in ornithophily. The concentration of the flowers to a highly compacted head and
the enlargement of the extrafloral bracts promote the attractiveness of the
blossom. In terms of phylogeny, the primary function of the bracts probably has
been to protect the flowers against rain (Knuth, 1904: 182) which was then followed
by another evolutionary step, the transformation of the whole blossom into an
attraction apparatus. A comparable shifting of function of plain bracts from phy-
logenetically old structures to attraction units can also be seen in the blossoms of
some Verbenaceae (Classen 1986).
Presumably the display of the blossom plays an important role in the efficient
attraction of a pollinator, which is able to vagabondize distantly. In Nicolaia elatior
it is the stem again which provides the bird with a lookout to exploit nearby flower
tubes. As the inflorescence axis slowly elongates during the flowering period, there
is a constant distance between the bird and the opened flowers. This is, of course, a
very favourable prerequisite to a successful bird pollination. It is the direct con-
Sequence of the aggregation of flowers. So, even the aggregation seems to be an
equipment of prior rank for bird pollination.
2. Nicolaia elatior provides a landing and sitting place to the bird. It is the top of
the blossom which is enlarged by the dense aggregation of bracts and flowers. And
it is the stem again which provides the bird with a lookout and to exploit nearby
flower tubes. As the inflorescence axis slowly elongates during the flowering
period, there is a constant distance between the bird and the opened flowers. This
is, of course, a very favourable prerequisite to a successful bird pollination. It is the
direct consequence of the aggregation of flowers. So, even the aggregation seems to
be an equipment of prior rank for bird pollination.
3. Nicolaia elatior produces a high amount of nectar to reward the bird for its
visit. The nectar is clear and liquid. The concentration of sugar (taken at random
with a field refractometer) was 18-28% in the open flowers and only 5-8% in the
4? Gard. Bull. Sing. 40(1) (1987)
nectar of just fading flowers, which are not visited any more by the bird. The
presentation of the contrasting yellow signal helps the bird to find the entrance to
the nectar store. Pollination should take place as soon as the bird has dipped its bill
into the flower tube: At first, the pollen brought along from another flower is wiped
off on the stigma and then the pollen of the flower actually under visitation is
deposited on the underbill (fig. 5). As the pollination mechanism is very much
the same for all kinds of pollinators, Knuth’s description (1904: 182) concerning
pollination by butterflies is acceptable for bird pollination too. The small membra-
nous lip below the stigma assures that the pollen will not touch the stigma when the
bill is retracted.
According to Knuth (1904: 183) the flowers are said to be damaged by the hard
and curved bill. This was the main reason that he did not consider that bird
pollination takes place in Nicolaia elatior — an objection which has already been
rejected by Werth (1915: 370). I did not notice any flower being damaged by the
bird — all staminodial tubes remained completely intact. Obviously, the flower is
very well adjusted to the bill, the staminodial tube and the bill showing a high
similarity in curvature (fig. 5, pl. 4). As the labellum is hardened on its abaxial side
and equipped with a slippery inner surface, its construction is just a slide and would
function as a leading groove for the bill. The ovary is protected by its hypogyny and
by the length of the flower tube (up to 5 cm). As the nectar 1s capable of rising
within the tube, it is by no means necessary that the bill (or tongue or proboscis of
any visitor) is as long as the tube. So, even bees and butterflies with quite short
sucking organs can become successful pollinators (see Knuth 1904: 182).
4. Nicolaia elatior flowers continuously. There are always several inflorescences
of different ages on any one plant and each of them has a flowering period of about
3 weeks. Thus the plant probably can make nectar available to its visitors through-
out the year.
Without doubt, the blossom of Nicolaia elatior is adapted to bird pollination.
But, it is not exclusively an ornithophilous blossom, it is visited as well by diverse
butterflies especially Lycaenidae and Danaidae. The insects sit down on one of the
outer bracts and insert their sucking organ bowlike from below into the flower
tube. Though the sucking behaviour is quite different, the success of pollination is
unquestionable. In general, ‘the differences between the psychophiles and
ornithophiles are rather indistinct’ (Faegri & v.d. Pijl 1979: 126) and so it is idle to
separate them artificially.
In terms of phylogeny we can draw a consistent line: recalling the specific
parameters which promote bird pollination it is clear that in the case of Nicolaia
elatior the transition of the zingiberaceous inflorescence into an inflorescence-
blossom (pseudanthium) has been the most important step. While psychophily —
based on the flower structure — is also possible without any special aggregation and
modification of the inflorescence, the development towards ornithophily only
depends on the special inflorescence-construction, especially on flower aggrega-
tion, bract integration and display by a shaft-like stem.
So, the flowering head of Nicolaia elatior shows an example for the evolution of
an inflorescence-blossom, one that opens new pollination chances for a certain —
species. This may also help to understand how the succession of well operating
pollinators happened in the evolutionary process.
Pollination of Nicolaia elatior 43
Acknowledgements
I am very grateful to Dr. Chang Kiaw Lan, Curator of the Herbarium at the
Singapore Botanic Gardens, for the working facilities at the Garden, and to Dr.
Sao Kyu Win, Singapore Polytechnic, for her hospitality during my stay in Singa-
pore. I also like to thank the Deutsche Forschungsgemeinschaft, which has given
financial support (Cl 81/2-1).
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King, B. Woodcock, M. & E.C. Dickinson (1984). A Field Guide to the Birds of
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Knuth, P. (1904). Handbuch der Bltitenbiologie I1./1. — Leipzig.
Kunze, H. (1985). Die Infloreszenzen der Marantaceen und ihr Zusammenhang
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Porsch, O. (1924). Vogelblumenstudien I. J. wiss. Bot. 63.
Smith, R. (1986). New combination in Etlingera Giseke (Zingiberaceae). Notes
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Ecology of the Trees of Bukit Timah Nature Reserve
WONG YEw Kwan*
89 Soo Chow Garden Road, Singapore 2057
Abstract
An inventory was carried out in Bukit Timah Nature Reserve to study the specific composition of the
trees. For the sampling a systematic layout was used. The sampling unit was a cluster, with 4 circular
tenth acre sub-units, so that the area of each cluster is 0.4 acre. Enumeration was done for all trees down
to 24 inches (c.61 cm) gbh but for each cluster, in one of the circular subunits enumeration was down to
12 inches (c.30.5 cm) gbh. The idea was to have smaller-girthed trees to indicate recruitment.
In all, the 20-cluster sample, covering some 8 acres (3.24 ha) yielded 889 trees, belonging to 44 known
families with 212 species. Twenty individual trees in 13 clusters could not be identified at all.
The specific composition of the forest conforms to that of a Coastal Hill Forest according to
Symington’s classification of the forests of Peninsular Malaysia. The dipterocarps show family domi-
nance with 125 individuals out of a total population of 889 trees while the family Euphorbiaceae show
the highest number of genera (11) and the greatest number of species (22). The other families with
species preponderance are the Anacardiaceae, Burseraceae, Lauraceae, Myrtaceae and Leguminosae.
A parameter, called the Distribution Index (DI), is used to indicate spatial distribution (lateral
spread) of the species within the community. This is simply the number of clusters in which the species
occurred, its number of individuals being ignored. It is reckoned that this parameter, tabulated side by
side with the sample total of the species affords a quick visual appreciation of the ecological influence of
the species. In the sample, 121 species have been found to have a DI of unity, 41 with 2, 13 with 3, etc.
Only 5 species have a DI equal to or exceeding 10.
A stand curve, and a species/area curve are given. The stand curve, an inverted “‘J’’, shows the typical
content of uneven-aged stands and indicates good representation of trees in different girth classes. The
species/area curve does not show flattening out, indicating that the sample has not reached a size which
would have included most of the species.
For trees with gbh. = 24 inches the mean density of the forest is 86 stems/acre compared to 103.3
stems/acre for some stands in a similar forest type in Malaysia. The difference is found to be statistically
significant at the 0.05 level. The Mischungsquotients, varying between 1.3 and 2.0, however, compare
well with those of stands in Malaysia and indicate great complexity of species.
The hypothesis is put forward that despite the small size of the Reserve and its isolated nature the
species which are peculiar to the Coastal Hill Forest type and wielding structural and numerical
dominance as a whole, would survive over a long period of time because their progeny have an innate
ability to grow up in the shade within the ambit of the parent trees. Because of this, at any one time they
have individuals of all sizes in the forest ranging from saplings through pole-sized to mature trees. This is
a guarantee to species survival. On the other hand, species with low density which are not widespread
and whose saplings require canopy opening to grow up, may gradually die out, the reason being that the
chance opening of the canopy in their vicinity may not coincide with their fruiting, and a tree may die
before such an opening. The demise of a tree itself can cause an opening in the canopy but again it may
occur at a time when its saplings are completely absent. All these factors may lead to a decline in the
overall floristic composition of the forest.
*Mr. Wong was Forest Ecologist, Silviculturist, and later Deputy Chief Research Officer of the Forest
Research Institute, Kepong, Malaysia. He was Commissioner for Parks and Recreation between 1974
and 1982 during which term of office he also served as a Trustee of the Singapore Nature Reserves
Board.
46 Gard. Bull. Sing. 40(1) (1987)
I. Introduction
The forest on Bukit Timah Hill was made a nature reserve just before the Second
World War and is protected under the Nature Reserves Act. The area is about 75
ha. (185 ac.) and the summit of the hill at 163.5 m (536 ft.) is reputed to be the
highest point in Singapore.
Geologically the area consists of granite and because this rock type is used
extensively as a road metal and construction material several quarries have made
extensive exploitation for the rock west and south of the Reserve. The topography
ranges from gradual to steep and in some places the land falls to merge with
steep-sided gulleys where granite outcrops and boulders abound. Outcrops can also
be seen in many other places within the Reserve and along its boundary.
200m 100m _ 0 200 m 400 m
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P.U.B. Catchment Area
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\ Hamstead +. Fern
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Legend
i
Contour OO ge ee yy
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South View Hut
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23
Clusters + +3¢
Fig. 1. Sketch Map of Bukit Timah Nature Reserve Showing Location of Sampling Units.
Ecology of the Trees of Bukit Timah Nature Reserve 47
Until very recently to all intents and purposes Bukit Timah could be considered
as part of the central catchment area in Singapore, being located on its western
fringe and separated from it only by a pipe-line carrying water from Johore to
Singapore. With the recent completion of the Pan Island Expressway, east of the
Reserve, however, it is now completely isolated with non-forested areas on all
sides. The Expressway is a six-laned major highway with ample road reserve.
The vegetation consists of stands of high forest and pockets of secondary forest
along the reserve boundary at different stages of ecological succession ranging from
open scrubland to pole-sized forest. The high forest appears undisturbed and is of a
primary nature. In terms of specific composition the forest should best be classified
as a Coastal Hill Forest (vide Symington (1942)). This is characterised by Shorea
curtisii occurring at low elevation and its association with species which would
otherwise occur only or predominantly in Hill Dipterocarp Forest, such as Anisop-
tera curtisii and Artocarpus lanceifolius.
At the time when Symington wrote his Manual of Dipterocarps it was thought
that Shorea curtisti was probably endemic to the Malay Peninsula. However, more
recent taxonomic works by both Meijer (1964) and Ashton (1982) have reported
the presence of Shorea curtis in the hills of Borneo and on islands off the Sumatran
coasts. There is, however, no ecological work to elucidate its association with other
species. It is, therefore, uncertain whether there exists a Coastal Hill type of forest
in these places, similar to that under reference. If none exists then Bukit Timah is
unique in that it is the southernmost habitat of a Coastal Hill Forest.
The Hill is botanically famous because it is the type locality of many species, the
result of its being the collecting ground of many famous botanists, such as Ridley,
Burkill, Holttum and Corner. In recent years teachers and students of both univer-
sities and secondary schools have made observations on the plant communities and
animals in the Reserve (see for example, Whitmore & Wong (1959). Recently a
guide book (1985) has been published in which common plants and animals are
featured. As far as it was known before this survey was carried out no quantitative
study had ever been made on the ecology of the trees. The survey was therefore
initiated so as to create a scientific record of the tree species and their distribution
in the Reserve. The work was carried out as an activity of the Singapore Branch of
the Malayan Nature Society and the field work spanned a period of some three and
a half years between June, 1974 and January, 1978. The actual area covered by the
survey was only 3.24 ha (8.0 ac.). The apparently long period required to achieve
this lies in the fact that field work was carried out as Sunday outings, once a month,
and sometimes inclement weather rendered some Sundays impossible for work.
II. Method
(a) Design and Layout
The sampling units were located systematically. A systematic layout was prefer-
red because although the Nature Reserve is relatively small there is great variation
in topography and in the vegetation to be studied. As the study was not intended to
be a statistical exercise but one which was aimed at revealing as much as possible
the character of the forests, a random sample might not give the uniform spread of
the sampling units over the Reserve. At the time of the survey a map of the Reserve
with an Imperial* scale of 1 inch = 150 yds (approx. 1 cm = 54 m) was used as the
base map for the sampling. A one-inch grid was laid over the map; the intersections
“The survey was initiated before the Metric System was adopted in Singapore. In order to facilitate
direct comparison with results obtained in Malaysia and to avoid sometimes meaningless conversions, it
was decided to present the results of the survey using the Imperial System.
48 Gard. Bull. Sing. 40(1) (1987)
of the grid were used as the centres of the sampling units. There were some 40
possible sampling units but only 20 could be completed during the period men-
tioned earlier. A map of the Reserve with the positions of the sampling units
appears in Fig 1. It will be noted that two of the units, No.31 and No.32 were
located outside the Reserve boundary but they were included in the survey as the
forest appeared uniform and continuous with the forest within the Reserve bound-
ary. (See, however, Section III(c) concerning No. 32.)
Each sampling unit is a cluster of 4 circular plots (sub-units) each with a radius of
37.25 ft. (c. 11.35 m) arranged along the cardinal directions (see Fig 2). The plot
centre 1s 55 ft. from the cluster centre. A circular plot of 37.25 ft. radius has an area
of one tenth acre, so that a cluster covers an area of 0.4 acre. Unless otherwise
stated, the cluster forms the basis of the various analyses presented in this report.
(b) Location of the Clusters and Circular Plots
The job was made easy by the many paths and junctions present in the Reserve
reference to which cluster positions could be located by the use of a compass and a
100-ft measuring tape. In order to correct for slopes the tape was pulled as
horizontally as possible when measuring distances. If need be, such as on rather
steep slopes, measurement was done step-wise for short distances instead of for a
whole 100 feet.
Once the cluster centre was located, the tree nearest to the spot was marked and
this was used as the working centre for the cluster. The centres of the 4 circular
plots were located with reference to this tree. Likewise, trees were used to mark
their centres. If they were of enumerable size they would be enumerated; otherwise
they would be ignored during the enumeration.
N
W
Fig. 2. Configuration and Size of the Sampling Cluster. O =
Cluster Centre, Ob = 55 ft., r = radius of sub-plot =
37.25 ft., a,b,c,d, are respectively centres of sub-plots
A,B,C,D. N,S,E,W are the cardinal points. Each sub-
plot has an area of 0.1 acre. The area of the cluster is
therefore 0.4 acre.
Ecology of the Trees of Bukit Timah Nature Reserve 49
A quick inspection was made to ensure that none of the circular plots of the
cluster would fall on man-made areas, such as a foot path or boundary. If any did,
then the cluster centre was adjusted so that the plot(s) would avoid such an object
as far as possible. Other than this, cluster location adhered strictly to what was
indicated by the map.
Marking the sampling units with existing trees made our field work much easier
than if we had to carry timbers or other similar materials to site for plot or cluster
demarcation. Further, the circular plot was not demarcated. The boundary was the
circumference, within seeing distance from the centre where the team leader, doing
the booking, would be standing. When in doubt, a boundary tree was checked with
the tape to see if it was in or out of the plot.
(c) The Enumeration
Within the circular plot all trees with a girth at breast height (gbh — 4 ft. 6 ins. or
c. 1.37m from the ground) of 24 inches (c. 61 cm) or larger were measured for girth.
But for each cluster, Plot A formed a special sub-sample in which girth measure-
ment included all trees down to 12 inches (c. 30.5 cm). The idea was to use this to
indicate recruitment and regeneration. For trees which had buttresses taller than 4
ft. 6 inches, measurement was done above the buttresses and if the buttresses were
too tall to be reached, then an estimate for the girth above buttresses was made. In
the case of species which have a tendency to produce coppiced shoots from ground
level, the girth of each coppiced stem, meeting the requirement of minimum girth
at breast height, was measured.
In the enumeration the book was closed after finishing each circular plot.
The choice of 24 inches as the minimum girth in the enumeration was somewhat
arbitary. In order to get a good picture of the composition of the forest in relation
to recruitment and regeneration, the lower the girth limit the more accurate the
information. Indeed for regeneration study enumeration of the seedling population
is essential. However, it is also a fact that for rain forest ecology, the smaller the
tree the more difficult it is to identify and the present study shows that if we had
lowered the enumeration girth for all plots to 12 inches instead of 24 inches, then
we would have had to deal with a population nearly 70% larger than what we have
got. In absolute terms we would have had to measure and identify an additional 600
odd trees. This would have been too cumbersome for Sunday outings!
(d) Species Identification
For this we relied mainly on sterile characters of the species. As we were working
in a nature reserve, bark cut for identification purpose was kept to a minimum.
Using such characters one had no difficulty identifying most of the larger trees, but
for smaller trees, in many instances, we had to collect the fallen leaves and send
them to the herbarium of the Botanic Gardens for further identification. In this
study some 20 trees could not be identified at all. In the various lists presented in
this report, the class called ‘Unknown’ is the sum total of such difficult cases. (See
Section III (b) for comments and plant names and their revisions.)
(e) Compilation and Analysis of Data
The bulk of the compilation and analyses was done using dBase III Plus, on an
IBM PC, with a capacity of 640 KB and with two disk drives. dBase III Plus is a
widely known and widely used software package but for our present study it has
been found that the features mentioned below are examples of useful commands
for our compilation and analyses. Using the “‘Index”” mechanism followed by the
“Total” command, one could generate a complete species list with sample total for
SO Gard. Bull. Sing. 40(1) (1987)
each species. Indexing in association with the ‘Set Unique” command also segre-
gates species into mutually exclusive groups in different clusters. This facilitates
species/area curve construction.
The programme is also extremely useful for the calculation of basal area by
cluster with each figure being achieved by a single command! It also enables blocks
of data, for example, lengthy tables, etc. to be transferred to a word-processor for
further editing and reporting.
III. Results
(a) Some Stand Parameters
Table 1(a) presents some stand parameters of different clusters. Column 2 in the
Table gives the actual number of stems encountered in the 0.4 acre cluster. Each
figure is multiplied by 2.5 to convert it to number of stems per acre and the
converted figures appear in Column 3. Column 4 gives the number of species
present in each cluster. The Mischungsquotient (see Richards, 1964) of Column 5 is
obtained by dividing Column 2 by Column 4 and it is simply the number of
individuals per species within the stand. This is used as a measure of the degree of
complexity of the stand. The larger this quotient is, the fewer the number of species
in the particular stand.
Table 1(a). Some Stand Parameters — Bukit Timah Nature Reserve.
(Based on trees with gbh >=24 inches).
Clust. Stem No. Stem No. Species Misc. Basal
Number In The Per Acre No. In Quotient Area/Acre
Cluster Cluster
(1) (2) (3) (4) (5) (6)
5 28 70 19 Ld 125
6 42 ii 28 1.5 168
7 38 95 22 1k 177
1] 42 105 oF | 1.6 136
14 37 92 25 1.5 160
15 39 97 27 1.4 118
16 30 75 25 ie 97
Ly 32 80 20 1.6 115
18 24 60 Zp 1.0 95
19 24 62 20 be 105
23 32 80 19 Pe? 149
Za 53 130 6 8.8 96
26 33 82 1] 4.7 70
3] 39 97 24 1.6 125
32 50 125 pe) 2.0 176
33 ZT 67 21 1:3 103
34 33 82 25 1.4 114
35 23 57 19 BZ 86
36 24 60 | 3.4 76
37 37 92 14 IA) 132
* Total 687 1720 405 2423
Mean as ee 86 20.2 121.1
Ecology of the Trees of Bukit Timah Nature Reserve a1
Column 6 gives the total basal area of each cluster. The basal area of a tree is
simply its cross-sectional area at breast height and that for the stand is the sum of
the basal area of all the trees within that stand. This is a density parameter.
Table 1(b). Densities of some Malaysian Stands of Coastal Hill Forest. (Trees with
gbh>=24 ins.)
Location Elevation Plot size Plot Stems/
(ft.a.s.1) (acre) Total acre
Jarak Is. 50 — 200 0.5 40 80
200 — 350 0.5 i 102
1.0 91 91
Rumbia Is. 20 — 150 0.5 59 118
150 — 300 0.5 5S 106
Kg. Gajah 300 — 500 0.5 4] 82
Perak 350 — 400 0.5 49 98
1.0 90 90
Banang 350 — 500 0.5 50 100
Perak 700 — 850 0.5 40 80
500 — 650 0.5 54 108
250 — 350 0.5 64 128
Pangkor Is. 50 — 200 0.5 54 108
Soga, Johore 350 — 500 0.5 61 122
900 — 1100 0.5 60 120
Gunong Raya 200 — 400 Os 60 120
Langkawi Is.
Total 1653
Mean 103.3
Source: Wyatt-Smith (1963), Table 12, Pg. III — 14/20, only figures for gbh>=24 ins have been
extracted for conversion into densities presented here. All the non-island Forest Reserves, except Kg.
Gajah, are within 3 to 6 miles from the sea. Kg. Gajah is within 15 miles from the sea.
Table 1(b) presents some figures on number of stems per acre derived from
different forest reserves with Coastal Hill forest in Peninsular Malaysia, viz Jarak
Island, Rumbia Island, Pangkor Island, Banang, Kampong Gajah, Soga and
Gunong Raya on Langkawi Island. The relevant figures are extracted from Wyatt-
Smith (1963).
The purpose is to compare this parameter with the figures presented in column 3
of Table 1(a).
The density figures of the Bukit Timah Forest show rather wide variation, with
the values ranging from 57 to 130 stems per acre. The mean of the 20 clusters is 86
stems per acre. A t-Test, dealing with unpaired variates but with the variance of the
two samples being equal, shows that this mean is significantly different from that of
103.3, the mean derived from the 16 stands of forests in Peninsular Malaysia, with
the significance just missing the 1 per cent level. The Bukit Timah clusters have
included 4 stands known to have disturbed or man made forest, viz. Clusters 25, 26,
36 and 37. Even if we exclude these from the t-Test, the conclusion is still the same.
32 Gard. Bull. Sing. 40(1) (1987)
Table 2. Stands of Disturbed or Man-made Forests. (All trees with gbh>=24
inches. See girth size equivalents in Section III(d)).
(a) Cluster 25 with Adinandra dumosa as the dominant species
Species Gf ~ G2-= G3 “G4 Ga ae Total
1. Adinandra dumosa 0 0 30 6 p! 0 38
2. Campnosperma auriculatum 0 0 1 0 0 0 1
3. Cratoxylon formosum 0 0 1 0 0 0 1
4. Gironniera parvifolia 0 0 0 1 0 0 1
5. Ixonanthes reticulata 0 0 1 1 2 1 5
6. Rhodamnia cinerea 0 0 4 1 0 0 5
(R. trinervia)
7. Timonius wallichianus 0 0 2 0 0 0 2
(b) Cluster 26 also with Adinandra dumosa as the dominant species
Species Gl G2. G3” 4 ee Total
-1. Adinandra dumosa 0 0 18 a 0 0 21
2. Cratoxylon formosum 0 0 0 0 0 1 1
3. Eugenia linocieroidea 0 0 1 0 0 0 1
4. Ixonanthes reticulata 0 0 0 0 2 a Ss
5. Ploiarium alternifolium 0 0 1 0 0 0 1
6. Rhodamnia cinerea 0 0 3 0 0 0 3
(R. trinervia)
7. Streblus elongatus 0 0 0 0 1 0 1
(Sloetia elongata)
(c) Cluster 36 with Palaquium gutta as the dominant species
Species Gh. 3G2 GS!» -Ge eee Total
1. Actinodaphne sesquipedalis 0 0 1 0 0 0 i
2. Artocarpus scortechinii 0 0 1 0 0 0 1
3. Litsea elliptica 0 0 1 0 0 0 1
4. Palaquium gutta 0 0 4 6 5 e) 18
5. Pellacalyx saccardianus 0 0 0 0 1 0 1
6. Pimelodendron griffithianum 0 0 1 0 0 0 1
7. Shorea leprosula 0 0 0 0 1 0 1
(d) Cluster 37 also with Palaquium gutta as the dominant species
Species Gt. dad, GE!) Gas ie Total
1. Adinandra dumosa 0 0 1 0 0 0 1
2. Antidesma coriaceum 0 0 0 1 0 0 1
3. Campnosperma auriculatum 0 0 0 1 0 1 7:
4. Cratoxylon formosum 0 0 0 1 0 0 1
5. Dysoxylon sp. 0 0 1 0 0 0 1
6. Eugenia linocieroidea 0 0 + 0 0 0 4
7. Gymnacranthera forbesii 0 0 1 0 0 0 1
8. Gynotroches axillaris 0 0 0 1 0 0 1
9. Ixonanthes reticulata 0 0 1 1 0 1 3
10. Litsea elliptica 0 0 0 1 0 0 1
11. Gluta wallichii 0 0 0 1 0 0 1
(Melanorrhoea woodsiana)
12. Palaquium gutta 0 0 7 7 1 3 18
13. Rhodamnia cinerea 0 0 1 0 0 0 1
(R. trinervia)
14. Sindora sp. 0 0 0 0 0 0 1
Ecology of the Trees of Bukit Timah Nature Reserve 53
It is, however, difficult to proffer an explanation as to why there should be a
significant difference. One is tempted to surmise the forest at Bukit Timah is not
that primary after all and that past creaming of the forest for small timbers by
villagers living in the vicinities could have reduced the density.
Coming to the Mischungsquotients in Column 5 of Table 1(a), if we leave out the
ones for clusters 25, 26, 36 and 37 for the time being, then we have them ranging
from 1.0 to 2.0. If the quotient is 1 then it is the same as saying that within the
particular stand every individual belongs to a different species! If it is 2 then we
would expect to find two individuals to every species. From the figures we have it
can be seen that the Bukit Timah high forest is very mixed and this is comparable to
stands found in different parts of Peninsular Malaysia. Figures given by Wyatt-
Smith (1963) for Pangkor F.R. Banang F.R., Bukit Lagong F.R. and Kampong
Gajah F.R. also range from 1.3 to 1.9.
The quotients for Clusters 25, 26, 36 and 37 are certainly larger than that of the
other clusters. As stated earlier large values indicate stands dominated by a few
species. Reference to Table 2 will show that this is the case. Clusters 25 and 26 are
dominated by a single species, Adinandra dumosa. These two stands are in dis-
turbed forest lying at the fringe of the reserve and undergoing succession. Clusters
36 and 37 are dominated by Palaquium gutta, a species of commercial importance
in the past because it yielded gutta percha, the raw material for chewing gum.
Corner (1952) made passing reference to the trees planted in the Taban Valley and
Clusters 36 and 37 were located therein.
In rain forest ecology one of the rather intriguing questions is how big a sample
one must take in order to include most of the species in an area under investigation.
One method employed to get some idea is the Species/Area Curve. Greig-Smith
(1964) states that there are commonly three ways in which to construct such a
curve, the most efficient being to take independent samples of different sizes within
the study area and then plot number of species encountered against area of the
respective sample. In this study of ours it was realised from the start that our
sample size would never be able to exhaust all the tree species in Bukit Timah,
bearing in mind that it has been variously estimated that in Peninsular Malaysia
alone there are no less than 3,000 species of trees with 12 inches girth and larger.
However, as a look-see exercise a plot is made here (Fig. 3) of species against area,
the area of successive classes being cumulative upon the previous total, a method
described by Greig-Smith (ibid) as being most inefficient.
Initial attempts to construct the curve included all 20 clusters and it produced a
curve that seemed to rise in three steps. This was due to the fact that the four
disturbed/man-made stands (see Table 2) mentioned previously, had few species
and when the curve hit those stands, it flattened out, only to rise again when it hit
again normal stands with more species. As it would not be logical to have a curve
covering different known forest types, with obviously different specific composi-
tion, the four abnormal stands were taken out and another curve constructed. The
resultant curve is shown in Fig. 3. It is noted that it is a relatively smooth curve with
the initial portion (within the first 0.8 acre) rising steeply as expected; it then
assumes a slightly more gradual slope but right till the end it is still on the rise,
showing that our sample of 6.4 acres is far from having included most of the species.
(b) Specific Composition of the Forest
A complete list of species by families arranged in alphabetical order is given in
Appendix 1(a). The ‘‘unknown”’ plants are those which we could not identify at all
and these are distributed in 13 of the clusters.
Quantitative plant ecological work in the Region has been in progress in the past
few decades, during which taxonomic revisions of the various families or genera,
54 Gard. Bull. Sing. 40(1) (1987)
Number of trees
Area (Acres)
Fig. 3. Species/Area Curve. All trees with gbh>=24 inches.
notably by the Flora Malesiana Foundation, have also taken place. This has
rendered some of the familiar and much used Latin names obsolete. In this study,
not only have the identities of the plants been checked by the Singapore Botanic
Gardens Herbarium, various relevant works have also been consulted to ensure the
use of the latest Latin names. However, in order to enable foresters and plant
ecologists of the Region, who might not have up-to-date knowledge of the plant
names but are nevertheless familiar with old names, to get a quick mental compari-
son on what they read, I have wherever possible included such names in parenth-
eses after the latest Latin names. This is done in both text and tables.
Appendix 1(b) gives a summary of the number of species and genera by family,
with the families arranged in order of species preponderence. There are 44 known
families and the totals show that the survey encountered 111 genera, 212 species
and 889 individual trees. Apart from the 20 ““unknown”’ trees there are another 20
trees in 11 genera which could be identified down to genus level only. That makes
849 trees which have been fully identified.
It can be seen that Euphorbiaceae has the highest number of genera (11) and the
highest number of species (22). In terms of species preponderence the Burseraceae,
Dipterocarpaceae, Myrtaceae and Lauraceae all tie second place with 13 species
each! Next come Leguminosae with 11 species and Annonaceae with 10 species,
and so on.
Ecology of the Trees of Bukit Timah Nature Reserve 2
In terms of sample total the Dipterocarpaceae, with 125 individuals, tops the list.
There is an anomaly in that the Theaceae takes second place with a sample total of
94 individuals. The species responsible for this is Adinandra dumosa, coming
mainly from two clusters located in disturbed forest, namely clusters 25 and 26 (see
Table 2). Surprisingly, the Sapotaceae also occupies a position of eminence, 5th in
the list. This is due in the main to the fact that clusters 36 and 37 were located in a
stand of planted Palaquium gutta. If we exclude these 4 stands from the analysis,
then the two families in question would go further down in the list.
Barring the anomaly created by the 4 stands just referred to, one could say that
the specific composition as encountered in Bukit Timah in the undisturbed stands is
quite typical of the Lowland Dipterocarp Forest, of which the Coastal Hill Forest
may be considered as a sub-type. As mentioned earlier, one of the outstanding
features of the Coastal Hill Forest is the presence of Shorea curtisii (Seraya) at low
elevation. In Bukit Timah its occurrence could be as low as 150 ft a.s.1. Associated
with Seraya are usually such species as Shorea gratissima, Shorea glauca, Dipter-
-ocarpus caudatus ssp. penangianus (D. penangianus) Anisoptera curtisii, Artocar-
pus lanceifolius, Swintonia schwenkii or S. spicifera. With the exception of Shorea
glauca these species are found within Bukit Timah but Anisoptera curtisii and
Swintonia have not been netted in by our sample.
The dipterocarps have in this study shown family dominance, a fact which is quite
well known from various studies made in Malaysia. In the Bukit Timah case this
dominance is very much the result of two dipterocarp species, viz Shorea curtisii,
with a total of 46 individuals, and Dipterocarpus caudatus ssp. penangianus (D.
penangianus) with 37 individuals. These two dipterocarps alone have contributed
83 individuals out of a total of 125 (vide Appendix 1(a) & the end of Appendix 4).
Looking at Appendix 1(b) if we remove the two clusters located in the secondary
forest dominated by Adinandra dumosa, then we would have the Euphorbiaceae
taking the second place with a total of 70 individuals. It also has a preponderance of
genera and species — with respectively 11 and 22 numbers. In terms of species, the
genera Aporusa and Baccaurea contribute most to the total number. Baccaurea
kunstleri and Pimelodendron griffithianum seem to be quite common and wide-
spread. Unlike the dipterocarps, however, trees of the Euphorbiaceae are small.
Even the larger ones have their crowns in the Main Storey of the canopy and many
of them stay in the C storey. In our sample, probably the largest trees are
Endospermum diadenum (E. malaccense) Blumeodendron tokbrai and Baccaurea
kunstleri, all growing up to about 70 ft.
The Anacardiaceae follows closely behind the Euphorbiaceae in the preponder-
ance of individuals with a sample total of 58, coming mainly from two species, viz
Gluta wallichii (Melanorrhoea woodsiana) and Campnosperma auriculatum, re-
spectively with 34 and 16 individuals. In Bukit Timah, it is somewhat surprising that
the genus Mangifera is not as common as it should be. Mangifera lagenifera, for
example, just like Swintonia schwenkii, is seen sporadically only in the Reserve, but
our sample has not been able to net it in.
Campnosperma auriculatum is an interesting species in that it has a very wide
ecological amplitude. It occurs in both primary and secondary forests, in the
lowlands, in mountains and in the peat and fresh water swamps! Such a feat seems
to be equalled only by Koompassia malaccensis(Leguminosae). In Bukit Timah, as
in other places with high forest nearby C. auriculatum is one of the first pioneers to
colonise secondary vegetation and in the primary jungles it can grow to a large
emergent.
Gluta wallichii (Melanorrhoea woodsiana), apart from being abundant, is also
the most wide spread species in Bukit Timah as the Distribution Index (see next
56 Gard. Bull. Sing. 40(1) (1987)
Section) will show, much to the woe of those who are allergic to its potent skin
irritant! Gluta wallichii, unlike Campnosperma, is not a very large tree. Even when
full grown its crown would be in the Main Storey of the canopy.
For more information on the relative importance of the various families and
species in our sample, the reader is referred to the various parts of Appendix 1.
(c) Spatial Distribution of Species and the Distribution Index (DI)
From the Mischungsquotient mentioned earlier, it can be seen that the Bukit
Timah forest, just like other Rain Forests in the Region, has a great diversity of
species. Yet there can be no doubt that some species are of very low density while
others show considerable dominance in numbers. Some species are vagrant while
others tend to be more gregarious. In such a situation tabulation to show the
presence of a species would not be too meaningful if we give only its sample total
for it could mean that the individuals were present in only one or two sampling
units, or they might be spread over many units. Because of this it is thought much
more useful to present a comprehensive list of the plants showing both their sample
totals as well as the extent to which they spread over the entire sample.
To show the latter phenomenon a parameter called the Distribution Index (DI) is
used. The DI of a species is simply the number of sampling units in which it occurs.
Thus, for example, in Appendix 1(a) it is seen that Dipterocarpus caudatus ssp.
penangianus (D. penangianus) has a DI of 12. It simply means that in the whole
sample, it occurs in 12 of the 20 clusters.
It has to be noted that it does not matter how many individuals occur in a cluster.
So long as there is one individual of the species, the species scores 1 for that
particular sampling unit. On the other hand even if there are 10 individuals in the
unit the score for DI is still 1. This brings us to the other point which is important
140
Number of species
Di Classes
Fig. 4. Species Distribution & Distribution Index (DI). (See Section III(c) of
text.)
Ecology of the Trees of Bukit Timah Nature Reserve a7
for showing dominance, i.e. the sample total of a particular species. Taking D.
caudatus ssp. penangianus again as an example, we find that its sample total from
all the clusters adds up to 37. This is indeed a very high number compared with
many other species. This, coupled with the fact that it is distributed in 12 of the 20
clusters, shows that it is wielding considerable influence within the community we
have sampled.
Appendix 1(a) shows the DI amongst other things but much more information
could be got at a glance by referring to Appendix 2. In here both sample total and
the DI of a species are given with the DI indexed in ascending order. From here it
can be seen that more than half of the species occur only once whereas others may
occur in one cluster only but within that cluster it occurs 5 times. Such is the case
with for example Aporusa bracteosa (Euphorbiaceae). Further, it can be seen that
as DI increases the number of species drops rapidly (see Fig. 4). There are 121
species with DI=1 but when DI 7 is reached, there are only 3! These are Pime-
lodendron griffithianum, Streblus elongatus (Sloetia elongata) and Timonius wal-
lichianus. Of the known species only 5 score a DI equal to or greater than 10. They
are Pellacalyx saccardianus (10), Artocarpus lanceifolius (11), Dipterocarpus
caudatus ssp. penangianus (12), Shorea curtisii (12) and Gluta wallichii (Melanor-
rhoea woodsiana) (15).
Species which have a DI of unity and small total, say 1 or 2 are likely to be
isolated vagrants while those with a high DI and high total are the really common
species of the community. A third condition can be that a species may have a low
DI but a very high total. Such a case indicates an extremely gregarious condition
and this may reflect the successional status of the particular community. Browsing
through Appendix 2 one could discern such a condition for Campnosperma auricu-
latum (TOT 16, DI 3) Rhodamnia cinerea (R. trinervia) (TOT 16, DI 3) Adinandra
dumosa (TOT 90, DI 4) Palaquium gutta (TOT 38, DI 4). In the case of Campnos-
perma, the master list (not included in this report) reveals that most of the
individuals (some 11 numbers) come from Cluster 32 alone. Inspection of the list of
species for the cluster shows that the associated species within that particular
cluster are matured individuals of high forest species. But a closer scrutiny reveals
that some 8 trees occur in Plot B alone within that cluster. It is therefore certain
that Cluster 32 had been placed straddling the boundary between high and regener-
ating (disturbed) forest.
In the case of Rhodamnia and Adinandra most of the individuals have come from
Clusters 25 and 26, which are located in disturbed vegetation. Palaquium gutta, as
indicated previously, comes mainly from the historical planted patch within the
Taban Valley.
(d) Structure of the Forest and Recruitment
For forest structure we shall use girth class distribution as an indicator. Appendix
2, inter alia, gives the girth class distribution of the various species and individuals
encountered, from G1 to G6. The equivalents in inches are given below:
Gl G2 G3 G4 G5 G6
12-12.9 13-23.9 24-35.9 36-47.9 48-59.9 >=60.0
As mentioned in the notes on enumeration, G1 and G2 were only measured in
the sub-sample of Plot A in each cluster. Appendix 2 has included all individuals
down to Gl.
Girth classes in a gigantic table are difficult to visualise. On account of this a
graph is produced by plotting number of trees against girth classes. Before doing so
58 ; Gard. Bull. Sing. 40(1) (1987)
the data were re-arranged so that the girth classes extend from 12 inches gbh to
over 144 inches gbh. As the 12-24 inch girth class was measured only in one of four
sub-plots, the number obtained was extrapolated by multiplying by 4 so as to bring
it in line with the other girth classes. The value of Girth Class 1 in Fig. 5 (796) has
been obtained in this way.
The graph in Fig. 5 shows a typical inverted “J” for girth class distribution of
individuals in a stand of uneven-aged forest. The characteristics are a very high
number of small trees and a low number of very large trees.
The graph suggests that the forest as a whole is well stocked with young trees to
take the place of the big giants when they die. This interpretation is likely to be an
over-simplification. As is already known the forest is extremely mixed. The ecosy-
stem is likely to be controlled by a fairly large number of species. For the present
forest to beget a similar forest in the future, it is necessary to have species that form
a large proportion of the present composition and which have vertical distribution
900 —O—O@—O0— _ Alltreesinsample
—~-K-— —xX— —K— — Excluding trees of
Clusters 25, 26, 36 & 37
800
700
600 All Clusters
Girth No. of
Class Inches Trees
1. 12— 23.9 796
p. 24 — 35.9 311
Z 3: 36 — 47.9 169
® 500 +1 4. 48 — 59.9 75
= 5) 60 — 71.9 48
° 6. 72— 83.9 35
¢ The 84— 95.9 22
E 8. 96 — 107.9 18
rad 400 8, 108 — 119.9 6
10. 120 — 131.9 4
Ast 132 — 143.9 0
2 > = 144.0 2
Girth Classes
Fig. 5. Frequency Distribution in Different Girth Classes.
Ecology of the Trees of Bukit Timah Nature Reserve 59
over the whole range of canopy strata. The data presented in the various sections in
Appendix 3 are meant to afford a glimpse into this problem.
Each section presents the results of a command put to the computer to list the
species with a definite girth class distribution pattern. In App. 3:a it was asked to
display species which are represented in all girth classes. Only one species, viz
Streblus elongatus (Sloetia elongata) met the condition. This species is likely to
survive well under even adverse conditions. Not only does it have a high total, it
also has a fairly high DI of 7, showing that it is fairly widely distributed. Indeed it is
common knowledge that this species occurs in all kinds of vegetation and is one of
the pioneers in regenerating forests.
App. 3:b lists species which have a continuous girth class distribution from G2
through to G6. For this 9 species meet the condition, including Streblus (Sloetia). It
is interesting to note that practically all the species are high forest species and they
are components of the different canopy strata — Shorea curtisii, Dipterocarpus
caudatus ssp. penangianus and Campnosperma auriculatum are emergents; Arto-
carpus lanceifolius, Gluta wallichii (Melanorrhoea woodsiana) and Artocarpus rigi-
dus are Main Storey species while the others are C-storey or Main Storey species.
Moreover, for all of them G2+G3 are very strongly represented. These are trees
with gbh from 13 inches to 36 inches. These are most likely to grow to maturity to
take over from the present giants. There is yet another plus factor for this group of
species, for it may be recalled that G1 & G2 were obtained using only 1 circular
plot. Conceivably their numbers are much larger if we had sampled down to G1 in
all the circular plots.
The total number of individuals in this group of trees number 241. Since Pala-
quium gutta comes from the planted plot we exclude its individuals. With this, 203
trees remain. The grand total for the whole sample is 889 trees. Taking proportion
we can see that these few species contribute to over 22% of the total population.
Not only that, with the exception of Palaquium and Campnosperma, they also have
high DI’s, indicating that they are widespread. They therefore exert considerable
structural and other ecological influence on the community.
For App. 3:d the display is of species with a continuous girth class distribution
from G4 to G6 and for this two more dipterocarps which are emergents and several
more Main Storey species are included. Here we can discern species, Shorea
macroptera and Palaquium hexandrum, which do not have small trees and we are
apt to jump to the conclusion that they therefore lack a vital factor to back up their
succession. However, this may not be necessarily so and this question is discussed
further in the next Section.
IV. Discussion
The present study has confirmed that the forest at Bukit Timah Nature Reserve
should best be considered as a Coastal Hill Forest according to Symington’s
classification (vide 1942, ibid) and as far as one could see a large proportion of the
forest is still of a primary nature. However, although there was no organised
commercial logging of the forest for timber, the significantly lower density of the
forest compared with some stands in such a forest type in West Malaysia, gives rise
to speculation that removal of small timbers by villagers living nearby could have
taken place in the past. The average density of the forest at Bukit Timah is some
16% less than that of the Malaysian stands quoted in Table 1(b).
The forest at Bukit Timah is indeed rich dipterocarp forest with large trees of
Shorea curtisii (Seraya) Dipterocarpus caudatus ssp. penangianus (Keruing) and
many other trees of dipterocarps and non-dipterocarps which loggers would sali-
vate over were they to see them in their concession area! Yet these giants seemed to
60 Gard. Bull. Sing. 40(1) (1987)
have escaped the saw. To hazard a guess one would say that one reason for their
having done so lies perhaps in their not belonging to the groups of naturally durable
heavy hardwood which loggers in the early days of timber utilisation sought after.
Most of the large and common trees in Bukit Timah would yield what the wood
technologist would classify as light or medium hardwood which found extensive
uses only after the War on account of advancement in wood preservation. By that
time, fortunately, Bukit Timah was already a nature reserve.
According to Symington, Coastal Hill Forest communities should have quite a
number of Balaus (the Heavy Hardwood dipterocarps) such as Shorea glauca and
S. maxwelliana. These, however, have not been met with during our enumeration
nor did the writer see any outside the sample. Could their absence be due to past
creaming? This is difficult to say but is unlikely for no sign of stumps could be seen
nor would the very natural and undisturbed nature of the environment suggest past
logging activities. Further, if such species had been in the forest before then
although selective logging would have removed the large trees, their progeny could
have survived in the forest and by now would have grown to sizeable individuals.
Their absence therefore suggests that the forest is inherently lacking in such
species.
_ Tropical Rain Forests are noted for their diversity of species. The present study
shows that the forest at Bukit Timah is no exception. Indeed with Mischungsquo-
tients ranging between 1.0 and 2.0 the forest is as diverse as any in Malaysia.
Furthermore the stand curve derived has shown that the forest also has girth class
distribution typical of uneven-aged stands. The characteristic inverted “‘J”’ indicates
the presence of a very large number of small trees, quite numerous larger trees and
a small number of very large trees.
Despite this, however, one of the nagging questions concerning a small area of
forest like Bukit Timah (185 acres) which is completely isolated, is whether it could
survive with unchanged character over a long period of time. The analysis in the
previous Section has already touched briefly on this subject. Based on the evidence
available it would appear quite a number of the high forest species show good
recruitment and are well represented in the whole range of girth classes studied. On
the other hand many of the giants have big trees only and are completely lacking in
smaller progeny. One is apt to jump to the simple conclusion that those with
smaller trees in say G1 to G3 are likely to survive while those without such small
trees will be in danger of dying out. In actual fact the situation is likely to be much
more complex and for a more complete picture, study of the seedling populations
and of trees of different size classes of various species, under a continuous inven-
tory with say annual reenumeration to assess changes in the populations, due to
normal mortality and recruitment or to catastrophes, with time would be necessary.
This of course would entail the laying down of permanent sampling units and
labelling of the plants permanently.
The optimist may argue that the forest at Bukit Timah has been there for
millennia and it has survived unchanged up to now despite its progressive isolation
into a small plot of land in recent times. It is certainly true that based on its present
composition, we can conclude that the forest is still similar to any other Coastal Hill
Forest. Also even if it is true that species of emergents have no pole-sized trees at
any one point in time that does not mean that there would not be such trees at a
later date. The seedlings of Shorea curtisii (Seraya) are known to be able to grow up
under their parents in rather shaded conditions and this may account for its
superiority in numbers and continuous girth class distribution over a wide range of
girth sizes (vide also Wong (1978)). On the other hand some species are known to
have a truncated girth class distribution in nature, over a sizeable area of forest.
Some of the emergents, e.g. Shorea leprosula, though needing forest conditions for
Ecology of the Trees of Bukit Timah Nature Reserve 61
its saplings to grow up, are known to be light demanders and they need an opening
in the canopy such as one resulting from the parent tree having been killed by a
natural catastrophe, to give them the additional light and or reduction in root
competition for growing up. If no disaster occurs over the area of forest in question
to create the gaps, for a long, long time, then only large trees will appear in a
sample. Poore (1968) has found in a lowland forest of 1 square kilometer in Jengka
F.R. in Pahang, Malaysia, that 8 — 10% of the forest was under regeneration in
some gaps. Further, he estimated that such gaps would take about 30 years to
mature, i.e. to produce uniform crown conditions again to merge with the adjacent
trees, and that such regenerated areas could last for a mean period of some 270
years! If this is so then uniform shaded conditions would prevail over this period
and it would preclude the possibility of light demanding seedlings/saplings from
growing up. It is little wonder therefore such species have a truncated girth class
distribution.
The example just considered relates to a case of the habitat being taken over by
the progeny of a tree formerly occupying it. However, this need not be always the
case. Indeed it is doubtful when a forest dies it would be replaced by its
replica. The demise of a big tree or group of trees could create conditions favour-
able for the progeny of nearby species to develop in the gap so created. It is entirely
a matter of chance as to what would seed up, or grow up in, such a gap apart from
saplings of the trees formerly occupying the site, as they might be there. Other
factors being equal proximity of species to the gaps and an inherently more
frequent fruiting habit, would definitely be an advantage. Thereafter the ensuing
fierce competition will also determine what would survive and grow up to maturity.
It is most probable that such chanced reproduction of species in regeneration
gaps in large ecosystems like the Lowland Dipterocarp Forest and the Peat Swamp
Forest, is one way of ensuring species survival and hence species diversity. Enormi-
ty of area would ensure superiority in numbers for practically all species and hence
a good chance for all to regenerate and survive. Thus although the composition of
the forest and the pattern of species distribution may change from generation to
generation, the same pool of species will remain practically unchanged.
The same mode of regeneration has been observed to take place in Bukit Timah
but in a small isolated community it is likely that the innate ability of the species to
regenerate and grow up in shade would be important for perpetuation. Perhaps it is
no coincidence that the few species shown in Appendix 3: b have individuals in the
whole range of girth class distribution and are numerically dominant as a group.
Indeed they are the very species (perhaps with the exception of Campnosperma)
that possess the innate quality of being able to reproduce and grow up in the shade
and in the ambit of the parent trees.
Shorea curtisii, Anisoptera curtisii, Artocarpus lanceifolius are essentially Hill
Forest species, in inland mountains occurring from about 1000 ft. to 2500 ft. a.s.1. in
Malaysia. They miss the vast areas of Lowland Dipterocarp Forest only to emerge
again in the Coastal Hills. Coastal Hills in contrast with inland mountain masses are
always small. In this respect I would say that the forest at Bukit Timah, at least the
typical Coastal Hill species viz Shorea curtisii, Dipterocarpus caudatus ssp. penan-
glanus, Anisoptera curtisii and to some extent Artocarpus lanceifolius, have always
faced isolation, even before the advent of Man as they could grow only in the
Coastal Hill habitat in Singapore. Presumably the other species which are common
to both Lowland Forest and the Coastal Hill Forest could have moved to and fro in
geological time, thus contributing to and ensuring the complexity of their kinds; but
not the few species just mentioned as they could only grow in the Hills. Granted
that places like Mt Faber, Bukit Gombak and some of the hills in the Pasir Laba
area, could have carried Coastal Hill species, knowing their characteristics they
62 Gard. Bull. Sing. 40(1) (1987)
could not have migrated to and fro to maintain their dominance in these places. In
other words these species must have persisted as isolated colonies, and their innate
ability to regenerate and grow up in the shade has ensured their survival and
dominance.
Looked upon in this light it is likely that these species that wield structural
dominance and to some extent numerical dominance (when considered as a group)
would be able to persist in perpetuity, provided of course Bukit Timah is preserved.
However, the physical isolation resulting from recent road and other development
could induce more drastic fluctuations in the critically important ecological factors
such as temperature and humidity. These may affect the periphery more but by and
large they are not likely to have much effect on the interior of the Reserve bearing
in mind that we are in the heart of the Humid Tropics. One thing though we must
guard against and that is the danger of fire. During a very dry year the scrubland on
the periphery may catch fire and this may cause destruction to the dried up interior
of the forest. Such an event actually happened to Cape Rachado, another Coastal
Hill Forest situated about 10 miles south of Port Dickson. The forest was badly
burnt when it caught fire during the unusually dry spell of 1963. Many of the giant
shoreas were killed and the place was invaded by Trema spp. For the other species
on Bukit Timah which are common to both the Lowland Forest and the Coastal
Hill Forest, particularly those with low frequencies and small DI’s, the diminutive
state of the Reserve does give cause for concern. For such species, the timing of a
seed year or the presence of seedlings or small trees and the chance demise of the
parent tree is very critical. This is particularly so when it is realised that most of the
large trees in the Rain Forest have evolved with long fruiting intervals of 2, 3 and as
long as, 7 years. For a small, isolated forest like Bukit Timah the scenario as
depicted below could happen.
Let us say a Dyera costulata, the Jelutong, (of which there are only a few
individuals in the Reserve) has no seedlings on the ground now. Neither are there
pole-sized trees around. It is killed by lightning. That could be the end of that tree
and species in that particular spot. Since the Reserve is small and since the
population of such a species is finite it is conceivable that after a few more of such
disasters, the species is extinct within the Reserve. This is the kind of mechanism
that could in the long run lead to retrogressive changes in the specific composition
of the forest, resulting in the decline in the number of species. Unlike the geological
past, before the advent of Man, there shall not be Jelutong trees near Bukit Timah
to effect migration back to it!
To conclude I think we can say that Bukit Timah is unique in that at the fringe of
a great metropolis lies the relict of a small Rain Forest community completely
isolated by non-forested areas. Ecologically it should be extremely interesting to
study. Scientific records obtained under a continuous inventory over the next
decade or two should yield preliminary information on changes in the community
with its component small animals one way or another. Would it be able to survive
with no loss in characteristics or would it degenerate into a secondary forest such as
many experts have predicted. Only time and some hard work will tell.
Acknowledgement
As mentioned in the introduction the field work of the project was carried out as
an activity of the Singapore Branch of the Malayan Nature Society. At the time the
Secretary was Mrs Lisette Henrey. She and her husband were indeed staunch
supporters. This could be. seen from the fact that Mr. Henrey had on many
occasions come with their toddler kid tied to his back Chinese style to help in the
enumeration! Other regular supporters were Prof. A.N. Rao of the Botany Depart-
ment, University of Singapore, and his colleagues, notably Prof. Hsuan Keng, and
Ecology of the Trees of Bukit Timah Nature Reserve 63
Dr. K.H. Chow, my then colleagues Lee Sing Kong & Teoh Teck Seng, and
students of the Ornamental School of Horticulture, Botanic Gardens. Occasionally
we were also joined by some other members of the Malayan Nature Society of
whom it will be too numerous to mention by names and to whom I should like to
express my grateful thanks for having helped in the field work.
I am also very much indebted to Mohd. Shah of the Botanic Gardens for having
assisted in the identification of materials in the herbarium. If not for his painstaking
work the magnitude of the ““Unknown” in our plant lists would have been much
larger. He and Ali Ibrahim also helped to up-date the Latin names of many of the
species and the authorities that go with them.
References |
Symington, C.F. (1942). Foresters’ manual of dipterocarps. Mal. Forest Rec. No.
16 — chart on pg XIV & Text Fig. 2. Caxton Press, Kuala Lumpur.
Meijer, W. & Wood, G.HLS. (1964). Dipterocarps of Sabah. Sabah Forest Record
No. 5. Forest Department, Sandakan.
Ashton, P.S. (1982). Dipterocarpaceae. Flora Malesiana Seri I Vol. Doe Pt-2.
Whitmore, T.C. & Wong, Y.K. (1959). Sunfleck and shade light in Tropical Rain
Forest. Mal. Forester, Vol. XXII, pg 50.
A Guide to Bukit Timah Nature Reserve. Published by Singapore Science Centre
in collaboration with the Nature Reserves Board. First published 1985.
Richards, P.W. (1964). The tropical rain forest. Cambridge Unviersity Press.
Wyatt-Smith, J. (1963). Manual of Malayan silviculture for inland forests. Mal.
For. Rec. No. 23. Vol. 2, pg III-14/14 et seq.
Corner, E.J.H. (1952). Wayside trees of Malaya. Govt. Printing Office, Singapore.
Pg 601.
Greig-Smith, P. (1964). Quantitative plant ecology. Butterworths, London. Pg 151
et seq.
Wong Y.K. (1978). Girth class distribution of Seraya (Shorea curtisii) as an
indication of its regeneration and recruitment in Bukit Timah Forest Reserve.
Mal. Nat. Journ. Vol. 31, pg 149-153.
Poore, M.E.D. (1968). Studies in Malaysia Rain Forest. I. The forest on Triassic
Sediments in Jengka Forest Reserve. J. Ecol. Vol. 56, pg 143 et seq.
64
Gard. Bull. Sing. 40(1) (1987)
Appendix 1: a. Complete list of species (gbh>=12”) by family, with the species’
sample totals and their distribution indices (DI).
(For explanation on DI see text Sect. III (c). Latin names in parentheses are obsolete but familiar
Rec. #
We WN
20%
30.
40).
Family
ANACAR
ANACAR
ANACAR
ANACAR
ANACAR
ANACAR
ANONA
ANONA
ANONA
ANONA
ANONA
ANONA
ANONA
ANONA
ANONA
ANONA
APOCYN
APOCYN
BOMBAC
BURSER
BURSER
BURSER
BURSER
BURSER
BURSER
BURSER
BURSER
BURSER
BURSER
BURSER
BURSER
BURSER
CELAST
CELAST
CONNAR
CONNAR
DILLEN
DIPTER
DIPTER
DIPTER
DIPTER
DIPTER
names.)
Species
Bouea macrophylla Griff.
Bouea oppositifolia (Roxb.) Meissn.
Buchanania sessilifolia BI.
Campnosperma auriculatum (BI.) Hk. f.
Gluta wallichi (Hk. f.) Ding Hou
(Melanorrhoea woodsiana King)
Parishia paucijuga Engl.
Anaxagorea javanica Bl.
Cyathocalyx ramuliflorus
(Maing. ex Hk. f & Th.) Scheff.
Cyathocalyx ridleyi (King) Sinclair
Fissistigma lanuginosum Hk. f. et Th.
Fissistigma latifoltum (Duval) Merr.
Polyalthia hookeriana King
Polyalthia rumphii (BI.) Merr.
Polyalthia sp.
Xylopia ferruginea
(Hk. fet Th.) Ak: fet Th.
Xylopia malayana Hk. f. et Th.
Alstonia angustifolia Wall. ex A.DC.
Dyera costulata (Miq.) Hk. f.
Durio griffithii (Mast.) Bakh.
Canarium grandifolium (Ridl.) H.J. Lam
Canarium sp.
Dacryodes costata (Benn.) H.J. Lam
Dacryodes laxa var typica (Benn.) H.J. Lam
Dacryodes rostrata (Bl.) H.J. Lam
Dacryodes rugosa (Bl.) H.J. Lam
Santiria apiculata Benn.
Santiria griffithii (Hk. f.) Engl.
Santiria laevigata BI.
Santiria rubiginosa BI.
Santiria tomentosa BI.
Santiria sp. A
Santiria sp. B
Bhesa paniculata Arn.
Kokoona reflexa (Laws.) Ding Hou
(Lophopetalum reflexum Laws.)
Agelaea borneensis (Hk. f.) Merr.
Ellipanthus tomentosus kurz
Dillenia grandifolia Wall. ex Hk. f. et Th.
(Dillenia eximia Miq.)
Anisoptera costata Korth.
Dipterocarpus kerrii king
Dipterocarpus caudatus
ssp. penangianus (Foxw.) Ashton
(Dipterocarpus penangianus Foxw. )
Dipterocarpus sublamellatus Foxw.
Hopea mengarawan Miq.
Total
wore
oe le Ln N —=
FS MO NOR kK Ne
KS Be KSB WN WON NK NN NN HR KK KK LY
DI
—
Ch GN =i
Nr ££
KS NOK KS KN B CO COR KN NR RRR RK ke LY
—
.
:
Ecology oj the Lrees Of Buku Liman Nature Keserve
(Appendix 1:a cont.)
Rec.#
43.
44.
45.
46.
90.
Family
DIPTER
DIPTER
DIPTER
DIPTER
DIPTER
DIPTER
DIPTER
DIPTER
EBENA
EBENA
EBENA
EBENA
ELAEOC
ELAEOC
ELAEOC
ELAEOC
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
EUPHOR
FAGA
FAGA
FAGA
FAGA
FAGA
FAGA
GUTTIF
GUTTIF
GUTTIF
GUTTIF
Species
Shorea bracteolata Dyer
Shorea curtisii Dyer ex King
Shorea gratissima (Wall. ex Kurz.) Dyer
Shorea leprosula Migq.
Shorea macroptera Dyer
Shorea pauciflora King
Vatica maingayi Dyer
Vatica sp. A
Diospyros buxifolia (Bl.) Hiern
Diospyros lanceifolia Roxb.
Diospyros maingayi (Hiern) Bakh.
Diospyros sp.
Elaeocarpus acronodia BI.
(Elaeocarpus mastersii King)
Elaeocarpus nitidus Jack
Elaeocarpus petiolatus Wall.
Elaeocarpus polystachyus Wall.
Antidesma coriaceum Tul.
Antidesma cuspidatum Muell. Arg.
Aporusa benthamiana Hk. f.
Aporusa bracteosa Pax. et Hoffm.
Aporusa prainiana King ex Gage
Aporusa sp. A
Aporusa sp. B
Baccaurea bracteata Muell. Arg.
Baccaurea kunstleri King ex Gage
Baccaurea maingayi Hk. f.
Baccaurea sumatrana Miq.
Baccaurea sp. A
Baccaurea sp. B
Blumeodendron tokbrai (BI.) Kurz
Elateriospermum tapos BI.
Endospermum diadenum (Migq.) Airy Shaw
(Endospermum malaccense Muell. Arg.)
Koilodepas longifolium Hk. f.
Macaranga conifera (Zoll.) Muell. Arg.
Macaranga lowii King ex Hk. f.
Mallotus penangensis Muell. Arg.
Pimelodendron griffithianum
(Muell. Arg.) Benth.
Ptychopyxis caput-medusae (Hk. f.) Ridl.
Castanopsis megacarpa Gamb.
Castanopsis wallichii King ex Hk. f.
Castanopsis sp.
Lithocarpus conocarpus (Oudem.) Rehd.
Lithocarpus encleisacarpus (Korth) A. Camus
Lithocarpus ewycku (Korth) Rehd.
Calophyllum curtisii King
Calophyllum ferrugineum Ridl.
Calophyllum pulcherrimum
Wall. ex Planch. et Triana
Calophyllum rubiginosum
Hend. et Wyatt-Smith
46
—
BNO NDN ON RRP RUNNY Wee PO Ns NN)
OMe YR
hhRWN WDNR KKH NN
i)
DI
—
NY NO
NOR RR NR RD WN
NNR RRR NOR WR Re
Mm bd
Mm He tO
ee ee
i)
WtrN RK WN RK —
66 Gard. Bull. Sing. 40(1) (1987)
(Appendix 1:a cont.)
Rec. # Family Species Total DI
91. GUTTIF Calophyllum wallichianum Planch. et Triana 1 1
92. GUTTIF Garcinia hombroniana Pierre 2 2
93. GUITIF Garcinia nigrolineata Planch. et T. Anders. Zz 2
94. GUTTIF Garcinia rostrata (Hassk.) Miq. l
95; HYPERI Cratoxylon formosum (Jack) Dyer 3 x
96. IXONAN Ixonanthes icosandra Jack ~ 4
OF: IXONAN Ixonanthes reticulata Jack 20 6
98. LAURA Actinodaphne malaccensis Hk. f. 1
eel LAURA Actinodaphne sesquipedalis 1
Hk. f. et Th. ex Meissn.
100.. LAURA Beilschmiedia maingayi Hk.f. 3 3
101. LAURA Cinnamomum zeylanicum Garc. ex BI. Il 1
102. LAURA Cryptocarya ferrea BI. l |
103. LAURA Crytocarya rugulosa Hk. f. 1 ]
104. LAURA Litsea castanea Hk. f. 8 5
105. LAURA Litsea costalis (BI.) Kosterm. 1 l
106. LAURA Litsea elliptica BI. 2) 3
107. LAURA Litsea gracilipes Hk. f. l
108. LAURA Litsea grandis Hk. f. 2
109. LAURA Litsea machilifolia Gamb. Z 2
110. LAURA Nothaphoebe umbelliflora (BI.) BI. rt l
ia LEGUMI Dialum kingii Prain. + 3
| 2s LEGUMI Dialium laurinum Baker 2 2
113. LEGUMI Dialium maingayi Baker
114. LEGUMI Dialium patens Baker Z l
113, LEGUMI Dialium platysepalum Baker 3 l
116. LEGUMI Dialium wallichii Prain 2 1
117. LEGUMI Koompassia malaccensis Maing. ex Benth. 10 6
118. LEGUMI Sindora coriacea Maing. ex Prain |
LTS: LEGUMI Sindora echinocalyx (Benth.) Prain l 1
120. LEGUMI Sindora velutina Baker l l
iZh. LEGUMI Sindora sp. l l
122: MAGNOL Aromadendron elegans BI. 2 2
1a: MELAST Memecylon coeruleum Jack l
124. MELAST Pternandra coerulescens Jack 2 2
125. MELAST Pternandra echinata Jack 7 5
126. MELIA Aglaia trichostemon C. DC. 2 2
127. MELIA Aglaia sp. 2 2
128. MELIA Chisocheton erythrocarpus Hiern. l l
129, MELIA Dysoxylon sp. S l
130. MELIA Sandoricum koetjape (Burm. f.) Merr. l l
at. MORA Artocarpus kemando Miq. l
132. MORA Artocarpus lanceifolius Roxb. 14 11
133. MORA Artocarpus rigidus BI. 9 6
134. MORA Artocarpus scortechinii King 5 4
135; MYRIST Gymnacranthera eugeniifolia (A. DC.) Sincl. 2 2
136. MYRIST Gymnacranthera forbesii (King) Warb. ]
137. MYRIST Horsfieldia brachiata (King) Warb. | 1
138. MYRIST Horsfieldia superba (Hk. f. & Th.) Warb. 1 1
139. MYRIST Knema hookeriana (Hk. f. et Th.) Warb. l 1
140. MYRIST Knema intermedia (BI.) Warb. l 1
141. MYRIST Myristica cinnamomea King 2 Ps
142. MYRSIN Ardisia tuberculata Wall. 2 1
143. MYRSIN Ardisia teysmanniana Scheff. l 1
Ecology of the Trees of Bukit Timah Nature Reserve
(Appendix 1:a cont.)
Rec. #
144.
145.
146.
147.
148.
149.
150.
151.
152.
23.
154.
155.
156.
157.
158.
159.
160.
161.
162.
163.
164.
165.
166.
167.
168.
169.
170.
Aft.
hid.
£73.
174.
175.
176.
a7.
178.
179.
180.
181.
182.
183.
184.
185.
186.
187.
188.
Family
MYRSIN
MYRTA
MYRTA
MYRTA
MYRTA
MYRTA
MYRTA
MYRTA
MYRTA
MYRTA
MYRTA
MYRTA
MYRTA
MYRTA
OCHNA
OLACA
OLACA
OLACA
POLYGA
POLYGA
POLYGA
RHAMNA
RHIZO
RHIZO
RHIZO
ROSA
ROSA
RUBIA
RUBIA
RUBIA
RUBIA
RUBIA
RUBIA
RUBIA
RUTA
SABIA
SAPIND
SAPIND
SAPIND
SAPIND
SAPIND
SAPIND
SAPIND
SAPOTA
SAPOTA
Species
Maesa ramentacea Wall.
Eugenia chlorantha Duthie
Eugenia filiformis Duthie
Eugenia glauca King
Eugenia linocieroidea King
Eugenia ngadimaniana Hend.
Eugenia palembanica (Miq.) Merr.
Eugenia rugosa (Korth) Merr.
Eugenia subdecussata Duthie
Eugenia sp. A
Eugenia sp. B
Eugenia sp. C
Rhodamnia cinerea Jack
(Rhodamnia trinervia Bl.)
Tristania merguensis Griff.
Gomphia serrata (Gaertn.) Kanis
Ochanostachys amentacea Mast.
Scorodocarpus borneensis Becc.
Strombosia ceylanica Gardn.
(Strombosia rotundifolia King)
Xanthophyllum ellipticum Korth. ex Miq.
(Xanthophyllum kingii Chodat)
Xanthophyllum stipitatum Benn.
Xanthophyllum sp.
Ziziphus calophylla Wall.
Gynotroches axillaris BI.
Pellacalyx axillaris Korth.
Pellacalyx saccardianus Scort.
Licania splendens (Korth.) Prance
Prunus polystachya (Hk. f.) Kalkman
Pertusadina eurhyncha (Miq.) Ridsdale
(Adina rubescens Hemsl.)
Diplospora malaccensis Hk. f.
Nauclea officinalis
(Pierre ex Pitard) Merr. et Chun.
(Nduclea junghuhnii Merr.)
Randia densiflora Benth.
Randia scortechinii King & Gamb.
Timonius wallichianus (Korth.) Valeton
Urophyllum glabrum Wall. ex Roxb.
Euodia glabra (BI.) BI.
(Evodia glabra BI.)
Meliosma pinnata (Roxb.) Walp
Nephelium glabrum Noronoh
Nephelium lappaceum L.
Euphoria malaiensis (Griff.) Radlk.
(Nephelium malaiense Griff. )
Nephelium rubescens Hiern
Nephelium sp.
Pometia pinnata Forst.
f. alnifolia (Bl.) Jacobs.
Xerospermum intermedium Radlk.
Ganua kingiana (Brace) van den Assem
Palaquium gutta (Hk.f.) Baillon
Total
—
— RP OmMndN eR NNONN FP UYNDN OR WR Fe
— eS Re CORR OR ke ff
Ne
DI
SS OR NR eR ff — ep SNR Re WrRNNN NR NN RK NR
NO Re
68 Gard. Bull. Sing. 40(1) (1987)
(Appendix I:a cont.)
Rec. # Family Species Total DI
189. SAPOTA Palaquium hexandrum (Griff.) Baillon 4 1
190. SAPOTA Palaquium microphyllum King & Gamb. = Zz
Si. SAPOTA Palaquium obovatum (Griff.) Engl. 2 Z
192. SAPOTA Palaquium semaram H.J. Lam 6 6
193. SAPOTA Payena obscura Burck 1 1
194. SAPOTA Planchonella maingayi (Clarke) van Royen 1 1
£95. SIMARU Eurycoma longifolia Jack 1 1
196. STERCU Heritiera elata Rid. 2 2
157. STERCU Scaphium macropodum 1 1
(Miq.) Beumee ex Heyne
(Scaphium affine Miq.)
198. STERCU Heritiera simplicifolia (Mast.) Kosterm. 1 1
(Tarrietia simplicifolia Mast.)
199. THEA Adinandra acuminata Korth. 1 1
200. THEA Adinandra dumosa Jack 90 4
201, THEA Ploiarium alternifolium (Vahl) Melchior 1 1
202. THEA Ternstroemia bancana Miq. 1 1
203. THEA Gordonia multinervis King 1 1
204. THEA Gordonia singaporeana Wall. ex Rid. 1 1
205. THYMEL Aquilaria malaccensis Lamk. 1 1
206. TILIA Grewia blattaefolia Corner 5 3
207. TILIA Pentace triptera Mast. S 2
208. MORA Ficus dubia Wall. ex King 1 i
209. ULMA Gironniera nervosa Planch. + 3
210. ULMA Gironniera parvifolia Planch. 8 5
241. ULMA Streblus elongatus (Miq.) Corner 19 7
(Sloetia elongata (Miq.) Koord.) ;
282. VERBEN Teijsmanniodendron coriaceum 2 1
(Clarke) Kosterm.
213. Unknown 20 13
Ecology of the Trees of Bukit Timah Nature Reserve
Appendix 1: b. Families arranged according to descending order of species
preponderence.
(All species with gbh > = 12 inches)
Family No. of No. of Sample
Genera Species Total
1. Euphorbiaceae 11 2D. 70
2. Burseraceae 3 13 4]
a Dipterocarpaceae 5 13 125
4. Myrtaceae a 13 46
=e Lauraceae 6 13 26
6. Leguminosae 3 1] 28
‘sf Anonaceae 5 10 20
8. Guttiferae 2 8 19
2} Sapotaceae 4 8 58
10. Sapindaceae - 7 10
11. Myristicaceae ~ 7 9
Pe. Rubiaceae 6 ws 3}
eo: Fagaceae 2 6 10
14. Moraceae 3 6 49
iS: Anacardiaceae 5 6 58
16. Theaceae 3 5 94
i. Meliaceae 4 5 11
18. Elaeocarpaceae 1 - 6
19. Ebenaceae 1 4
20. Melastomaceae 2 3
21. Sterculiaceae 2 3
Z2: Myrisinaceae 2 3
23. Rhizophoraceae Zz 3
24. Olacaeae 3 3
2S. Polygalaceae 3
26. Ixonanthaceae 1 2
27. Ulmaceae 1 z 2
28. Connaraceae 2 2 2
29. Rosaceae 2 fi 2
30. Apocynaceae 2 2 2
31. Tiliaceae 2 2 7
32. Celastraceae 2 Z 5
53. Rutaceae 1 1 5
34. Hypericaceae 1 1 3
5. Sabiaceae 1 l 2
36. Verbenaceae 1 1 2
af. Ochnaceae 1 | Z
38. Dilleniaceae 1 1 2
39. Magnoliaceae 1 1 2
40. Bombacaceae 1 l l
41. Rhamnaceae 1 l l
42. Simarubaceae 1 | l
43. Ternstroemiaceae l | l
44, Thymeliaceae l l l
45. Unknown 0 0 10
Total: 111 212 889
70 Gard. Bull. Sing. 40(1) (1987)
Appendix 2. Girth class distribution and spatial distribution of species as indicated
by the Distribution Index (DI). ;
All species with girth>=12”’.
G1 & G2 obtained from only 1 of 4 sub-plots of each cluster.
For explanation on DI and girth size (G) equivalents see Sect. III (c) & III (d).
“TOT” is the Sample Total of the Species
Species Code G1 G2 G3 G4 G5 G6 TOT ODI
A. Actinodaphne malaccensis ACMA 0 1.0 ta 1 1
2: Actinodaphne sesquipedalis ACSE 0 0 1 0 ae 1 1
3: Adinandra acuminata ADIN 0.0 .0 3a 1 u
4. Agelaea trinervia AGET O-.0 1 Oe ae 1 1
5. Alstonia angustifolia ALAN 0 1-90 33 1 1
6. Anaxagorea javanica ANAX 0. O: Oy tae 1 1
ip Anisoptera costata ANIS 0. 0-0: hia 1 1
8. Antidesma cuspidatum ANTC 0. 0: 1°00 1 il
9; Aporusa bracteosa APBR 1 2.22.0) ae 5 1
10. | Aporusa prainiana APPR 0... 0 0. eae 1 1
kit. Aporusa sp. A APZA fh 0 2 tS 1 1
i? Aporusa sp. B APZB 0.0 1° Oe 1 1
13: Aquilaria malaccensis AQMA 0-0 0 te 1 1
14. Ardisia teysmanniana ARDT 0 0 the i 1
15. Ardisia tuberculata ARDI 0-2. 2 ia 2 1
16. Artocarpus kemando ARKE Q 40 0b 1 1
Ly. Baccaurea sp. B BAZB tL 0, 0 aia 1 1
18. Bouea macrophylla BOMA 0 0 1 Oye 1 1
19. Bouea oppositifolia BOOP 00° "0; “tycGeao 1 1
20. Calophyllum rubiginosum CALR 0.2) Oe ae 2 1
ZA; Calophyllum wallichianum CALW O°: O20 0.00, ae 1 1
me; Canarium grandifolium CANA 0.0 Ostia 1 1
ZS. Canarium sp CANP 0-1. ON psa 1 1
24. Castanopsis megacarpa CASM 02:0 Oe ae 1 1
23 Castanopsis wallichii CAST 0. 90. 0:54 Re 1 1
26. Castanopsis sp. CAZP Or 0 Oo Tate 1 1
a). Chisocheton erythrocarpus CHER oo 130 aa 1 1
28. Cinnamomum zeylanicum CIZE 0-000 OF he ae 1 1
29. Cryptocarya ferrea CRFE 0.5 0-0 1S ea 1 1
30. Cryptocarya rugulosa CRRU OA SO eee 1 1
a1: Cyathocalyx ridleyi CYRI 00>. 1. a eee 1 1 |
a2; Dacryodes costata DACO Oo © ol. 22a 2 1
33. Dacryodes rugosa DARU 0-10 Se i: ae
34. Dialium maingayi DIMA 0 0. fF Sais 1 1
So Dialium patens DIPA 0 0. f eae 2 1
36. Dialium platysepalum DIPL 0 t 1 Aelia 3 1
Rie Dialium wallichii DIWA 0 0 0: 0.32 2 1
38. Diospyros lanceifolia DOLA 0 0 0 ACS 1 1
39. Diospyros maingayi DOMA 0 0 + aw 1 1
40. Diospyros sp. DOSP 0: - 0. O20. a 1 1
41. Diplospora malaccensis DPMA 0 0.0 SO 2 1 1
42. Dipterocarpus sublamellatus DPRS 0 0.3 0430 3 1
43. Durio griffithii DUGR Oo L010 ae 1 1
44, Dyera costulata DYCO 0. 1. eee 1 1
45. Dysoxylon sp. DYSP 0 4 1 809 @ 5 1
46. Elaeocarpusnitidus ELAN 0.0 2a 1.
47. Elaeocarpus petiolatus ELAP 0 &-@? dA gam 1 1
48. Elaeocarpus polystachyus ELAS 1 0 Deas 1 1
Ecology of the Trees of Bukit Timah Nature Reserve
(Appendix 2 contd.)
Species
49. Ellipanthus tomentosus
50. Eugenia chlorantha
a1. Eugenia glauca
a2. Eugenia palembanica
53. Eugenia sp. C
54. Euphoria malaiensis
SS. Eurycoma longifolia
56. Ficus dubia
a7. Fissistigma lanuginosum
58. Fissistigma latifolium
59: Garcinia rostrata
60. Gomphia serrata
61. Gordonia multinervis
62. Gordonia singaporeana
63. Gymnacranthera forbesii
64. Horsfieldia brachiata
65. Horsfieldia superba
66. Knema hookeriana
67. Knema intermedia
68. Koilodepas longifolium
69. Kokoona reflexa
70. Licania splendens
71. Lithocarpus ewyckii
ae. Litsea costalis
73. Litsea gracilipes
74. Litsea grandis
fey Macaranga conifera
76. Macaranga lowil
77. Mallotus penangensis
78. Maesa ramentacea
9. Memecylon coeruleum
80. Meliosma pinnata
81. Nephelium glabrum
82. Nephelium rubescens
83. Nephelium sp.
84. Nothaphoebe umbelliflora
85. Palaquium hexandrum
86. Payena obscura
87. Pellacalyx axillaris
88. Pertusadina eurhyncha
89. Ploiarium alternifolium
90. Planchonella maingayi
ol: Polyalthia hookeriana
92. Polyalthia rumphii
93. Polyalthia sp.
94. Pometia pinnata f. alnifolia
95. Prunus polystachya
96. Randia densiflora
97. Sandoricum koetjapi
98. Santiria apiculata
B, Santiria sp. A
100. Santiria sp. B
101. Scaphium macropodum
102. Shorea bracteolata
103. Sindora coriacea
104. Sindora echinocalyx
Code
ELTO
BUCH
EUGL
EUPA
EUZD
NEMA
EVLO
FICD
FILA
FILT
GARR
GOMS
GOMU
GOSI
GYFO
HORB
HOSU
KNHO
KNIN
KOLO
LORE
LICA
LIEW
Le
LITG
LITN
MACO
MALO
MAPE
MARA
MECO
MEPI
NEGL
NERU
NESP
NOUM
PAHE
PAYA
PEAX
ADIA
PLAL
PLMA
POHO
POLR
POLS
POMA
PRPO
RADE
SAND
SANE
SAZA
SAZB
SCAF
SHBR
SICO
SIEC
G1 G2 G3 G4 G5 G6 TOT
cococooororocoococoooocooooooooooooorcocococococoococooocoocoocoocoooeeoorse se eee
CoooCoOoOrFroNwconocoocoocooosoroococororwrocococorrooroocoocooocoooeor er onrr SOS
RB OOrrKRroocoonocorrorocoocoocooororrrvooooororoooorroroonorroroneooor
Oem ooororocoocoooroocooooonoorrvocooocooocororrOorocooocooorcocorcoreseeseeeee
coco CoCCoCocooooooocooooorOCSCoCOCSOCOOCOOCOOrFONF COCOoOOrFCOCOOCOOCOCOCCOCCOOCOCOCCOCCOCOCCOSC eer eee e
SCOrcoocooocoornocororrrrocoococoocooocoocooocooocooooooooocsoooorceoceorsce se eeee
See eee NEF OE NNNRP RP RFP RP RP RP BRP RPP NN RP UNONNRP RP NRF RP RP RP RP RP RP RP RP RP NP PNR RRP NN Pre
77
=
FR OO ee een ne NN Ne ce cel cee el ce ae ee cee ce ce cc ee ee
72 Gard. Bull. Sing. 40(1) (1987)
(Appendix 2 contd.)
Species Code G1 G2 G3 G4 G5 G6 TOT ODI
105. Sindora velutina SIVE 0 Oooo Cm 1 1
106. Sindora sp. SIZP 0 0 Oeenorag ey 1 1
107. Strombosia ceylanica STRO 0. 0 YT Oetiee 1 1
108. Heritiera simplicifolia TASI 0 0 eh oOr'eiea 1 1
109. Ternstroemia bancana TEBA 0 0 0 1° aree 1 1
£10: Teijsmanniodendron coriaceum TJCO 0 2 O00" 2 1
it. Tristania merguensis TRME 0 0 0" Ot 1 1
£2. Urophyllum glabrum URGL 0 1 0 0 lea 1 1
£13. Vatica maingayi VASA 0 ©) fe) Gram 1 1
114. Vatica sp. A VASB 0... 1-205 grae 1 1
£15; Xanthophyllum ellipticum XAKI 0:0: 0 Arete 1 |
£16. Xanthophyllum sp. XAZP 0 0 0 “fiegaes 1 1
My. Xerospermum intermedium XEIN 0. 0 Oooo ter 1 1
118. Xylopia ferruginea XYFE 0° OPO eee 1 1
P19. Ziziphus calophylla ZICA Os | al Re 1 1
120. Aglaia trichostemon AGLT 0 0 -O >Re eae 2 2
Ae Aglaia sp. AGSP 0 0 2°00 tie 2 2
B22. Aporusa benthamiana APBE 0 0 De hager ys 2
£23. Aromadendron elegans AREL 1 0 Oe retes 2 2
124. Baccaurea bracteata BABR 1 0° 2405 eee 2 2
125. Baccaurea maingayl BAMA 0 0 2 “Or egea 2 r
126. Baccaurea sp. A BAZA 0 0 1LOOrriad® 2 2
WA Bhesa paniculata BHPA 0 2 2 yet 4 v4
128. Buchanania sessilifolia BUSE 2 0 0 Gyaiere 2 y
be. Calophyllum curtisii CALC 0-0. 0 “OEteeee 3 y
130. Dacryodes laxa var typica DALA 0 1.0. ee 2 =
if. Dacryodes rostrata DARO lL. 0. Oe 2 2
132. Dialium laurinum DILA 0.) 0... 0 SO ire 2 2
135, Dillenia grandifolia DLEX 0. 1 Soares 2 2
134. Diospyros buxifolia DOBU 0 1 aes 2 <
£35. Elaeocarpus acronodia ELAM 0 ORR OVaiy ss 2 y
136. Elateriospermum tapos ELTA 0 .0 Oe 2 2
137. Eugenia filiformis EUFI 0 2. sbi aire 3 2
138. Eugenia linocieroidea EULI 0 O°. S/S ee 2 2
139. Eugenia ngadimaniana EUNG 0 1. 4. 50840008 2 2
140. Eugenia subdecussata EUSU 0 0 4 eee 4 P
141. Eugenia sp. A EUZA 0 0 DeDBe- tee 2 2
142. Eugenia sp. B EUZB 0 .0 2 SarGraG 2 2
143. Euodia glabra EVOD 0 1 ‘beesekiae 5 2
144. Garcinia hombroniana GARH 0 I 4 Reipet 2 2
145. Garcinia nigrolineata GARN 0 bb (bope iis 2 2
146. Gymnacranthera eugeniifolia GYEU 0 1 <)iaw 2 2
147. Heritiera elata HEEL 0 1 Ue" paw 2 2
148. Lithocarpus conocarpus LICO 0 0 Db pore 2 2
149. Litsea machilifolia LITS 0 0 2 Ones 2 2
150. Myristica cinnamomea MYCI 0 1-0 “eae ps 2
151. Nauclea officinalis NAJU 0 0 2 ari 2 2
152. Nephelium lappaceum NELA 0.0 Latest 2 4
153. Palaquium microphyllum PALM 0 0'°0 8 eh 2 2
154. Palaquium obovatum PALO 0 1 tee 2 2
155. Pentace triptera PETR 0 0 2 Diab 3 ys
156. Pternandra coerulescens PTCO 0 0 Io de eee 2 2
157. Ptychopyxis caput-medusae PTYC 1 0. tees 4 2
158. Santiria tomentosa SANT 1 0 eee 3 2
159. Shorea gratissima SHGR 0 0 Oe 3 2
160. Xylopia malayana XYMA 0 2 Rat ere Ps 2
hs / ee
Ecology of the Trees of Bukit Timah Nature Reserve
Ps
(Appendix 2 contd.)
Species Code Gi1G2/G3 G4 G5, G6, TOV. Di
161. Antidesma coriaceum ANOM VSR ae Be i 3 3
162. Beilschmiedia maingayi BEMA Sra a rh, DS 3 4]
163. Calophyllum pulcherrimum CALP ree AO te ye - 3
164. Campnosperma auriculatum CAMA We tbe AGe ISR 52.78 16 3
165. Cratoxylon formosum CRFO Ose Gy deta Os. 1 3 3
166. Dialium kingii DIKI ee he 2 Oe TO 4 3
167. Dipterocarpus kerii DPRI Dit Oi ek py On a2 5 3
168. Ganua kingiana GANK Oe ta een D..-0) - 3
169. Gironniera nervosa GINE B29. dee Oe Os 0 < 3
170. Grewia blattaefolia GRBL Pg aa. oe | SR a | 2 3
M71. Lithocarpus encleisacarpus LIEN OF 8 ch yeaeArn © 3 3
172. Litsea elliptica LITE Op eae Qe 0 3 3
£73. Rhodamnia cinerea RHTR be SR Puyo 16 3
174. Shorea macroptera SHMA PO. GA eee: 1 - 3
£75. Adinandra dumosa ADIU OPE 692. 9 MSZ 0 90 -
176. Artocarpus scortechinii ARSC er Oe baer By 1 5 ~
RYT. Blumeodendron tokbrai BLTO Oh: Raeantabs (1 %.-0 5 -
178. Calophyllum ferrugineum CALF Me ee.) Be Be 10 - -
1/9. Endospermum diadenum ENDO Oe ee ae EO <'O +
180. Hopea mengarawan HOME aes, oe) OD 4 -
181. Ixonanthes icosandra IXIC Bt Os -Ds. 0 - =
182. Palaquium gutta PAGU Oe Sn 29 7 38 4
183. Parishia paucijuga PARP Bae Os Td 4 4
184. Santiria rubiginosa SANR Pa ero Out One 2 5 -
185. Scorodocarpus borneensis SCBO Beacon Ge ie 3 ~
186. Xanthophyllum stipitatum XAST Re Are on ae Ot" 'D 4 a
187. Baccaurea kunstleri BAKU eit ON te 9 5
188. Baccaurea sumatrana BASU Ote2e 2c ce Ut 0 6 5
189. Eugenia rugosa EURA say, hia a | > 5
190. Gironniera parvifolia GIPA ieee ono. 1) eo 8 5
191. Gynotroches axillaris GYNO Pie oe oe Ot O 6 5
192. Litsea castanea LITA pe A a OO 8 5
73. Ochanostachys amentacea OCAM A ems eae alla 5 5
194. Pternandra echinata PTEC ITI oe 8 Se: ae ase etagal | Rea 7 5
2. Shorea leprosula SHALE Riera. Sy Sa 4 10 5
196. Artocarpus rigidus ARRI eee a ret 2 9 6
197. Cyathocalyx ramuliflorus CYRA OA ae BN, OD 7 6
198. Ixonanthes reticulata IXRE aoe ee ee ee TO 20 6
199. Koompassia malaccensis KOMA apg Pel Le OS 10 6
200. Palaquium semaram PALS IS se, Na ea oa 6 6
201. Randia scortechinii RASC ls: Salat, i aah | a 13 6
202. Shorea pauciflora SHPA Ea ae Soe | ees Sie, 9 6
203. Pimelodendron griffithianum PIGR COE Pe: uy | a | 9 7
204. Streblus elongatus SLOE Fa ae ESS oe ee 19 7
205. Timonius wallichianus TIWA PET, al ae Se, | SE, 13 7
206. Santiria griffithi SANG He AS HO AY i aie 12 8
207. Santiria laevigata SANL erg Mls <2" vo Oa E 8 8
208. Pellacalyx saccardianus PESA Be Se. 62 "7 28: 10
209. Artocarpus lanceifolius ARLA RS Tee tie Seas ee 6.
210. Dipterocarpus caudatus DPRP Os 84.7 a ees a (in 0
ssp. penangianus
ei. Shorea curtisii SHCU i he i a 46 12
zi. Unknown ZNKN boi Bai he ie p | ae
zi. Gluta wallichii MELW ee PS Ne eos 2 oe, ke
74 Gard. Bull. Sing. 40(1) (1987)
Appendix 3. Species with individuals in specified ranges of girth classes.
“TOT” is the Species’ Sample Total.
For DI and G see Sect. III (c) & III (d)
a Species with individuals in the whole range of girth classes.
Species Code G1, G2 G3 G4 G5 G6. tO
i. Streblus elongatus SLOE 1 SoS Gre eee 19 ¥
b Species which must have individuals from G2 thr’u to G6.
Species Code G1 G2 G3 G4 G5 G6 Tor
{. Artocarpus lanceifolius ARLA O' “3 “Alaa 14 il
Zz Artocarpus rigidus ARRI On syle Ms Se Gale 9
3. Campnosperma auriculatum CAMA O V1 26 “SS 16 3
4. Dipterocarpus caudatus DRRP 0. *8 >: Paes Oo] ae
ssp. penangianus
Ss: Gluta wallichii MELW 0 5 “he ae 34. 8S
6. Palaquium gutta PAGU 0 210s eae 38 4
ve Pellacalyx saccardianus PESA 0 °5@0 9) SOUS 28 S10
8. Shorea curtisii SHCU 0 oS LOrrks Wire 46 12
D: Streblus elongatus SLOE l 3h SOi2e 19 7
c Species which must have individuals from G3 thr’u to G6.
Species Code Gi G2 G3 G4'GS GGA toa
i. Atrocarpus lanceifolius ARLA 0 3. 40a 14). a4
Lr. Artocarpus rigidus ARRI 0 vd ea ae 9
a: Artocarpus scortechinil ARSC 0) OD ck ee 5 4
4. Campnosperma auriculatum CAMA 01 \ O<558\Ggee 16 3
a Dipterocarpus caudatus DPRP 0 8 =e cee 37 cnhe
ssp. penangianus
6 Ixonanthes reticulata IXRE 0 0 S40 4aee 20 6
Ts Gluta wallichii MELW 0 5 ASS2O eee 34> nS
8. Palaquium gutta PAGU 0.°2, 10 Ta are 38 4
2: Palaquium semaram PALS 0. 0° decd aeah ee 6 6
10. Pellacalyx saccardianus PESA 0° °6 Qu bot arene 28 0
ii, Shorea curtisil SHCU 0. 5 10 (26) peeare 46 2
‘2. Streblus elongatus SLOE lL 3 disse 19 4
d Species which must have individuals from G4 thr’u to G6.
Species Code G1 G2 G3 G4 GS G6. TOT) Wa
i Artocarpus lanceifolius ARLA 0. 3 of eee 14 oe
2 Artocarpus rigidus ARRI 0 1,2 30g 9
2. Artocarpus scortechinil ARSC 0 0. 4d) 2. 5 4
4. Campnosperma auriculatum CAMA 0 .1 6 73a 16 3
>. Dipterocarpus caudatus DPRP 0 8 7. aaa oT he
ssp. penangianus
6 Ixonanthes reticulata IXRE 0.0 4 2a 20 6
re Gluta wallichii MELW 0 5 35 ee aoe 34
8. Palaquium gutta PAGU 0 2 dO. ee 38 4
9. Palaquium hexandrum PAHE O 0 it ee 4 ]
10. Palaquium semaram PALS 0 0. lee oes 6 6
11. Pellacalyx saccardianus PESA QO 6, OB 28. 10
12. Santiria griffith SANG | &« 2 See 12 8
M3. Shorea curtisii SHCU 0 .5 40, See ae 46 12
14. Shorea leprosula SHLE 0 . 2.0. ae 10 5
b. Shorea macroptera SHMA 0 0 @ gees 4 3
16. Streblus elongatus SLOE 1 3.:\ghe ia 19 2
Ecology of the Trees of Bukit Timah Nature Reserve
“TOT” is the Species’ Sample Total.
girth class G2.
For G and D1 see Sect. III (c) & III (d) for explanation.
_
_
ee
Nee Sc ee ate a fe
tw
a
WWNONNINNN NM WN LY
BSSRBIAARONS
ow OS Bee to Garo Go's
SAURRESHSHESSRZARAEGH
ee Se See ee
Species
Actinodaphne malaccensis
Adinandra dumosa
Alstonia angustifolia
Aporusa bracteosa
Aporusa sp. A
Ardisia tuberculata
Aromadendron elegans
Artocarpus lanceifolius
Artocarpus rigidus
Baccaurea bracteata
Baccaurea kunstleri
Baccaurea sumatrana
Baccaurea sp. B
Bhesa paniculata
Blumeodendron tokbrai
Buchanania sessilifolia
Calohyllum pulcherrimum
Calophyllum rubiginosum
Campnosperma auriculatum
Canarium sp
Chisocheton erythrocarpus
Cryptocarya rugulosa
Cyathocalyx ramuliflorus
Dacryodes laxa var typica
Dacryodes rostrata
Dacryodes rugosa
Dialium kingii
Dialium platysepalum
Dillenia grandifolia
Diospyros buxifolia
Dipterocarpus caudatus
ssp. penangianus
Durio griffithii
Dyera costulata
Dysoxylon sp.
Elaeocarpus polystachyus
Endospermum diadenum
Eugenia chlorantha
Eugenia filiformis
Eugenia glauca
Eugenia ngadimaniana
Eugenia palembanica
Eugenia rugosa
Euodia glabra
Ganua kingiana
Garcinia hombroniana
Garcinia nigrolineata
Gironniera nervosa
Gironniera parvifolia
Appendix 4. Number of species which have individuals in girth class G1 and/or
Sl? G25G3s- G4 GS. G6 - TOT
"yooh a Yn Ny i oe ey ee ne orgs
DS) SS SS CS) BS BS
=)
0)
0
l
\O
=)
Nn —
—
Re nAON OHO FN NY
—
Ww eRe NO DN ADR RK OW LN MN
os)
“J NM YO
mM NMR Were fe nee
frNONM SN
io 0)
©
fous
NNO De NOR RR Re fe
Nn
Nn
WN Ne
mM Mm hw KR WR YN OR RR De
—"
eo ae et
No
M©mmDWNN WN Nn —
76 Gard. Bull. Sing. 40(1) (1987)
(Appendix 4 contd.)
Species Code GI G2-G3 G4 G5 G6" Tore:
49, Grewia blattaefolia 0 Lf 4° 0 7o 5 3
50. Gymnacranthera eugeniifolia 0. 1. wa ae 2 2
Sf. Gynotroches axillaris 0 (2.92 2 6 5
D2: Heritiera elata 0 1 0... eae Zz 2
33. Hopea mengarawan O° oP a ee * a
54. Horsfieldia brachiata 0 1 SOO ae 1 1
33: Ixonanthes icosandra 0 2°°2° ee 2 4
56. Knema intermedia 0 1 =O ee 1 1
a7. Koilodepas longifolium 0 1. 0 De 1 1
58. Koompassia malaccensis 0 SS. 4S 10 6
Dep Macaranga lowi 0 Pf 2 eee 1 il
60. Mallotus penangensis L340 Sie 5 1
61. Maesa ramentacea Oot oy Oe eee 1 1
62. Gluta wallichii 0 5 Bere os 34-1 aS
63. Meliosma pinnata 0 4. Si See 2 2
64. Myristica cinnamomea 0 1° 0 a eee 2 ys
65: Euphoria malaiense 1 0 0 3" re 1 1
66. Nothaphoebe umbelliflora 0.2. 20 ae 1 1
67. Ochanostachys amentacea 0 2° ae 5 5
68. Palaquium gutta 0 200 =a 38 4
69. Palaquium obovatum eT a at See mee 2 f)
70. Pellacalyx saccardianus 06. "SS Gir eter 20 eat
Th, Pimelodendron griffithianum ONE VS “2 ee 2 7
(va Polyalthia sp. 0... 2..°)0) SOR 2 1
73: Pternandra echinata OTe oY sae eee i 5
74. Ptychopyxis caput-medusae L, OO ee 5 %
79: Randia densiflora LS 2 Sa 6 1
76. Randia scortechinii 0. 7" BS eee 13 6
7. Rhodamnia cinerea 1.6 48% TE Sa 16 3
78. Santiria apiculata 1° od: OY OS 2 1
79. Santiria griffithii 140) ae 12 8
80. Santiria tomentosa LO Oa eee 2 1
81. Shorea curtisii 0.75. 10° oe AG Ae
82. Shorea leprosula 0. <2. Fr wee 10 <i
83. Streblus elongatus Ll 3 Se 6 19 7
84. Timonius wallichianus l 6) 3.3 13 7
85. Teijsmanniodendron coriaceum 0 2 20a y 1
86. Urophyllum glabrum 0.1 0-0 1 1
87. Vatica sp. A 0 1 0] Ct 1 1
88. Xylopia malayana Q 2°00 toe 4 2
89. Unknown 1 6 3) ole 1S Ta
The list above contains 87 known species and the “unknown” group with 20 trees. The individuals are
distributed in the various girth classes as shown below:
G1 G2 G3 G4 G5 G6 TOT
21 181 202 102 45 82 633
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A)
THE GARDENS’ BULLETIN
SINGAPORE
VOL. 40 (Part 2) 1 December 1987
CONTENTS
cr
| PAGES ‘
BIDIN, AZIz: \
A Preliminary Survey of the Fern Flora of Langkawi Islands .....................4. 77-102 &®
LEONG, CHEE CHIEW: \
MSMaA for Controlling Cyperus kyllingia, Axonopus compressus and i
Brachiara distachya in Tifgreen Bermuda Grass Turf .................cceeceseeeeeeees 103-112 \
KENG, HSUAN:
Annotated List of Seed Plants in Singapore (XI) ...............cccccceseeeeeeeeeeeeees 113-132
FOONG, THAI WU and YANG, CHENG NOI:
Resolving Iron Deficiency in Wrightia religiosa by Foliar Analysis
and its Amelioration Using an Iron Chelate as a Soil Additive ................ 133-137
LEONG, CHEE CHIEW:
Chemical Growth Retardation of Baphia nitida with PP333 ..................0e000s 139-144
VERHEIJ, EW.M.:
Growth and Yield of Mango cv Golek in Java Over 25 Years .............sceeeeeeees 145-152
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THE GARDENS’ BULLETIN
SINGAPORE
VOL. 40 (Part 2)
1 December 1987
CONTENTS
BIDIN, AZIz:
A Preliminary Survey of the Fern Flora of Langkawi Islands .............
LEONG, CHEE CHIEW:
MSMA for Controlling Cyperus kyllingia, Axonopus compressus and
Brachiara distachya in Tifgreen Bermuda Grass Turf ....................005:
KENG, HSUAN:
Annotated List of Seed Plants in Singapore (XI) .................. cece eee e ees
FOONG, THAI WU and YANG, CHENG NOI:
Resolving Iron Deficiency in Wrightia religiosa by Foliar Analysis
and its Amelioration Using an Iron Chelate as a Soil Additive .......
LEONG, CHEE CHIEW:
Chemical Growth Retardation of Baphia nitida with PP333 ...............
VERHEIJ, E-W.M.:
Growth and Yield of Mango cv Golek in Java Over 25 Years ...............
Published by the Botanic Gardens
Parks and Recreation Department
Ministry of National Development
Cluny Road, Singapore 1025.
PAGES
Se eee 77-102
de 103-112
Rem A: 113-132
daa we 133-137
ea ape es 139-144
sae 3 145-152
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A Preliminary Survey of the Fern Flora
of Langkawi Islands
AZIZ BIDIN
Botany Department, Faculty of Life Sciences, Universiti Kebangsaan Malaysia
Bangi, 43600 UKM Bangi, Selangor, Malaysia
Abstract
A survey of the fern flora of Langkawi Islands is presented, including a summary of their habitats
and ecology. Of the 145 infrageneric taxa recorded from the islands, 7 are introduced or naturalized and
6 are presumed rare or endemics. Species are listed for both indigenous and secondary plant communities.
Introduction
Langkawi, the natural paradise in the northwestern coast of Peninsular Malaysia
is noted for its sun, unspoilt white sandy beaches and clear blue water. This group
of islands lie between 22 and 54 km off the coast of Perlis, the northern-most state
of Peninsular Malaysia, and 112 km north of Penang. Compared to the mainland,
these 99 islands are relatively free of any kind of pollution. Only the blasting of
limestone in the Kubang Badak Quarry, may have caused considerable noise and dust
pollution within its operational area. Traffic density is light and the only noticeable
pollution is smoke from the burning padi straws and rice husks during harvesting
season. The white sandy beaches of Pantai Cenang, Tanjung Rhu and Teluk Barau
are Only a few of the major tourist attractions of Langkawi. The beaches have been
free of oil spillage from ships, rubbish or sewage. The islands are covered with gent-
ly rolling hills of lush tropical rainforest. They are sheltered from the effects of tropical
storms by the chains of islands not very far from the shore lines of the main island.
Langkawi is one of the great natural assets of Malaysia.
Geography and Geology
Langkawi comprises six moderate sized islands and ninety-three other small islets
(Fig. 1). These islands, formed of and flanked by towering masses of limestone as
well as quartzite and granite are situated at 6°10’ to 6°27’ and 99°37’ to 99°56’
and lie about 48 km off the coast of mainland Kedah. Langkawi Island is by far
the biggest and more important, with an area of about 939 square km. Next in size
is Dayang Bunting Island, 24 km long, with Tuba Island on the east, separated by
a strait so narrow that is appears like a river. West of Dayang Bunting is Singa Besar
Island, Beras Basah Island and many islets. On the southwest of Langkawi Island
is Rebak Besar and Tepur Islands; on the east Langgun and Tembus Islands, while
on the southeast is Timun Island, separated from the main island by the deep and
narrow Strait of Panchor. With the exception of Langkawi, Dayang Bunting and
Tuba, the other islands are either uninhabited or support a handful of temporary
occupants of fishermen.
The centre of Langkawi Island, composed of granite and dominated by Gunung
Raya — the highest peak on the island, is almost entirely primary forest with few
77
78 Gard. Bull. Sing. 40(2) (1987)
access routes into it. It is the only Virgin Jungle Reserve (VJR) with an area of
149 ha demarcated in Langkawi (Putz, 1978). The primary forest is undisturbed and
representative of the forest here. The slopes are characteristically steep and severely
drained. Large trees are found in scattered patches where milder slopes have allowed
deeper soil to develop. North of Gunang Raya towards the coastal strip known as
Pantai Pasir Hitam, is secondary forest, much of which has been cleared to make
way for settlement and a cement factory in Kampung Ewa. Towards the northwest,
there is a long serrated quartzite ridge with about fifteen peaks clad with virgin jungle
which is appropriately called ‘‘Gunung Machinchang’’, the Chopped Mountain.
The middle-west and centre of Langkawi Island consist solely of granite. Again
there is granite in Dayang Bunting, Tuba and Bumbun islands. Fine crystalline
marble is found at the northeast end of Dayang Bunting, Rebak, Tepur, Beras Basah
and Singa Besar islands and numerous islets in the vicinity are composed of quartzite
and shale.
Limestone rock least affected by the granite is seen in certain islets such as Jong
and Kora as well as in the bigger islands such as Langgun, Tanjung Dindang, Timun,
Dayang Bunting and the northeast coast of Langkawi. The limestone is part of the
carboniferous limestone formation that is also found in Sumatra, north of Penin-
sular Malaysia, Thailand, Indochina and Burma. The limestone hills rise from flat
or undulating plains often with steep or over-hanging cliffs and some of them rise
sheer from the water’s edge where part of or even the entire coast is entirely limestone.
Climate
The tropical islands of Langkawi share the same climatic patterns with northern
Kedah in the Malay Peninsula. Day temperature is high and lies within a fairly
narrow range of 35°-37° (Jones, 1978). Water temperature is around 26°-28°.
Rainfall recorded for ten years (1965-1975) from four districts average 260 cm
annually (Jones, 1978). The islands experience a distinct and more severe dry season
from November/December to March than is associated with the most part of Penin-
sular Malaysia. This corresponds to the northeast monsoon when the average monthly
rainfall for January and February hardly exceeds 5 cm compared with an average
for the year of about 260 cm. Two wet seasons occur during the change of mon-
soons, that is, during April and May when the monthly rainfall averages over 22.5
cm, and August to October when the average reaches over 27.5 cm per month. Bet-
ween these two wet seasons occurs another dry period in June and July correspon-
ding to the southwest monsoon when the rainfall averages between 17 and 20 cm.
Observations
Langkawi has been the subject of many general biological investigations in the
last hundred years or so (Maxwell, 1887; Ridley, 1908 and 1912; Henderson, 1939;
Igo and Koike, 1966; Wilcocks, 1969; Yadab and Ratnasabapathy, 1974; Balgooy
et al., 1977; Chin, 1977; Kuthubutheen, 1981). However comprehensive accounts of
any particular group of plants especially ferns, are still lacking. Sporadic collections
have been made from time to time and among the notable fern collectors of the islands
are M.R. Henderson, H.C. Robinson, C. Curtis, Md. Haniff, Md. Nur, H.N. Ridley,
R.E. Holttum and recently Chin, R. Jaman and A. Bidin. Most of the specimens
are kept at the herbaria in Singapore (SING), The Universiti Kebangsaan Malaysia
Bangi (UKMB), University of Malaya (KLU) and Kew (K).
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79
{S P. CHORONG
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80 Gard. Bull. Sing. 40(2) (1987)
Present Account
The present survey is based on the results of a number of field trips carried out
in January, February, April and May 1986 by the author and a number of field
assistants under a project sponsored by the Universiti Kebangsaan Malaysia in Bangi,
as well as on observations of herbarium specimens deposited at the University of
Malaya, The Universiti Kebangsaan Malaysia, Bangi and The Royal Botanic Gardens,
Kew.
Field Observations
A striking feature of the scenery in Langkawi is the occurrence of two main types
of vegetation, limestone and granite. The vegetation of the limestone is primary forest,
and only slightly disturbed. Only where the area has been disturbed by quarrying
and similar activities does secondary forest appear. Much of the limestone cliffs are
honeycombed with caves. On exposed headlands of Langkawi and Dayang Bunting
islands as well as the entire surfaces of some of the rocky islets, the vegetation is
often stunted and reduced, the rocks being covered with grasses, bamboos and occa-
sional stunted trees. Only in areas sheltered from the wind is the vegetation more
luxuriant with a denser growth of larger trees. Among the common ferns found on
the rocky headlands with constant exposure to sea sprays are Davallia denticulata,
Pyrrosia spp., Drynaria sparsisora and Asplenium spp.
The granite ridges on the other hand have a much softer outline, the vegetation
is markedly more luxuriant, consisting of many trees, shrubs, climbers and epiphytes.
Most of the alluvial areas of the coastal plains of Langkawi Island have been cleared
and cultivated, and sometimes abandoned. Here the most abundant and conspicuous
of all the sun-loving ferns are the well-known ‘‘Resam’’ Dicranopteris linearis and
‘*Ribu-ribu’’ Lygodium microphyllum. D. linearis covers considerable tracts right up
to the edges of the forest where it produces dense thickets, while L. microphyllum
scrambles over other plants, and through the grasses. The rice-fields abound with
Marsilea crenata, Azolla pinnata and Ceratopteris thalictroides. C. thalictroides is also
commonly found in irrigation canals. In the mangrove, which covers more than 1474
hectares (nearly 8 per cent of the Island’s land surface), the two species of Acrostichum,
namely A. aureum and A. speciosum are abundant. On earth banks where exposure
to sunlight is maximal, Nephrolepis biserrata and Blechnum orientale are common.
The plain to the north of Kuah is largely rubber plantation and scattered settle-
ment areas. Here, the common ferns are both terrestrial and epiphytic. Nephrolepis
biserrata and Stenochlaena palustris, with their ability to reproduce vegetatively,
flourish. Patches of D. linearis are everywhere, especially where the rubber canopy
is thin. Platycerium coronarium, P. holttumii, Goniophlebium persicifolium, Davallia
denticulata, Drynaria sparsisora, D. rigidula, Drymoglossum piloselloides, Asplenium
nidus and Vittaria ensiformis are commonly found as epiphytes on the trunks of Hevea
brasiliensis as well as on other trees by the Kuah-Kisap Road. Doryopteris ludens,
Adiantum latifolium (introduced from Tropical America but naturalised and
widespread not only in Langkawi but also all over Peninsular Malaysia) and Microlepia
speluncae are abundant on stream banks in the rubber plantations. Besides the two
species of Platycerium mentioned, the new species Platycerium platylobum (Bidin
and Jaman, 1986) is only found on this plain, although uncommon. Helminthostachys
zeylanica and Schizaea digitata are very common in lightly shaded areas of the plan-
tation, growing among the grasses. The aerial part of the former is eaten as a vegetable
while the rhizomes of both species are used alone or by mixing with other herbs in
traditional medicines, supposedly as an aphrodisiac. The most popular vegetable fern
(pucuk paku) among the locals is Stenochlaena palustris. The reddish juvenile frond
*
a
Fern Flora of Langkawi Islands 8]
is eaten raw or cooked with coconut milk. The fern is abundant in open areas such
as settlements, inhabiting wet places, especially stream banks.
Along the tracks leading to ‘‘Telaga Tujuh’’ (Seven Wells) (situated at the ex-
treme western end of Langkawi Island) and Gunung Raya (the highest point on the
Island overlooking the town of Kuah) the presence of Preris scabripes with its whitish
streaks near the frond margins is very conspicuous. The fern has also been observed
on other parts of Langkawi Island but not on any other islands in the group.
Gunung Raya, a granite peak situated in the middle of Gunung Raya Forest Reserve
is highly accessible. Most of the collections were made here. It is extremely rich in
Trichomanes, Hymenophyllum, Asplenium and Bolbitis. Bolbitis malaccensis is
elsewhere known only from one collection in Peninsula Thailand and from Tioman
Island, on the eastern side of Peninsular Malaysia. Microlepia strigosa is found at
the summit of Gunung Raya as well as the foot of Malut Hill (c. 100 m) and the
collections made constitute the third location for the species recorded for Peninsular
Malaysia. Among the members of the family Thelypteridaceae, the genus Christella
is present from lower elevations (C. papilio) right through to the summit of Gunung
Raya (C. subpubescens).
Pseudodrynaria coronans is collected from the Gunung Raya Forest Reserve, near
the summit. The plant has its distributional area on the Asian mainland.
In Langkawi and Dayang Bunting it is observed that the sporophyte of some species
of ferns (Cheilanthes tenuifolia and Adiantum philippense) perennate during the dry
season especially in January and February by means of their rhizome while the fronds
completely die down. Other species maintain at least a few of their leaves in an ex-
tremely dehydrated but living state, e.g. Pyrrosia penangiana and Humata pectinata,
both of which are ferns of the limestone rock.
Drynaria bonii (a new record for Peninsular Malaysia) is found in Langgun Island
only, living epiphytically on stunted trees or on limestone rocks with direct exposure
to sunlight and sea sprays. Records at herbaria show that the plant is distributed mainly
in Thailand as well as in Indo-China.
Much of Timun Island is inaccessible and the common ferns found on the limestone
cliffs are Adiantum stenochlamys, A. caudatum, A. philippense, Tectaria variolosa
(probably the only location for Malesia (Holttum, pers. comm., 1986, Holttum, 1985),
Drynaria sparsisora and Davallia denticulata.
The genus 7richomanes is abundant in Langkawi with eight species, most of them
collected in the Gunung Raya Forest Reserve vicinity.
Of the fifteen genera in Thelypteridaceae in Malaya (Holttum, 1980), six are found
in Langkawi. All are found in the granite area and mostly in the lowlands. Pro-
nephrium is confined to the lowlands only whilst Amphineuron and Christella are
the ferns of lower elevations as well as the mountains. The mountain species are Am-
phineuron immersum and Christella subpubescens — both found near the summit
of Gunung Raya.
Among the Dennstaedtiaceae, Lindsaea (7 spp.) Pteris (7 spp.), Asplenium (9 spp.)
Tectaria (5 spp.) are well represented in Langkawi. Lindsaea which prefers moist deep
shade, is common near the many streams in Gunung Raya Forest Reserve, whilst
Pteris, Asplenium and Tectaria inhabit a variety of habitats. As for 7ectaria, a brief
mention should be made of two lowland species namely 7: brachiata and T: variolosa,
wrongly united by Holttum (1981b; 137). 7: brachiata is widely distributed on granite
in light shade in Langkawi: it is common in rubber plantations in Kisap as well as
in forested areas along trails where the canopy is slightly open. It has a distributional
area from NE. India to S. China and Taiwan, Thailand, Vietnam, northern Malaya
and Jaya. Holttum (1981b) reported that the species is adapted to a climate with a
seasonal dry period, which Langkawi certainly has. It would be interesting to see
82 Gard. Bull. Sing. 40(2) (1987)
if the species is present on Tioman Island on the eastern side of Peninsular Malaysia
which has a similar climate. 7: variolosa, a limestone fern which occurs from NE
India southwards into Burma, Thailand and Vietnam is found in Langkawi on
limestone cliffs and in crevices by the sea on Timun Island. It is a new record for
Peninsular Malaysia.
Ferns of Economic Importance
Ornamentals
Some of the ferns of Langkawi have also found their way into homes and gardens.
Among the popular species are Asplenium nidus, A. phyllitidis, Platycerium coro-
narium, Microsorium punctatum and Adiantum stenochlamys. Quite a number of
ornamental ferns which are not of local origin are also popular as pot as well as garden
plants. These are Adiantum trapeziforme, A. peruvianum, A. mathewsianum, A.
tenerum, A. capillus-veneris and Nephrolepis biserrata var. exaltata. All are Tropical
American ferns which flourish in Langkawi as well as in other parts of the country
and which sometimes escape from cultivation and become part of the natural
vegetation.
Other Uses
In the daily life of the Malays in Langkawi, the fern Lygodium has a variety of
uses. The long climbing rachises are used as twine and said to be long-lasting. Peel-
ings from the rachises are used to make handbags, purses, mats and fish traps.
List of Ferns Collected and Recorded from Langkawi Islands
The following list of ferns is arranged systematically. For each species the scien-
tific name and the original author with its reference is given. Collection numbers and
localities are cited: PL numbers refer to specimens collected during the course of
the studies while specimens from herbaria and their collectors are mentioned also.
Vernacular names are given where known.
The classification adopted follows Holttum (1968) and his subsequent reappraisal
of the families Gleicheniaceae, Schizaeaceae, Cyatheacease and Thelypteridaceae in
Flora Malesiana (1959, 1963, 1978, 1981a). Names of 7ectaria follow Holttum (1981b
and 1985), and of Lindsaea follow Kramer (1971). Materials I have not seen are in-
dicated by relevant literature citations, while records based upon field identification
by the author without voucher specimens being collected are indicated as sight records.
All materials collected are lodged in UKMB and duplicates in K (where indicated).
OPHIOGLOSSACEAE
Helminthostachys zeylanica (Linn.) Hook., Gen. Fil. t.47b. 1840.
Paku tunjuk langit; akar paku.
PL 78: Kisap; Boon Siew Estate. Terrestrial, in light shade, c. 40 m elevation.
UKMB, duplicate at K.
MARATTIACEAE
Angiopteris evecta (Forst.) Hoffm., Comm. Soc. Reg. Gott. 12: 29, t.5. 179b.
Paku gajah.
~--
Fern Flora of Langkawi Islands 83
PL 122: Singa Besar Island. Terrestrial, in deep shade, c. 5 m elevation. UKMB.
PL 281: Gunung Raya Forest Reserve; Genting Palas, Terrestrial, on slope, in deep
shade, c. 550 m elevation. UKMB, duplicate at K.
SCHIZAEACEAE
Schizaea dichotoma (L.) J.E. Smith, Memb. Acad. Turin 5: 442. t.9. 1793; Holttum
1959: 41.
Paku tumbak; paku jarum.
No. 7145. Md. Haniff & Md. Nur. Gunung Raya. Terrestrial. 13 November 1921. K.
Schizaea digitata (L.) Swartz, Syn. Fil.: 150, 380. t.4. f.1. 1896; Holttum 1959: 41.
Janggut adam; misal ikan keli.
PL 148: Road junction of Kampong Buta and Kampong Kubang Badak. Ter-
restrial, in light shade, c. 90 m elevation. UKMB.
PL 282: Gunung Raya Forest Reserve; Genting Palas. Terrestrial, in light shade,
c. 550 m elevation. UKMB, duplicate at Ky ~
Kisap; Boon Siew Estate. Terrestrial, in light shade, c. 40 m elevation. Not collected.
Lygodium polystachyum Wall. ex Moore, Gard. Chron. 1859: 671; Holttum 1959: 46.
PL 30: Bukit Malut Forest Reserve. Terrestrial and scrambling, in light shade,
c. 30 m elevation. UKMB, duplicate at K.
PL 113: Road to Kuala Temoyong. Terrestrial, in light shade, c. 10 m elevation.
UKMB.
PL 19]: Bumbun Besar Island. Scrambling on bushes, exposed, c. 20 m elevation.
UKMB.
PL 257: Selat Panchor Forest Reserve; track to Tanjung Apau. Terrestrial, in light
shade, c. 110 m elevation. UKMB.
Lygodium microphyllum (Cav.) R. Brown, Prod. FI. Nov. Holl: 162. 1810; Holttum
1959: 47.
Ribu-ribu selada; ribu-ribu; salada.
PL 115: Kedawang; Kampong Lubuk Buaya. Terrestrial, in light shade, c. 50 m
elevation. UKMB, duplicate at K.
Lygodium flexuosum (L). Sw., Schrad. J. Bot. 1800(2); 106. 1801; Holttum 1959: 53.
Akar sidin; ribu-ribu gajah; darai paya.
PL 128: Seven Wells, by trackside. Scrambling, in light shade, c. 100 m elevation.
UKMB.
PL 180: Kisap. Terrestrial, on roadside, exposed, c. 30 m elevation. UKMB, 2
sheets.
Lygodium flexuosum (L.) Sw. X Lygodium salicifolium Presl; Holttum 1959: 53.
PL 166: Kampong Sungai Taru. Scrambling on river bank, exposed, c. 20 m eleva-
tion. UKMB.
PL 40: Kisap; Boon Siew Estate. Scrambling, exposed, c. 40 m elevation. UKMB.
Lygodium salicifolium Presl, Suppl, Pterid.: 102, 1845; Holttum 1959: 57.
No. 2739. E.A. Turnau, 5 May 1961. K.
H.C. Robinson. Dayang Bunting Island. Terrestrial. November 1916. K.
84 Gard. Bull. Sing. 40(2) (1987)
PL 29: Bukit Malut Forest Reserve. Terrestrial, scrambling on 7remma, by road-
side, in light shade, c. 30 m elevation. UKMB, duplicate at K.
PL 18]: Gunung Raya Forest Reserve: Lubuk Semilang. Terrestrial, scrambling,
in light shade, c. 40 m elevation. UKMB, duplicate at K.
PL 226: Kisap; Boon Siew Estate, near Kedah Marble Factory. Terrestrial, scrambl-
ing, in light shade, c. 40 m elevation. UKMB.
Lygodium circinnatum (Burm.f.) Sw., Syn. Fil.: 153. 1806; Holttum 1959: 59.
Paku jari merah; ribu-ribu duduk; ribu-ribu bukit.
PL 183: Kisap; Kuala Kisap. Scrambling, in light shade, c. 5 m elevation. UKMB.
GLEICHENIACEAE
Dicranopteris linearis (Burm.f.) Underw., Bull. Torr. Bot. Cl. 34. 249: 1907. var.
linearis; Holttum 1959: 33.
, Resam.
PL 92: Klebang-Ulu Melaka Road. By roadside. Terrestrial, exposed, c. 50 m eleva-
tion. UKMB, duplicate at K.
PL 185: Kisap; Kuala Kisap. Terrestrial, in light shade, c. 5 m elevation. UKMB.
Dicranopteris speciosa (Presl) Holtt., Reinwardtia 4: 273. 1957; Holttum 1959: 32.
PL 175: Gunung Raya Forest Reserve: Lubuk Semilang. Terrestrial, in light shade
c. 40 m elevation. UKMB.
’
HYMENOPHYLLACEAE
Hymenophyllum denticulatum Sw., Schrad. Journ. 1800 (2): 10. 1801.
No. 7168. Md. Haniff & Md. Nur. Gunung Raya. Epiphyte. 13 November 1921. K.
No. 7051. Md. Haniff & Md. Nur. Gunung Raya. Epiphyte. 13 November 1921. K.
Trichomanes digitatum Sw., Syn. Fil. 370. 1806.
No. 7122. Md. Haniff & Md. Nur. Gunung Raya. Epiphyte. 13 November 1921. K.
PL 134: Gunung Raya Forest Reserve; Durian Perangin Waterfall. On rock, in
light shade, c. 200 m elevation. UKMB.
Trichomanes proliferum Bl., Enum. Pl. Jav. 224. 1828.
No. 1064. Md. Haniff. Telaga Tujuh. Epiphyte. 18 September 1914. K.
Trichomanes pallidum Bl., Enum. Jav. 225. 1828.
PL 217: Summit of Gunung Raya. On rock, in streambed, in deep shade, c. 950 m
elevation. UKMB.
Trichomanes bipunctatum Poiret, in Lamarck, Encycl. 8: 60. 1808.
PL 202: Gunung Raya Forest Reserve; Genting Palas. On rock, in light shade,
c. 800 m elevation. UKMB, duplicate at K.
Trichomanes latemarginale Eaton, Proc. Am. Acad. 4: 111. 1259.
No. 15700. M.R. Henderson. Adang Island. Epiphyte. 1911. K.
Trichomanes javanicum Blume, Enum. Pl. Jav. 224. 1828.
No. 2433. C. Curtis. Machinchang. Epiphyte. February 1899. K.
No. 15699. M.R. Henderson. Adang Island. Epiphyte. 1911. K.
No. 15672. M.R. Henderson. Seven Wells. Epiphyte. 1911. K.
Fern Flora of Langkawi Islands 85
PL 21: Bukit Malut Forest Reserve. Terrestrial, in deep shade, c. 160 m elevation.
UKMB.
PL 264: Gunung Raya Forest Reserve; Sungai Kelian. On rock, in deep shade,
c. 150 m elevation. UKMB.
Trichomanes maximum Bl., Enum. Pl. Jav. 228. 1828.
No. 15484. Md. Haniff & Md. Nur. Gunung Raya. February 1911. K.
PL 273: Gunung Raya Forest Reseve; Sungai Kelian. On mossy rock in the
middle of Sungai Kelian, in deep shade, c. 460 m elevation. UKMB, duplicate at K.
Trichomanes obscurum Bl., Enum. Pl. Jav. 227. 1828.
PL 209: Near summit of Gunung Raya. Terrestrial, in deep shade, c. 900 m eleva-
tion. UKMB, duplicate at K.
CYATHEACEAE
Cyathea contaminans (Hook.) Copel., Phil. J. Sci. 4c: 60. 1909; Holttum 1963: 135.
PL 86: Kisap; Boon Siew Estate. Terrestrial, in light shade, near stream, c. 40 m
elevation, UKMB, duplicate at K.
Cyathea borneensis Copel., Phil. J. Sci. 6: 135. 1911; Holttum 1963: 110; 1968: 631.
PL 62: Near summit of Gunung Raya. Terrestrial, in deep shade, c. 900 m eleva-
tion. UKMB, duplicate at K.
PL 67: Gunung Raya Forest Reserve; track to summit. Terrestrial, in deep shade,
c. 700m elevation. UKMB.
Cyathea glabra (Bl.) Copel., Phil. Sci. 4c: 35. 1909; Holttum 1963: 120.
PL 93: Gunung Raya Forest Reserve; Lubuk Semilang, near pump-house. On
streambank in deep shade, c. 50 m elevation. UKMB, duplicate at K.
PL 263: Gunung Raya Forest Reserve; Sungai Kelian. Terrestrial, in deep shade,
c. 150 m elevation. UKMB, duplicate at K.
PL 165: Gunung Raya Forest Reserve; Lubuk Semilang. Terrestrial, in deep shade,
c. 40 m elevation. UKMB, 2 sheets.
POLYPODIACEAE
Platycerium coronarium (Koenig) Desv., Prodr. 213. 1827.
Rumah langsuyar.
PL 117: Kisap; Boon Siew Estate. Epiphyte on Hevea brasiliensis, in light shade,
c. 40 m elevation. UKMB.
Platycerium holttumii D. Jonch. & Hennipm., Brit. Fern Gaz. 10, 3. 116: 1970.
PL 79: Kisap; Boon Siew Estate: Epiphyte on Hevea brasiliensis, in light shade,
c. 40 m elevation. UKMB.
PL 176: Kisap; Boon Siew Estate. Epiphyte on Hevea brasiliensis, in light shade,
c. 40 m elevation. UKMB.
PL 150: Kisap; Boon Siew Estate (near Kedah Marble Factory). Epiphyte on Hevea
brasiliensis, in light shade, c. 20 m elevation. UKMB, 2 sheets.
Platycerium platylobum Aziz Bidin & Razali Jaman, Gard. Bull. 39(2). pl. 1 & 2. 1986.
PL 149: Kampong Padang Lunas. Epiphyte on Hevea brasiliensis, in light shade,
c. 30 m elevation. UKMB.
86 Gard. Bull. Sing. 40(2) (1987)
PL 124: Kisap; Boon Siew Estate. Epiphyte on Hevea brasiliensis, in light shade,
c. 50 m elevation. Endemic. UKMB.
Platycerium sp. (unidentified).
PL 11: Bukit Malut Forest Reserve; near proposed dam site. On fallen tree, in
light shade, c. 100 m elevation. Endemic. UKMB.
Pyrrosia adnascens (Sw.) Ching, Bull. Chin. Bot. Soc. 1: 45. 1935. (Hovenkamp (1986)
places both PR adnascens and P. varia as synonyms of P. lanceolata (L) Farwell).
Sakat batu; tetumpang.
PL 37: Bukit Malut Forest Reserve; near beach. Epiphyte, in light shade, c. 2 m
elevation. UKMB.
PL 169: Machinchang Forest Reserve; Tanjung Datai. Epiphyte, in light shade,
c. 5 m elevation. UKMB.
PL 287: Bumbun Besar Island. On rock, exposed, c. 3 m elevation. UKMB.
_ PL 154: Tanjung Rhu. On limestone, exposed, c. 50 m elevation. UKMB.
Pyrrosia varia (Kaulf.) Farewell, Am. Midl. Nat. 12: 302. 1931.
PL 255: Tanjung Dagu Forest Reserve; track to Tanjung Apau. On rock, in light
shade, c. 140 m elevation. UKMB.
Pyrrosia penangiana (Hook.) Holtt., Fl. Malaya II. 146-147, pl. 62. 1968.
Tanjung Rhu. On limestone by the sea. In light shade, c. 8 m elevation. Slight
record, not collected.
Pyrrosia longifolia (Burm.) Morton, Wash. Acad. Sci. 36: 168: 1946.
PL 85: Kisap; Boon Siew Estate. Epiphyte, in light shade, c. 30 m elevation.
UKMB, duplicate at K.
Pyrrosia stigmosa (Sw.) Ching, Bull. Chin. Bot. Soc. 1: 67. 1935.
PL 234: Dayang Bunting Island. On rock, exposed. c. 5 m elevation. UKMB.
Drymoglossum piloselloides (Linn.) Presl, Tent. Pterid. 227. 1836.
Duit-duit; sisik naga; sakat ribu-ribu.
PL 109: Kampong Belanga Pecah. Epiphyte, in light shade, c. 30 m elevation.
UKMB.
PL 237: Kisap; Boon Siew Estate. Epiphyte, in light shade, c. 30 m elevation.
UKMB.
Belvisia mucronata (Fée) Copel., Gen. Fil. 192. 1947.
PL 77: Near summit of Gunung Raya. Epiphyte, in light shade, c. 850 m eleva-
tion. UKMB, duplicate at K.
Loxogramme avenia (Bl.) Presl, Tent. Pterid. 215. 1836.
PL 267: Gunung Raya Forest Reserve; Genting Palas. On mossy rock, in light
shade, c. 450 m elevation. UKMB, duplicate at K.
Microsorium punctatum (Linn.) Copel, Univ. Cal. Publ. Bot. 16: 111. 1929.
PL 5: Bukit Malut Forest Reserve. Epiphyte on Pandanus, in light shade, c. 30
m elevation. UKMB.
PL 177: Kisap; Boon Siew Estate. Epiphyte on Hevea brasiliensis, in light shade;
c. 30 m elevation. UKMB, duplicate at K.
Fern Flora of Langkawi Islands 87
PL 190: Kisap; Kuala Kisap. Epiphyte on Hydrocarpus, in light shade, c. 5 m eleva-
tion. UKMB.
PL 245: Gunung Raya Forest Reserve; Lubuk Semilang. Epiphyte, in light shade,
c. 100 m elevation. UKMB, duplicate at K.
PL 235: Dayang Bunting Island, foot of Gua Langsir. On rock, in light shade,
c. 10 m elevation. UKMB.
Microsorium heterocarpum (Bl.) Ching, Bull. Fan. Mem. Inst. 4: 295. 1933.
PL 275: Gunung Raya Forest Reserve; Genting Palas. On rock, in deep shade,
c. 50 m elevation. UKMB.
Colysis acuminata (Bak.) Holtt., Fl. Malaya II. 162-163. 1968. var. angustata Holtt.
PL 161: Gunung Raya Forest Reserve; Lubuk Semilang. On rock, in stream bed,
in deep shade, c. 50 m elevation. UKMB.
PL 199: Gunung Raya Forest Reserve; Genting Palas. On rock, in stream bed,
in deep shade, c. 800 m elevation. UKMB, duplicate at K.
PL 240: Gunung Raya Forest Reserve; Lubuk Semilang. On rock, in light shade,
c. 90 m elevation. UKMB.
Colysis pedunculata (Hook. et. Grev.) Ching, Bull. Fan. Mem. Inst. 4: 321. 1933.
PL 15: Bukit Malut Forest Reserve. Epiphyte, in deep shade, c. 130 m elevation.
UKMB.
Drynaria sparsisora (Desv.) Moore, Ind. Fil. 348. 1862.
PL 36: Bukit Malut Forest Reserve. Epiphyte, exposed, c. 5 m elevation. UKMB.
PL 167: Kampung Sungai Taru; Klebang-Lubuk Semilang Road. Epiphyte, ex-
posed, c. 40 m elevation. UKMB.
PL 196: Lentang Jalan Island. On rock, exposed, c. 4 m elevation. UKMB.
PL 178: Kisap. Epiphyte, in light shade, c. 30 m elevation. UKMB.
PL 246: Gunung Raya Forest Reserve; Lubuk Semilang. Epiphyte, in light shade,
c. 100 m elevation. UKMB, duplicate at K.
Drynaria rigidula (Sw.) Beddome, Ferns Brit. Ind. t.314. 1869.
PL 84: Kisap; Boon Siew Estate. Epiphyte on Hevea brasiliensis, in light shade,
c. 30 m elevation. UKMB, duplicate at K.
PL 179: Kisap; Boon Siew Estate. Epiphyte on Hevea brasiliensis, in light shade,
c. 30 m elevation. UKMB, duplicate at K.
Drynaria bonii Chr., Not. Syst. 1: 186. 1910.
PL 205: Langgun Island. On rock, exposed, c. 20 m elevation. Rare. UKMB,
duplicate at K. Also collected by Chin (1977). KLU.
Pseudodrynaria coronans (Wall.) Ching, Sunyatsenia 5, 4: 262. 1940.
PL 58: Gunung Raya Forest Reserve. Epiphyte, in light shade, c. 750 m elevation.
UKMB, 2 sheets.
PL 212: Near summit of Gunung Raya. Epiphyte, in light shade, c. 800 m eleva-
tion. Rare. UKMB.
Phymatodes nigrescens (Bl.) J. Smith, Ferns Br. & For. 94. 1966.
Paku sumpah; paku ciai.
PL 102: Gunung Raya Forest Reserve; Durian Perangin Waterfall. On rock, ex-
posed, c. 130 m elevation. UKMB.
88 Gard. Bull. Sing. 40(2) (1987)
PL 244: Gunung Raya Forest Reserve; Lubuk Semilang. On rock, in light shade,
c. 100 m elevation. UKMB, duplicate at K.
Phymatodes scolopendria (Burm.) Ching, Contr. Inst. Bot. Nat. Acad. Peiping 2:
63, 1933.
PL 168: Machinchang Forest Reserve; Tanjung Datai. Terrestrial, exposed, c. 5
m elvation. UKMB.
Crypsinus trilobus (Houtt.) Copel., Gen. Fil. 206. 1947.
PL 74: Summit of Gunung Raya. Epiphyte, in light shade. UKMB.
Lecanopteris sinuosa Copel., Univ. Calif. Publ. Bot. 16: 123. 1929.
PL 110: Kampong Belanga Pecah. Epiphyte by roadside, exposed, c. 30 m eleva-
tion. UKMB, duplicate at K.
Goniophlebium persicifolium Bedd. 1870 (not in Handb.)
PL 71: Near summit of Gunung Raya, Epiphyte, in light shade, associated with
Vittaria and Drynaria, c. 900 m elevation. UKMB, duplicate at K.
PL 213: Summit of Gunung Raya. Epiphyte, in light shade, c. 960 m elevation.
UKMB, duplicate at K.
THELYPTERIDACEAE
Mesophlebion chlamydophorum (C. Chr.) Holttum, Blumea 22: 321: 1975; Holttum
198la: 384.
PL 280: Gunung Raya Forest Reserve; Genting Palas. Terrestrial, on stream bank,
in light shade, c. 550 m elevation. UKMB.
Cyclosorus interruptus (Willd.) H. Ito, Bot. Mag. Tokyo 51: 714: 1937; Holttum
198la: 386.
PL 256: Selat Panchor Forest Reserve; track to Tanjung Apau. Terrestrial, in light
shade, c. 160 m elevation. UKMB.
Sphaerostephanos heterocarpus (Bl.) Holttum in Nayar & Kaur, Compl. to Beddome:
280. 1974; Holttum 198la: 457.
PL 286: Guan Thong Estate. Terrestrial, exposed, c. 20 m elevation. UKMB,
duplicate at K.
Sphaerostephanos penniger (Hook.) Holttum in Nayar. & Kaur, Compl. to Beddome:
209. 1974; Holttum 1981la: 461.
PL 53: Gunung Raya Forest Reserve. Terrestrial, in light shade, c. 650 m eleva-
tion. UKMB, duplicate at K.
RJ 2101 Razali Jaman: Guan Thong Estate. Terrestrial, in light shade, c. 40 m
elevation. UKMB.
Pronephrium asperum (Presl.) Holttum, Blumea 20: 112. 1972; Holttum 1981la: 512.
PL 50: Gunung Raya Forest Reserve. Terrestrial, near track to summit, in light
shade, c. 650 m elevation. UKMB.
PL 2: Bukit Malut Forest Reserve. Terrestrial, in light shade, c. 120 m elevation.
UKMB.
PL 10]: Gunung Raya Forest Reserve; Durian Perangin Waterfall. Terrestrial, in
light shade, c. 50 m elevation. UKMB.
~
et a
Fern Flora of Langkawi Islands 89
PL 129: Track to Seven Wells. Terrestrial, in light shade, c. 90 m elevation. UKMB,
duplicate at K.
Pronephrium repandum (Fée) Holttum, Blumea 20: 109. 1972; Holttum 1981a: 533.
PL 136: Seven Wells. Terrestrial, on slope, in light shade, c. 200 m elevation.
UKMB, duplicate at K.
PL 262: Gunung Raya Forest Reserve: Sungai Kelian. Terrestrial, in light shade,
c. 150 m elevation. UKMB, duplicate at K.
Christella papilio (Hope) Holttum in Nayar & Kaur, Compl. to Beddome, 208. 1974;
Holttum 1981la: 556.
PL 69: Gunung Raya Forest Reserve. Terrestrial, in light shade, c. 600 m eleva-
tion. UKMB, duplicate at K.
Christella subpubescens (Bl.) Holttum, Webbia 30: 193. 1976; Holttum 1981la: 558.
PL 200: Gunung Raya Forest Reserve: Genting Palas. Terrestrial, on humus-rich
stream bank, in deep shade, c. 800 m elevation. UKMB, duplicate at K.
Christella parasitica (L.) Lev., Fl. de Kouy-teheou: 475. 1915; Holttum 1981a: 559.
PL 38: Guan Thong Estate. Terrestrial, in light shade, c. 20 m elevation. UKMB,
duplicate at K.
PL 10: Bukit Malut Forest Reserve. Terrestrial, in light shade, c. 100 m elevation.
UKMB.
PL 33: Bukit Malut Forest Reserve. Terrestrial, in light shade, c. 55 m elevation.
UKMB, duplicate at K.
PL 143: Machinchang Forest Reserve. Terrestrial, in light shade, c. 90 m eleva-
tion. UKMB.
Christella dentata (Forsk.) Brownsey & Jermy, Brit. Fern Gaz. 10: 338. 1973; Holt-
tum 198la: 557.
PL 17: Bukit Malut Forest Reserve. Terrestrial, on slope, in light shade, c. 450 m
elvation. UKMB, duplicate at K.
PL 160: Gunung Raya Forest Reserve; Lubuk Semilang. Terrestrial, in light shade,
c. 45 m elevation. UKMB.
Amphineuron terminans (Hook.) Holttum, Amer. Fern. 63: 82. 1973; Holttum
198la: 545.
PL 18: Bukit Malut Forest Reserve. Terrestrial, on slope, in light shade, c. 450 m
elevation. UKMB, duplicate at K.
PL 119: Dayang Bunting Island, track to the lake. Terrestrial, in light shade,
c. 5 m elevation. UKMB, duplicate at K.
PL 236: Kisap; Boon Siew Estate. Terrestrial, exposed, c. 40 m elevation. UKMB.
Amphineuron opulentum (Kaulf.) Holttum, Blumea 19: 45. 1971; Holttum 198la:
548.
PL 27: Bukit Malut Forest Reserve. Terrestrial, in light shade, c. 30 m elevation.
UKMB, duplicate at K.
PL 151: Gunung Raya Forest Reserve; Kampong Batu Asah. Terrestrial, in light
shade, c. 30 m elevation. UKMB.
Amphineuron immersum (Bl.) Holttum, in Nayar & Kaur, Compl. to Beddome: 203.
1974; Holttum 1981la: 547.
90 Gard. Bull. Sing. 40(2) (1987)
PL 22: Bukit Malut Forest Reserve. Terrestrial, in light shade, c. 120 m elevation.
UKMB.
PL 52: Near summit of Gunung Raya. Terrestrial, in deep shade, c. 950 m eleva-
tion. UKMB, duplicate at K.
DENNSTAEDTIACEAE
Dennstaedtioideae
Microlepia strigosa (Thb.) Pr., Epimel. Bot. 95. 1849.
PL 61: Near summit of Gunung Raya. Terrestrial, in light shade, c. 900 m eleva-
tion. UKMB, duplicate at K.
PL 215: Summit of Gunung Raya. Terrestrial, in deep shade, c. 950 m elevation.
UKMB, duplicate at K.
Microlepia speluncae (L.) Index XCIII. 1857. var. hancei.
PL 8: Bukit Malut Forest Reserve. Terrestrial, exposed, c. 100 m elevation. UKMB,
duplicate at K.
PL 87: Kisap; Boon Siew Estate. Terrestrial, on stream bank, in light shade,
c. 40 m elevation. UKMB, duplicate at K.
Microlepia speluncae (L.) Moore, Index XCIII. 1857 var. villosissima C. Chr. Gard.
Bull. Str. Settl. 4: 399 (1929).
PL 31: Bukit Malut Forest Reserve. Terrestrial, in light shade, c. 60 m elevation.
UKMB.
PL 144: Machinchang Forest Reserve. Terrestrial, exposed, c. 90 m elevation.
UKMB.
Lindsaeoideae
Lindsaea repens (Bory) Thwaites, En. Pl. Zeyl.: 388. 1864; Kramer 1971: 237.
H.C. Robinson. Langkawi. November 1916. K.
PL 158: Gunung Raya Forest Reserve; Lubuk Semilang. On rock near stream bank,
in deep shade, c. 40 m elevation. UKMB.
PL 278: Gunung Raya Forest Reserve; Genting Palas. Climbing in deep shade,
c. 520 m elevation. UKMB.
Lindsaea repens (Bory) Thwaites var. pectinata (Blume) Meet. ex Kuhn; Kramer 1971:
239.
PL 278: Gunung Raya Forest Reserve; Genting Palas. Epiphyte, in deep shade,
c. 520 m elevation. UKMB.
Lindsaea lucida Bl., Enum. Pl. Jav. 216. 1828; Kramer 1971: 233.
PL 277: Gunung Raya Forest Reserve; Genting Palas. On rock near stream bank,
in deep shade, c. 520 m elevation. UKMB.
Lindsaea ensifolia Sw. Schrad. Journ. 1800/2: 77. 1801; Kramer 1971: 211.
PL 89: Kampong Batu Asah. Terrestrial, in light shade, c. 35 m elevation. UKMB.
PL 132: Seven Wells. On rock, in light shade, c. 200 m elevation. UKMB.
Lindsaea heterophylla Dryand., Trans. Linn. Cos. 3; 41. pl. 8. f. 1, 1791; Kraan
O71 ZV:
i —-—-——
Fern Flora of Langkawi Islands 9]
PL 20: Bukit Malut Forest Reserve. Terrestrial, in light shade, c. 200 m elevation.
UKMB.
PL 141: Machinchang Forest Reseve. Terrestrial, in light shade, c. 200 m eleva-
tion. UKMB. .
PL 187: Kisap; Kuala Kisap. Terrestrial, in light shade, c. 30 m elevation. UKMB.
Lindsaea parasitica (Roxburgh ex Griffith) Hieron, Hedwigia: 62. 14. 1920; Kramer
1971: 244.
PL 204: Near summit of Gunung Raya. Epiphyte, in light shade, c. 900 m eleva-
tion. UKMB, duplicate at K.
Tapeinidium pinnatum (Cav.) C.Chr., Ind. Fil. 213. 1905; Kramer 1971: 191.
PL 210: Near summit of Gunung Raya. Terrestrial, in light shade, c. 950 m eleva-
tion. UKMB.
Davallioideae
Davallia denticulata (Burm.) Mett., Kuhn, Fil. Dec. 27. 1867.
Paku tertutup.
PL 73: Gunung Raya Forest Reserve. Epiphyte, in light shade, c. 650 m elevation.
UKMB.
PL 156: Bukit Sawak Forest Reserve. Epiphyte, in light shade, c. 150 m elevation.
UKMB.
PL 174: Padang Lalang. Epiphyte, exposed, c. 5 m elevation. UKMB, duplicate
at K.
PL 197: Lentang Jalan Island. On rock, in light shade, c. 6 m elevation. UKMB,
duplicate at K.
PL 232: Dayang Bunting Island. On rock, by the beach, exposed. UKMB, duplicate
at K.
Davallia trichomanoides Bl., Enum. Pl. Jav. 238. 1828.
PL 285: Track to summit of Gunung Raya. Epiphyte, in deep shade, c. 600 m
elevation. UKMB.
Davallia trichomanoides Bl. var lorrainii (Hance) Holtt.
PL 216: Summit of Gunung Raya. Epiphyte, in deep shade, c. 960 m elevation.
UKMB, duplicate at K.
Davallia divaricata Bl., Enum. Pl. Jav. 237. 1828.
PL 54: Gunung Raya Forest Reserve. Epiphyte, in light shade, c. 660 m elevation.
UKMB, duplicate at K.
Humata pectinata (Sm.) Desv., Prodr. 323. 1827.
Tanjung Rhu. On limesone cliff by the beach, in light shade, c. 8 m elevation.
Sight record.
Humata vestita (Bl.) Moore, Index XCIII. 1857.
PL 131: Seven Wells. On rock in light shade, c. 200 m elevation. UKMB, duplicate
at K.
Humata repens (L.fil.) Diels, Nat. Pflanzenfam. 1/4: 209. 1899.
PL 75: Track to summit of Gunung Raya. Epiphyte, in deep shade, c. 900 m eleva-
tion. UKMB.
92 Gard. Bull. Sing. 40(2) (1987)
Oleandroideae
Nephrolepis biserrata (Sw.) Schott, Gen. Fil. ad t. 3. 1834.
PL 116: Kedawang; Kampong Lubuk Buaya. Terrestrial, exposed, c. 5 m eleva-
tion. UKMB.
PL 120: Singa Besar Island. Terrestrial, exposed, c. 5 m elevation. UKMB.
Nephrolepis biserrata (Sw.) Schott var. exaltata.
Kuah. Ornamental plant. Sight record.
Nephrolepis hirsutula (Forst.) Pr., Tent. Pterid. 79. 1836.
PL 184: Kisap; Kuala Kisap. Terrestrial, exposed, c. 30 m elevation. UKMB.
Oleandra pistillaris (Sw.) C. Chr., Ind. Fil., Suppl. III: 132. 1834.
PL 214: Near summit of Gunung Raya. Terrestrial, in light shade, c. 950 m eleva-
tion. UKMB.
Pteridioideae
Pteris vittata Linn., Spec. Pl. 2: 1074. 1753.
PL 83: Kisap; Boon Siew Estate. Terrestrial, in light shade, c. 30 m elevation.
UKMB.
PL 153: Kisap. Terrestrial, in light shade, c. 40 m elevation. UKMB.
Pteris ensiformis Burm., Fl. Ind. 230. 1768.
Mukut; paku lemukut.
PL 47: Kisap; Boon Siew Estate. Terrestrial, in light shade, c. 20 m elevation.
UKMB.
PL 107: Gunung Raya Forest Reserve; Durian Perangin Waterfall. Terrestrial, in
light shade, c. 100 m elevation. UKMB.
PL 258: Selat Panchor Forest Reserve; track to Tanjung Apau. Terrestrial, in light
shade, c. 110 m elevation. UKMB.
Pteris scabripes Wall. apud. Hook., Spec. Fil. 2: 165. 1858.
No. 3381. C. Curtis. Foot of Gunung Raya. Terrestrial, April 1896. K.
No. 15789. M.R. Henderson. Teluk Barau. Terrestrial. April 1911. K.
PL 219: Bukit Sawak Forest Reserve. Terrestrial, in light shade, c. 80 m elevation.
UKMB.
PL 260: Gunung Raya Forest Reserve: Sungai Kelian. Terrestrial, exposed,
c. 55 m elevation. UKMB.
Pteris venulosa Bl., Enum. Pl. Jav. 209. 1828.
PL 127: Seven Wells. Terrestrial, in deep shade, c. 90 m elevation. UKMB, duplicate
at K.
Pteris biaurita Linn., Spec. Pl. 2: 1076. 1753.
No. 6361: H.C. Robinson. Terutau. November 1916. K.
No. 29178: R.E. Holttum. Kisap. 30 November 1931. K.
PL 41: Guan Thong Estate. Terrestrial, in light shade, c. 20 m elevation. UKMB.
PL 88: Kisap; Boon Siew Estate. Terrestrial, in light shade, c. 40 m elevation.
UKMB, duplicate at K.
UL ——-
Fern Flora of Langkawi Islands 93
PL 142: Machinchang Forest Reserve. Terrestrial, in light shade, c. 90 m eleva-
tion. UKMB, duplicate at K.
Pteris mertensioides Willd., Sp. Pl. 5:394. 1810.
PL 23: Bukit Malut Forest Reserve. Terrestrial, in light shade, c. 120 m elevation.
UKMB.
Pteris longipinnula Wall., Cat. No. 108 (nom. nud.); Agardh, Recen. Pterid. 19. 1839.
PL 48: Gunung Raya Forest Reserve. Terrestrial, exposed, c. 500 m elevation.
UKMB, duplicate at K.
Acrostichum aureum Linn., Spec. Pl. 2: 1069. 73. 1753.
Larat; piai lasa; piai raja.
PL 112: Kuala Temoyong. In mangrove swamp, c. 0 m elevation. UKMB, duplicate
at K.
Acrostichum speciosum Willd., Spec. Pl. 5: 117. 1810.
PL Ill: Kuala Temoyong. In mangrove swamp, by the beach. UKMB, duplicate
at K.
PL 121: Singa Besar Island. In mangrove swamp, by the beach. UKMB.
Stenochlaena palustris (Burm.) Bedd., Ferns Brit. Ind. Suppl. 26. 1876.
PL 14: Bukit Malut Forest Reserve. Climbing, in light shade, c. 100 m elevation.
UKMB.
Asplenioideae
Asplenium nidus Linn., Spec. Pl. 2: 1079. 1753.
Daun semum; paku langsuyar.
PL 108: Gunung Raya Forest Reserve; Durian Perangin Waterfall. Epiphyte, in
light shade, c. 140 m elevation. UKMB.
Asplenium phyllitidis Don, Prodr. Fl. Nep. 7. 1825.
PL 13: Bukit Malut Forest Reserve. Epiphyte, in light shade, c. 100 m elevation.
UKMB.
PL 163: Gunung Raya Forest Reserve; Lubuk Semilang. Epiphyte on Knema
laurina, in light shade, c. 5O m elevation. UKMB.
PL 173: Machinchang Forest Reserve; Teluk Datai. Epiphyte, in light shade,
c. 5 m elevation. UKMB.
Asplenium phyllitidis Don subsp. malesicum Holttum; Gard. Bull. Sing. 27: 153. 1974.
PL 70: Near summit of Gunung Raya. Epiphyte, in light shade, c. 870 m eleva-
tion. UKMB, duplicate at K.
Asplenium salignum Bl., Enum. Pl. Jav. 175. 1828.
PL 266: Gunung Raya Forest Reserve; Genting Palas. On rock, in deep shade,
c. 450 m elevation. UKMB, duplicate at K.
PL 270: Gunung Raja Forest Reserve; Genting Palas. On rock, in deep shade,
c. 450 m elevation. UKMB, duplicate at K.
Asplenium spathulinum J. Smith, J. Bot. 3: 408. 1813.
PL 55: Gunung Raya Forest Reserve. Epiphyte, in deep shade, c. 720 m eleva-
tion. UKMB.
94 Gard. Bull. Sing. 40(2) (1987)
PL 57: Near track to summit of Gunung Raya. Epiphyte, in deep shade, c. 750 m
elevation. UKMB.
PL 254: Tanjung Dagu Forest Reserve. Epiphyte, associated with A. nidus, in deep
shade, c. 90 m elevation. UKMB, duplicate at K.
Asplenium pellucidum Lam., Encyl. 2: 305. 1786.
PL 241]: Gunung Raya Forest Reserve; Lubuk Semilang. Epiphyte, in deep shade,
c. 95 m elevation. UKMB, duplicate at K.
Asplenium macrophyllum Sw., Schrad. Journ. 1800/2: 52. 1801.
PL 182: Gunung Raya Forest Reserve; Lubuk Semilang. On rock, in light shade,
c. 500 m elevation. UKMB.
PL 233: Dayang Bunting Island. On rock, in light shade, c. 2 m elevation. UKMB.
PL 247: Gunung Raya Forest Reserve; Lubuk Semilang. On rock, in light shade,
c. 190 m elevation. UKMB. 2 sheets.
PL 269: Gunung Raja Forest Reserve; Genting Palas. On rock, in deep shade,
c. 450 m elevation. UKMB, duplicate at K.
Asplenium tenerum Forster, Prodr. 80, 1786.
PL 68: Near summit of Gunung Raya. Epiphyte, in deep shade, c. 900 m eleva-
tion. UKMB, duplicate at K.
Asplenium paradoxum Bl., Enum. Pl. Jav. 179. 1828.
No. 15486. H.N. Ridley. Gunung Raya. February 1911. K.
Blechnoideae
Blechnum orientale Linn., Spec. Pl. 2: 1077. 1753.
Paku ikan.
PL 7: Bukit Malut Forest Reserve. Terrestrial, on exposed earth bank, c. 100 m
elevation. UKMB, duplicate at K.
Lomariopsidoideae
Lomariopsis lineata (Presl.) Holttum Novit. Bot. Inst. Prag. 1968 (1969) 9; Holttum
1978: 262.
PL 159: Gunung Raya Forest Reserve; Lubuk Semilang. On rock, in deep shade,
c. 40 m elevation. UKMB.
Elaphoglossum callifolium (Bl.) Moore, Ind. Fil. (1857) 7.
PL 76: Near summit of Gunung Raya. Epiphyte, in deep shade, c. 900 m eleva-
tion. UKMB.
Bolbitis heteroclita (Pr.) Ching, in C. Chr., Ind. Fil. Suppl. III, 48. 1934.
PL 250: Selat Panchor Forest Reserve. Terrestrial, in deep shade, c. 50 m eleva-
tion. UKMB, duplicate at K.
PL 51: Gunung Raya Forest Reserve, near track to summit. Climbing, in deep
shade, c. 600 m elevation. UKMB.
PL 66: Gunung Raya Forest Reserve, near track to summit. Terrestrial, in deep
shade, c. 650 m elevation. UKMB.
PL 251: Selat Panchor Forest Reserve, track to Tanjung Apau. Climbing in deep
shade, c. 5O m elevation. UKMB.
Fern Flora of Langkawi Islands 95
PL 265: Gunung Raya Forest Reserve; Sungai Kelian. On rock, in deep shade,
c. 50 m elevation. UKMB.
PL 272: Gunung Raya Forest Reserve; Genting Palas. On rock, in deep shade,
c. 460 m elevation. UKMB.
Bolbitis malaccensis (C.Chr.) Ching, in C. Chr., Ind. Fil. Suppl. III, 49. 1934.
Langkawi. R.E. Holttum (1968). Rare.
Bolbitis appendiculata (Willd.) Iwatsuki, Acta Phytotax. Geobot. 18: 48. 1959.
(Egenolfia appendiculata in Holttum 1968: 459.).
PL 12: Bukit Malut Forest Reserve. On rock, in light shade, c. 100 m elevation.
UKMB.
PL 95: Gunung Raya Forest Reserve; Lubuk Semilang. On rock, in light shade,
c. 50 m elevation. UKMB.
PL 118: Dayang Bunting Island. Terrestrial, by trackside to the lake, in light shade,
c. 5 m elevation. UKMB.
PL 130: Path to Seven Wells. On rock, in light shade, c. 110 m elevation. UKMB.
PL 140: Machinchang Forest Reserve. On rock, in deep shade, c. 60 m elevation.
UKMB, duplicate at K.
Bolbitis virens (Wall.) Schott, Gen. Fil. at t. 14. 1834.
PL 220: Bukit Sawak Forest Reserve. On rock, in deep shade, 0.50 m elevation.
UKMB.
PL 276: Gunung Raya Forest Reserve; Genting Palas. Terrestrial, in deep shade,
c. 500 m elevation. UKMB. 2 sheets.
Dryopteridoideae
Polystichum prolificans V.A.V.R., Bull. Jard. Bot. Buitenz., III ser., 2: 170. 1920.
PL 72: Near summit of Gunung Raya. Terrestrial, in deep shade, c. 900 m eleva-
tion. UKMB, duplicate at K.
Tectarioideae
Tectaria brachiata (Zoll. & Mor.) Morton, Contr. U.S. Nat. Herb. 38. 217. 1973;
Holttum 1981b: 137, excl. Aspidium variolosum and its synonyms.
PL 24: Bukit Malut Forest Reserve. Terrestrial, in light shade, c. 120 m elevation.
UKMB.
PL 34: Bukit Malut Forest Reserve. Terrestrial, in light shade, c. 10 m elevation.
UKMB, duplicate at K.
PL 45: Kisap; Boon Siew Estate. Terrestrial, in light shade, c. 20 m elevation.
UKMB, duplicate at K.
PL 97: Ulu Melaka; Kampong Bukit Hantu. Terrestrial, in light shade, c. 60 m
elevation. UKMB, duplicate at K.
Tectaria variolosa (Hook.) C. Chr., Contr. U.S. Nat. Herb. 26 (1931) 289.
PL 222: Timun Island, Tanjung Timun. In rock crevices, exposed to sunlight and
sea-sprays, c. 6 m elevation. UKMB, duplicate at K. Rare.
Tectaria semipinnata (Roxb.) Morton, Contr. U.S. Nat. Herb. 38 (1974) 286.
(7. maingayi in Holttum 1981b: 513).
PL 49: Gunung Raya Forest Reserve. Terrestrial, in light shade, c. 650 m eleva-
tion. UKMB, duplicate at K.
96 Gard. Bull. Sing. 40(2) (1987)
PL 100: Gunung Raya Forest Reserve; Durian Perangin Waterfall. Terrestrial, in
light shade, c. 50 m elevation. UKMB, duplicate at K.
Tectaria angulata (Willd.) Copel., Sarawak Mus. J. 2: 370. 1917.
PL 164: Gunung Raya Forest Reserve; Lubuk Semilang, near pumphouse. Ter-
restrial, in light shade, c. 40 m elevation. UKMB.
PL 123: Singa Besar Island. Terrestrial, in light shade, c. 7 m elevation. UKMB.
Tectaria rumicifolia (Ridley) C. Chr. in Holttum 1968: 519, 636; 7: oligophylla
(Rosenst.) C. Chr.
PL 137: Machinchang Forest Reserve. Terrestrial, on slope, in deep shade,
c. 240 m elevation. UKMB, duplicate at K.
Cyclopeltis crenata (Fée) C. Chr., Ind. Fil. Suppl. 64. 1934.
PL 253: Tanjung Dagu Forest Reserve; track to Tanjung Apau. Terrestrial, in
deep shade, c. 60 m elevation. UKMB, duplicate at K.
Arcypteris irregularis (Pr.) Holtt., Reinwardtia 1: 193. 1951.
PL 82: Kisap; Boon Siew Estate, track to Gunung Raya. Terrestrial, in light shade,
c. 30 m elevation. UKMB, duplicate at K.
Ctenitis mannii (Hope) Ching, Bull. Fan. Mem. Inst. Bot. 8: 289. 1938.
PL 201: Gunung Raya Forest Reserve; Genting Palas. On rock, in light shade,
c. 800 m elevation. UKMB, duplicate at K.
Ctenitis subobscura (Christ) Holttum, Fern Gaz. 12: 320 (1934); Blumea 31: 23 (1985).
No. 15548. Md. Haniff. Gunung Raya. 1911. K.
Athyrioideae
Diplazium esculentum (Retz) Sw., Schrad. J. Bot. 1801/2: 312. 1803; Holttum 1968,
App. II: 637.
Paku besar.
PL 125: Kuah; Kampong Padang Lalung. Terrestrial, in swamp, exposed, c. 50 m
elevation. UKMB.
Diplazium malaccense Pr., Epim. Bot. 86. 1849; Holttum 1968, App. II: 637.
PL 207: Near summit of Gunung Raya. Terrestrial, in light shade, c. 950 m eleva-
tion. UKMB, duplicate at K.
Diplazium bantamense Bl., Enum. Pl. Jav. 191. 1828; Holttum 1968, App. II: 637.
PL 60: Track to summit of Gunung Raya. Terrestrial, in deep shade, c. 900 m
elevation. UKMB.
ADIANTACEAE
Ceratopteris thalictroides (Linn.) Brongn., Bull. Soc. Philon. 1821: 86.
Paku dodok.
PL 14: Kampong Kubang Badak. In rice-field, c. 5 m elevation. UKMB.
PL 90: Kampong Batu Asah. On stream-bed, in light shade, c. 30 m elevation.
UKMB.
PL 114: Kampong Lubuk Buaya. In mud, exposed, c. 5 m elevation. UKMB,
duplicate at K.
Fern Flora of Langkawi Islands 97
PL 147: Kampong Kubang Badak. In rice-field, c. 5 m elevation. UKMB.
PL 224: Kisap; Boon Siew Estate. In mud, exposed, c. 30 m elevation. UKMB.
Taenitis blechnoides (Willd.) Sw., Syn. Fil. 24, 220. 1806.
Paku bulu.
PL 3: Bukit Malut Forest Reserve. Terrestrial, in light shade, c. 100 m elevation.
UKMB.
PL 4: Bukit Malut Forest Reserve: Terrestrial, in light shade, c. 100 m elevation.
UKMB.
PL 135: Seven Wells. Terrestrial, on stream bank, in light shade, c. 200 m eleva-
tion. UKMB.
PL 261]: Gunung Raya Forest Reserve; Genting Palas. Terrestrial, in light shade,
c. 100 m elevation. UKMB.
PL 283: Gunung Raya Forest Reserve; Genting Palas. Terrestrial, in light shade,
c. 550 m elevation. UKMB.
No. 15786: M.R. Henderson. Teluk Barau. April 1911. K.
Cheilanthes tenuifolia (Burm.) Sw., Syn. Fil. 129, 332. 1806.
Resam padi; temangah; telur belangkas.
No. 2039. Abdul Samat Abdullah. Kuah. KLU, duplicate at K.
PL 43: Kisap; Boon Siew Estate. Terrestrial, in light shade, c. 20 m elevation.
UKMB, duplicate at K.
PL 138: Kampong Kubang Badak. Terrestrial, exposed, c. 40 m elevation. UKMB.
Pityrogramma calomelanos (L.) Link, Handb. Gew: 3:20. 1833.
Paku hijau.
PL 9: Bukit Malut Forest Reserve. Terrestrial, exposed, c. 100 m elevation. UKMB.
Doryopteris ludens (Wall.) J. Sm., Hist. Fil. 289. 1875.
H.C. Robinson. Dayang Bunting Island. November 1916. K.
No. 29066. M.R. Henderson. Selat Panchor, c. 50 m elevation. November 1934. K.
PL 46: Kisap; Boon Siew Estate. Terrestrial, in light shade, c. 20 m elevation.
UKMB.
PL 22]: Timun Island. In rock crevices, c. 1 m elevation. UKMB.
Hemionitis arifolia (Burm.) Moore, Ind. Fil. 114. 1859.
PL 32: Bukit Sawak Forest Reserve. On rock, in light shade, c. 50 m elevation.
UKMB.
PL 46: Kisap; Boon Siew Estate. Terrestrial, in light shade, c. 20 m elevation.
UKMB, duplicate at K.
PL 225: Kisap; Boon Siew Estate, near Kedah Marble Factory. Terrestrial, in light
shade, c. 30 m elevation. UKMB.
Adiantum stenochylamys Bak., Am. Bot. 5: 29. 1891.
No. 15788. M.R. Henderson. Teluk Barau. April 1911. K.
Timun Island. On rock, exposed, c. 6 m elevation. Not collected. Sight record.
Adiantum caudatum Linn., Mantissa 308. 1771.
PL 230: Dayang Bunting Island. Terrestrial, in light shade, c. 10 m elevation.
UKMB, duplicate at K.
98 Gard. Bull. Sing. 40(2) (1987)
Adiantum philippense Linn., Spec. Pl. 2: 1094. 1753.
Paku sisik; paku mega.
No. 29074. M.R. Henderson. Selat Panchor, c. 100 m elevation. 23 November
1934. K.
PL 44: Kisap; Boon Siew Estate. Terrestrial, in light shade, c. 70 m elevation.
UKMB; duplicate at K.
PL 229: Timun Island; Tanjung Timun. In rock crevices, exposed, c. 6 m eleva-
tion. UKMB, duplicate at K.
PL 231: Dayang Bunting Island. Terrestrial, in light shade, c. 10 m elevation.
UKMB, duplicate at K.
Adiantum capillus-veneris Linn., Spec. Pl. 2: 1096. 1753.
Kuah. Ornamental plant. Sight record.
Adiantum mathewsianum Hook., Sp. Fil. 2: 35. 1858.
Kuah. Ornamental plant. Sight record.
‘Adiantum peruvianum K\l., Linnaea 18: 555. 1845.
Kuah. Ornamental plant. Sight record.
Adiantum trapeziforme Linn., Spec. Pl. 2: 1097. 1753.
Kuah. Ornamental plant. Sight record.
Adiantum tenerum Swartz., Prodromus 135. 1788.
Kuah. Ornamental plant. Sight record.
Adiantum latifolium Lam, Encycl. 1: 43. 1783.
PL 42: Kisap; Boon Siew Estate. Terrestrial, in light shade, c. 20 m elevation.
UKMB.
PL 81: Kisap; Boon Siew Estate. Terrestrial, in light shade, c. 30 m elevation.
UKMB.
Adiantum polyphyllum Willd., Sp. Pl. 5: 454. 1828.
Kuah. Ornamental plant. Sight record.
Antrophyum callifolium Bl., Enum. Pl. Jav. III. 1828.
No. 6359. H.C. Robinson. Terutau. April 1911. K.
PL 16: Bukit Malut Forest Reserve. On rock, in light shade, c. 500 m elevation.
UKMB, duplicate at K.
PL 103: Gunung Raya Forest Reserve; Durian Perangin Waterfall. On rock, in
deep shade, c. 100 m elevation. UKMB.
PL 242: Gunung Raya Forest Reserve; Lubuk Semilang. On rock, in light shade,
c. 100 m elevation. UKMB, duplicate at K.
PL 274: Gunung Raya Forest Reserve; Genting Palas. On rock, in deep shade,
c. 500 m elevation, UKMB.
Vittaria ensiformis Sw., Ges. Nat. Fr. Berl. Neu. Schr. 2: 134, t. 7. f. 1. 1799.
No. 15831. H.C. Robinson. Pulau Adang: April 1911. K.
PL 106: Gunung Raya Forest Reserve; Durian Perangin Waterfall. Epiphyte, ex-
posed, c. 100 m elevation. UKMB, duplicate at K.
PL 139: Machinchang Forest Reserve. Epiphyte on Pentacalyx, in light shade,
c. 50 m elevation. UKMB.
PL 172: Machinchang Forest Reserve; Teluk Datai. Epiphyte, in light shade,
c. 5 m elevation. UKMB, duplicate at K.
99 Fern Flora of Langkawi Islands
Vittaria ensiformis Sw. var. latifolia Holttum.
PL 56: Gunung Raya Forest Reserve. Epiphyte, in light shade, c. 720 m elevation.
UKMB. 2 sheets.
PL 193: Bumbun Besar Island. On rock, exposed, c. 3 m elevation. UKMB.
Vittaria angustifolia Bl., Enum. Pl. Jav. 199. 1828.
PL 249: Gunung Raya Forest Reserve; Lubuk Semilang. Epiphyte, in light shade,
c. 90 m elevation. UKMB.
MARSILEACEAE
Marsilea crenata Presl, Rel-Haenk. 1: 84, t. 12, f. 13. 1825.
PL 98: Hulu Melaka; Kampong Bukit Hantu. In rice-field, exposed, c. 50 m eleva-
tion. UKMB, duplicate at K.
PL 99: Hulu Melaka; Kampong Bukit Hantu. In rice-field, exposed, c. 50 m eleva-
tion. UKMB, duplicate at K.
PL 146: Kampong Kubang Badak. In rice-field, exposed, c. 5 m elevation. UKMB.
AZOLLACEAE
Azolla pinnata R. Br., Prodr. Fl. N. Holl. 167. 1810.
Hulu Melaka; Kampong Bukit Hantu. In rice-field, exposed, c. 5 m elevatior
Sight record.
The breakdown of the fern flora is as shown in Table 1 below:
Table 1
Breakdown of the Langkawian Fern Flora.
Families/Subfamilies Genera No. of species (incl. subspecies,
varieties and hybrids)
OPHIOGLOSSACEAE Helminthostachys
MARATTIACEAE Angiopteris
SCHIZAEACEAE Schizaea
Lygodium
GLEICHENIACEAE Dicranopteris
HYMENOPHYLLACEAE Hymenophyllum
Trichomanes
CYATHEACEAE Cyathea
POLYPODIACEAE Platycerium
Pyrrosia
Drymoglossum
Belvisia
Loxogramme
Microsorium
Colysis
Drynaria
Pseudodrynaria
Phymatodes
Crypsinus
Lecanopteris
Goniophlebium
nme WOrR NAN —
100
Table 1 (Contd)
Families/Subfamilies
THELYPTERIDACEAE
DENNSTAEDTIACEAE
Dennstaedtioideae
Lindsaeoideae
Davallioideae
_Oleandroideae
Pteridioideae
Asplenioideae
Blechnoideae
Lomariopsidoideae
Dryopteridoideae
Tectarioideae
Athyrioideae
ADIANTACEAE
MARSILEACEAE
AZOLLACEAE
Genera
Mesophlebion
Cyclosorus
Sphaerostephanos
Pronephrium
Christella
Amphineuron
Microlepia
Lindsaea
Tapeinidium
Davallia
Humata
Nephrolepis
Oleandra
Pteris
Acrostichum
Stenochlaena
Asplenium
Blechnum
Lomariopsis
Elaphoglossum
Bolbitis
Polystichum
Tectaria
Cyclopeltis
Arcypteris
Ctenitis
Diplazium
Ceratopteris
Taenitis
Cheilanthes
Pityrogramma
Doryopteris
Hemionitis
Adiantum
Antrophyum
Vittaria
Marsilea
Azolla
Total number: 59
Conclusion
Gard. Bull. Sing. 40(2) (1987)
No. of species (incl. subspecies,
varieties and hybrids)
wWBRNN-
—
Total number: 145
The Langkawian fern flora consists of some 134 species, 2 subspecies, 8 varieties
and 1 hybrid representing about 29% of the total number of species found in Penin-
sular Malaysia, and comprising diverse elements of Malesia, Indochina, India, and
Tropical America (introduced).
The introduced elements (some escaped from cultivation and successfully form-
ing part of the natural vegetation, e.g. Adiantum latifolium, number only 7 (six of
Fern Flora of Langkawi Islands 101
them belong to the genus Adiantum) or 4.8% of the total fern flora of Langkawi.
Some of the species which have their centre of distribution in mainland Asia have
their southern limit in Langkawi as exemplifed by Platycerium holttumii,
Pseudodrynaria coronans and Tectaria variolosa. Ferns which may be absent from
mainland Malaysia occur in Langkawi due to the preference for a seasonal climate,
e.g. Bolbitis malaccensis and Adiantum philippense. It is apparent from the number
of species collected and recorded that Langkawi is rich in its fern flora — thanks
to its favourable climate and the varied habitats available.
The stunning beauty of the islands and the cool, refreshing and pollution-free air
and sea in Langkawi, have encouraged the authorities to put forward a plan to develop
a massive resort city on the north-east portion of the main island, covering c. 566
hectares, centred around a crescent-shaped beach near Tanjung Rhu, as well as at
a few other places in this group of islands. The construction of roads, airport, hotels,
etc., have already started. The after-effects, such as erosion and landslips, are in-
deed posing a real threat to the indigenous vegetation not only in terms of area deple-
tion but also extensive degradation and invasion of exotic plants. With these activities,
it is becoming apparent that the flora and fauna of Langkawi urgently require in-
vestigations before the primary character becomes drastically modified. As survival
depends upon preservation of their habitats, conservation of the existing nature
reserves is of the utmost importance.
Acknowledgements
The author is grateful to Prof. R.E. Holttum for his guidance in the identifica-
tion of some of the specimens, to the Keeper and Curator of the Herbaria at Royal
Botanic Gardens Kew and University of Malaya respectively for the use of materials
and to the staff of the Office of Forest, Langkawi for allowing the author to carry
out a survey of the ferns of Langkawi Islands. The field work was supported by
Universiti Kebangsaan Malaysia, Research Grant No. 123/85. The author also wishes
to thank Encik-Encik Razali Jaman, Ahmad Zainuddin and A. Hamid for their com-
petent assistance in the field.
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Malaysia. Gard. Bull. Sing. 39(2): 149-151.
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Chin, S.C. (1977). The limestone hill flora of Malaya I. Gard. Bull. Sing. 30: 165-219.
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Br. Roy. As. Soc. 17: 13-87.
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1: 1-61.
Holttum, R.E. (1963). Cyatheaceae. Fi. Malesiana Ser. II. Vol. 1: 65-176.
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MSMaA for Controlling Cyperus kyllingia, Axonopus
compressus and Brachiara distachya in
Tifgreen Bermuda Grass Turf
LEONG CHEE CHIEW *
Parks & Recreation Department
Botanic Gardens, Singapore 1025
Abstract
Several herbicides were tested for selective control of grass and broadleaf weeds in Tifgreen Bermuda
grass turf in golf courses. MSMA (monosodium methanearsonate) was found to selectively control
Cyperus kyllingia, Axonopus compressus and Brachiara distachya, three of the most troublesome weeds
in the golf courses. Metribuzin selectively controlled Euphorbia thymifolia.
Introduction
The high frequency of mowing, together with the low cutting height in golf courses
should be expected to keep many species of weeds under control. However, there
are those weeds which can thrive even under close mowing because they have leaf
forms that lie under the cutting blade. These if left unchecked can intermingle and
eventually take over the sward from the finer grasses (Ellis 1986). At a point where
manual weeding cannot keep up with spread of such hardy weeds, use of selective
herbicides may be the alternative.
In the use of herbicides various factors need consideration, perhaps the most im-
portant being the control of weeds without excessive damage to the turf. Johnson
(1983) reported that some herbicides like napropamide and prosulfalin reduced root
growth of Bermuda grass. Ability to resist herbicide damage was found to differ
among turfgrasses eg. among the triploid hybrid Bermuda grasses, Tifway hybrids
exceed the Tifgreen hybrids in tolerance to 2,4-D (Hanna 1986).
Examples of chemical weed control in Bermuda grass include the use of diclofop
for controlling goosegrass (Eleusine indica) in Tifdwarf putting greens (Murdock
and Nishimoto 1982). Batten (1984) reported the control of goosegrass by post-
emergence applications of MSMA or metribuzin. Sedges such as the purple nutsedge
were reported to be controlled post-emergence on the basis of a continual reduction
of the leaves and plant with repeated MSMA or bentazon applications; as new shoots
recur from rhizomes and tubers, another herbicide application will reduce them.
MSMA, especially effective in controlling established Johnsongrass, was applied
repeatedly between three to eight times per growing season in order to achieve effec-
tive control (Anderson 1977).
*The author is currently with the Strategic Planning Division of the Ministry of National Development,
Singapore.
103
104 Gard. Bull. Sing. 40(2) (1987)
Weeds invading Bermuda grasses used in the Serapong Golf Course of Sentosa
and Tanah Merah Golf Course were described by Wong (1986). This paper reports
the experiments carried out to control some of these weeds in the Tifgreen Bermuda
grass fairways and driving range of the Golf Courses.
Materials and Methods
Apart from herbicides which were already available to the experimentor, 12 locally-
based agrochemical companies were also requested to recommend selective herbicides
for use in Bermuda grass swards. Response was limited but a total of five herbicides
were finally chosen for the trial. They were metribuzin (Sencor WP 70), dicamba
(Fez PE 400), bentazon (Basagran 39.6%), 2,4-D (2,4-D amine 720), and MSMA
(MSMA 500).
Trials on Tifgreen Bermuda grass and weeds were conducted separately to deter-
mine the effects of the various herbicides on each. Trials on Tifgreen Bermuda grass
were conducted in relatively weed free areas in a fairway at the Serapong Golf Course
and those on weeds in the Tanah Merah Golf Course driving range. The weeds
initially monitored were Axonopus compressus, Cyperus kyllingia, and Euphorbia
thymifolia. Brachiara distachya was included at a later stage.
The randomised complete block design was used for each trial. The five herbicides
were each tested at 4 levels of concentration. They were:- metribuzin (Sencor WP
70) — 0, 0.2, 0.6 and 1.2 g/1; dicamba (Fez Pe 400) — 0, 1.0, 2.0 and 3.0 ml/];
bentazon (Basagran 39.6%) — 0, 2.0, 6.0 and 10.0 ml/1; 2,4-D (2,4-D amine 720)
— 0, 2.0, 4.0 and 8.0 ml/]; MSMA (MSMA 500) — 0, 2.0, 6.0 and 10.0 ml/I. In
each trial, 3 blocks containing 20 treatment plots each were constructed; each treat-
ment plot was 0.4 m by 0.4 m in size. Treatments were completely randomised within
each block.
Application of herbicide was carried out using Gardena pressure sprayers. Each
treatment plot was sprayed with 100 ml herbicide with teepol added as wetting agent.
Effects of herbicides on turf and weeds were monitored after herbicide applica-
tion by recording visual ratings of phytotoxicity, adopting the method used by Mur-
doch and Nishimoto (1982). In the case of Tifgreen Bermuda grass, ratings were
based on a scale of 1-10 with ‘1’ being no phytotoxicity and ‘10’ being complete kill
of turf. For weeds, ‘1’ meant no control of weeds while a ‘10’ rating meant complete
weed control. Visual ratings were recorded by the same person throughout. Results
presented below are all average readings of three blocks/replications.
Results and Discussion
Phytotoxicity or weed control ratings in each trial were monitored for a period
of one month. Ratings of phytotoxicity for Bermuda grass were based on a scale
of 1-10. Ratings of ‘1’ and ‘2’ are to be considered as showing no phytotoxicity;
in some instances ‘imperfections’ in the turf were caused by slight scalping during
mowing etc and a rating of ‘2’ instead of ‘1’ was given. Ratings of ‘3’ and ‘4’ in-
dicate mild phytotoxicity and ‘5’ shows 50% turf burn. Ratings of ‘6’ and ‘7’ in-
dicate increasing levels of unacceptable burn while plots given ratings of ‘8’ and above
were very badly burnt and unsightly.
Phytotoxicity symptoms in Bermuda grass, when they occurred, were in most cases
very quickly apparent i.e. within two days from herbicide application (Fig. 1). It was
also found that in most cases, turf that were burnt to ratings of ‘7’ and above were
able to recover to ratings of ‘4-6’ within a month from herbicide application. It must
be noted though that the seemingly rapid recovery from burn was probably due to
Fig. 1.
AVERAGE PHYTOTOXICITY RATING
AVERAGE PHYTOTOXICITY RATING
10 20 30
NUMBER OF DAYS AFTER HERBICIDE APPLICATION
Effects of herbicides on Tifgreen Bermuda grass.
Each herbicide was tested at four levels of concentration with 0 ml or 0 g/l as controls against
which the effects of other treatments were compared. Herbicides were applied once at the start
of the trial after which turf quality was monitored. This is expressed as phytotoxicity ratings on
a scale of 1-10 where ‘1’ means no phytotoxicity symptoms and ‘10’ means complete killing of
turf. Results presented are averages of three blocks/replicates. Herbicide concentrations are
represented by the following symbols:-
(x —————x ), ( « --------- Fey Loe SCG x) — metribuzin (Sencor WP 70) at 0.2, 0.6
and 1.2 g/l respectively; (oo. ),, ( 0 --------- aye csi kt ©) — dicamba (Fez
PE 400) at 1.0, 2.0 and 3.0 ml/l; (e——————e), ( @ --------- ye. en ee e)—
bentazon (Basagran 39.6%) at 2.0, 6.0 and 10.0 ml/1; (a—————a), (4 -------- A),
RE 4) — 2,4-D (2,4-D amine 720) at 2.0, 4.0 and 8.0 ml/l; (a—————a),
( a --------- 8 5 eae 4) — MSMA (MSMA 500) at 2.0, 6.0 and 10.0 ml/I.
105
10 re Se ee ee a
AVERAGE WEED CONTROL RATING
AVERAGE WEED CONTROL RATING
10 20 30
NUMBER OF DAYS AFTER APPLICATION
Fig. 2. Effects of herbicides on Cyperus kyllingia.
106
The procedure for this trial and symbols for herbicides and concentrations are as found in
Fig. 1. Weed control was expressed as weed control ratings of 1-10, with ‘1’ meaning no control
and ‘10’ meaning total killing of weeds. Results presented are averages of three blocks/replicates.
Ss
MSMA For Controlling Weeds in Tifgreen Bermuda Grass Turf 107
the fact that even when most of the turf within a treatment plot was killed, turf sur-
rounding it continued to grow, sending new shoots into the damaged plot. Thus
recovery from herbicide burn of ratings ‘7’ and above, under trial conditions, should
not be taken to mean that the same will be expected to occur if large areas of turf
were blanket sprayed with overdoses of herbicides. Spot spraying appears to be the
safer method of application and if blanket spraying is to be carried out then lower
dosages of herbicide should be used, repeating the application when necessary.
Results shown in Fig. 1 are summarised in Table 1 which shows herbicide concen-
trations that produced phytotoxicity ratings of ‘4’ and below in Bermuda grass. The
rating of ‘4’ was chosen as the upper limit for acceptable turf because the visual
appearance of grass with this rating and below was reasonably good and recovery
from the effects of herbicide was rapid. With this as the criterion it is seen that
metribuzin WP70 at 0.2 g/l, bentazon 39.6% at 2.0, 6.0 and 10.0 ml/l, and MSMA
500 at 2.0 and 6.0 ml/1 could be ‘safely’ sprayed on Tifgreen Bermuda grass.
Table 1
Herbicide concentrations that could be ‘safely’ used on Tifgreen Bermuda grass.
Metribuzin (Sencor WP 70)
g/l
Dicamba (Fez PE 400)
ml/]
Bentazon (Basagran 39.6%)
ml/]
2,4-D (2,4-D amine 720)
ml/]
MSMA (MSMA 500)
ml/1
Concentrations of herbicides that caused burn up to a upper limit of phytotoxicity rating ‘4’ are marked
(X); these are considered ‘safe’ for use on the turf under the experimental conditions employed. Results
shown here are extracted from data in Fig. 1. Plots of turf showing phytotoxicity ratings of ‘4’ and below
were not excessively unsightly and recovered from burn rapidly.
Figures 2-4 show the effects of herbicides on Cyperus kyllingia, Euphorbia
thymifolia and Axonopus compressus respectively. Weeds that were controlled at
ratings of ‘1-5’ were found to recover quickly from the effects of the herbicides.
Increasingly effective control was noted with ratings of ‘6-8’ while in cases where
ratings of ‘9-10’ were achieved, excessive damage to Bermuda grass also occurred.
Herbicide concentrations that resulted in control ratings of ‘7’ were therefore con-
sidered potentially useful for selective weed control.
r&
« °4 2. °%2 = = 2 * ee eee
. (eo) = re A st i Se al
4
h -_ >>
i os
AVERAGE WEED CONTROL RATING
AVERAGE WEED CONTROL RATING
10 20 30
NUMBER OF DAYS AFTER APPLICATION
Fig. 3. Effects of herbicides on Euphorbia thymifolia.
The explanation for this figure follows that in Fig. 2.
108
on ili —
AVERAGE WEED CONTROL RATING
AVERAGE WEED CONTROL RATING
NUMBER OF DAYS AFTER APPLICATION
Fig. 4. Effects of herbicides on Axonopus compressus.
The explanation for this figure follows that in Fig. 2.
109
110 Gard. Bull. Sing. 40(2) (1987)
Table 2 summarises herbicide concentrations that affect weeds by at least a
minimum control rating of ‘7’.
For the sake of convenience, it would be useful to have a herbicide which will
control all three weeds (A. compressus, E. thymifolia, C. kyllingia) at the same time.
Using the rating of ‘7’ as the criterion, it is seen that herbicides like dicamba and
2,4-D could only control E. thymifolia but not the other two weeds. Similarly, ben-
tazon at its highest concentration tested controlled C. kyllingia and E. thymifolia but
not A. compressus. Dicamba, 2,4-D and bentazon were each not versatile enough
for the purpose of providing control of all three weeds. Only applications of metribuzin
at 0.6 or 1.2 g/l, or MSMA at 6.0 or 10.0 ml/I could be expected to control all three
weeds at the same time. Results in Fig. 1 however show that metribuzin at 0.6 and
1.2 g/l, and MSMA at 10 ml/I caused excessive burn in Tifgreen Bermuda grass.
This left only MSMA at 6 ml/I as the potential for use in universal control of all
three weeds.
MSMaA (6 ml/Il) was sprayed on 3 m X 1 m plots at the rate of 55 ml spray solu-
tion per square metre using CP15 Knapsack sprayers. Plots were either sprayed only
once at the start of the experiment or once a week for 3 weeks in succession. Since
metribuzin at 0.6 g/l could control all three weeds but was too phytotoxic for Tifgreen
Bermuda grass, it was decided to test metribuzin at 0.2 g/l to determine if repeated
spraying could bring about the same effects as found for the higher concentration.
Results show that repeated weekly spraying of MSMA for 3 weeks resulted in selec-
tive control of A. compressus, C. kyllingia and another weed Brachiara distachya;
Table 2
Herbicide concentrations that effectively controlled C. kyllingia,
E. thymifolia and A. compressus.
Metribuzin
(Sencor WP 70)
g/l
Dicamba
(Fez PE 400)
ml/1
= = =
r) SS) 'S)
Ww
cc)
Bentazon
(Basagran 39.6%)
ml/]
2,4-D
(2,4-D amine 720)
ml/1
MSMA
(MSMA 500)
ml/1
i)
©
~ “wal ee: ;
© ole
Concentrations of herbicides that controlled the 3 weeds by at least a control rating of ‘7’ are marked
(X); these are considered effective for control of the respective weeds. Results shown here were extracted
from Figs. 2-4.
fe lie =
MSMA For Controlling Weeds in Tifgreen Bermuda Grass Turf 111
E. thymifolia were weak in growth, ceased spreading but were not totally killed.
Cyperus radians, a coarse, tufted sedge with wiry leaves and inflorescence stalks (Wong
1986) was affected by MSMA to the extent that tips of leaves and inflorescences were
burnt but the tuft remained alive; growth was impeded although total killing of the
weed did not occur. Metribuzin when sprayed at 0.2 g/l weekly for 2 weeks in suc-
cession resulted in total control of E. thymifolia but had no effect on A. compressus,
B. distachya or C. kyllingia even after 3 applications (weekly) in succession. To con-
firm the selectiveness of MSMA at 6 ml/I, it was sprayed repeatedly at weekly inter-
vals for 5 weeks on a9 m X 2m plot (55 ml spray solution per square metre).
A. compressus, C. kyllingia, B. distachya were all controlled while the Tifgreen Ber-
muda grass began to grow, spreading itself to cover areas formerly covered with weeds.
The entire plot looked darker green than the surrounding areas (Plate 1) by virtue
of the cover eventually provided by Tifgreen Bermuda grass within the plot; surroun-
ding areas were lighter green because they were still infested with weeds that gave
the lawn a lighter green color.
Plate 1. Weed control and the accompanying growth of Tifgreen Bermuda grass in MSMA-treated plots.
The darker green plot of turf in the figure resulted from the eradication of weeds like C. kyil-
ingia, A. compressus and B. distachya accompanied by active growth of Bermuda grass which
was not affected by the herbicide and was also darker green in colour than the weeds. MSMA
500 (6 ml/l) was sprayed at weekly intervals for 5 weeks in succession.
AZ Gard. Bull. Sing. 40(2) (1987)
Conclusion
Chemical control of weeds by selective herbicides is possible. Care must however
be taken to ensure the application of correct dosages. MSMA, a pentavalent arsenical
herbicide considered to be of a low order of toxicity to man and animals, and which
is inactivated upon contact with soil (Anderson 1977), was found to be highly selec-
tive in this trial. It controlled A. compressus, B..distachya, C. kyllingia and to a lesser
extent E. thymifolia without damaging Tifgreen Bermuda grass. Metribuzin, an
asymmetrical-triazine herbicide which is detoxified by microbial action in the soil
(Anderson 1977), was more effective on EF. thymifolia.
Acknowledgements
I wish to thank Mr Rehan bin Yusoff and Mr Ang Peter for their technical
assistance.
References
Anderson, W.P. (1977). Weed Science: Principles. West Publishing Co., New York:
277-280.
Batten, S.M. (1984). Those irrepressible, incredible, impossible grassy weeds! USGA
Green Section Rec 22(5): 1-4.
Ellis, G. (1986). War on weeds! Horticulture Week 199(2): 38.
Hanna, W.W. (1986). A Bermuda grass primer and the Tifton Bermuda grass. USGA
Green Section Rec 24(1): 11-13.
Johnson, B.J. (1983). Tolerance of Bermuda grass (Cynodon dactylon) putting greens
to herbicide treatments. Weed Sci. 31: 415-418.
Murdoch, C.L. and R.K. Nishimoto (1982). Diclofop for goosegrass control in Ber-
muda grass putting greens. HortScience 17(6): 914-915.
Wong, Y.K. (1986). The use of Tifgreen and Tifdwarf Bermuda grasses in two
Singapore golf courses. Gard. Bull. Sing. 39(2): 203-214.
es
Annotated List of Seed Plants in Singapore (XI)*
HSUAN KENG
c/o Department of Botany, National University of Singapore
Index to Families
Page Page
Agavaceae 6 Tridaceae 123
Alismataceae 113 Liliaceae 115
Amaryllidaceae 119 Najadaceae 115
Bromeliaceae 126 Palmae 126
Burmanniaceae 123 Philydraceae 123
Butomaceae 113 Pontederiaceae 122
Commelinaceae 124 Smilacaceae 118
Dioscoreaceae 12] Taccaceae 12]
Eriocaulaceae 126 Triuridaceae 115
Flagellariaceae | W25) Xyridaceae 126
Hydrocharitaceae 114 Zannichelliaceae 115
Hypoxidaceae 120
II. Angiospermae-Monocotyledons
143. ALISMATACEAE
Sagittaria sagittifolia L.
Aquatic herb; to 50 cm tall; leaves basal, with petiolate sheath and arrow-shaped
blade (15-30 cm long); flowers white, in panicles. Underground tubers globose,
4-5.5 cm across, edible; mostly imported from S. China especially during the
Chinese New Year Season, sometimes cultivated as an ornamental. Vern. Ubi
Keladi, Chinese arrowhead, hh .
144. BUTOMACEAE
Limnocharis flava (L.) Buch.
Aquatic herb, 50-70 cm tall; leaves petiolate, the blade ovate or suborbiculate,
6-28 cm long; flowers yellow, 2-8 or more in a stalked umbel. Native to S. America,
sometimes spreading along ditches or in pools.
*Continued from Gard. Bull. Sing. 39 (1): 67-95. 1986.
The author is indebted to Dr. Richard T. Corlett for going through the manuscript and for his suggestions.
113
114 Gard. Bull. Sing. 40(2) (1987)
145. HYDROCHARITACEAE
Key to the genera
A. Fresh water plants
B. Leaves spirally arranged or radical. .is.5...::4252. «sacnsjsbele 9h ou vitae. Gens eet ee Blyxa
B. Leaves in whorls of 358 . .. «6666 ¢ esses ayn uve « hieis Beceem Sedans ape = nee ee Hydrilla
A. Marine plants
C. Plants very delicate; leaves opposite, in spaced pairs on slender stems, ovate to lanceolate
cid gain ecm wb dhe v aceg yo. 5, buna a) 4 ie apm Oye eee egeae eee cate nel ae nee Halophila
C. Plants coarse; leaves arranged in 2 rows, ribbon-like
D. Rhizome covered with soft black fibres; leaves 30-150 cm long............... Enhalus
D. Rhizome nearly naked; leaves. 10-30'cm long... ...; 2: 2:2 05. . 22 ee Thalassia
Blyxa auberti Rich.
Formerly called Blyxa malayana Ridl. Freshwater plant, leaves radical, narrow
(less than 1 cm broad), 10-50 cm long; flowers bisexual, small, on a long stalk
_ surrounded by a tubular spathe. In ponds and ditches, Nee Soon, Serangoon Road
(Ridley 8413).
Blyxa alternifolia (Mig.) Hartog
Formerly called Enhydrias angustipetala Ridl. Leaves linear, 2-5 cm long, arranged
along the stems. In muddy ponds and ditches, Chua Chu Kang, (Ridley s. n. in
1905).
Enhalus acrocoides Royle
Formerly called Enhalus koenigii Rich. Marine plant; rhizome thick, covered with
stiff black fibres; leaves 2-3, strap-shaped, 0.5-1 m long; flowers small, unisexual,
dioecious, the female in coiled, long-stalked inflorescence. In shallow sea water,
Bajau (Goodenough 3952), Teluk Paku, Pulau Senang, Tanah Merah Besar,
Changi.
Halophila minor (Zoll.) Hartog
Small marine plant; stem slender, lateral shoots short, usually with one pair of
petiolate leaves; leaf-blades oblong-ovate, 0.7-1.4 cm long, with 3-8 pairs of side
veins; flowers small, dioecious, solitary, axillary. Below tidal level on muddy
bottom, Ponggol, (Holttum s.n. in 1924), Pasir Laba, Sentosa, Pulau Pawai,
Labrador.
Hal. ovalis (R. Br.) Hook. f.
Like the above, but the leaf-blades ovate, larger (1-4 cm long), with more pairs
(12-25 pairs) of side veins. Changi (Ridley s.n. in 1890), Woodlands, Pulau Senang.
Hal. spinulosa (R. Br.) Aschers.
Lateral shoots with 10-20 pairs of sessile distichous leaves; leaf-blades spathulate-
oblong, serrulate, 1-2.3 cm long. Pulau Tekong (Henderson & Corner, s.n. June
16, 1929), Beting Kusa, Tanah Merah Besar.
Hydrilla verticillata (Roxb.) Royle
Fresh water plant; stems slender, much branched; internodes 0.3-5 cm long; leaves
sessile, in whorls of 3-8, linear, 0.8-4 cm long, green with brown spots; flowers
very small, unisexual. In ditches and pools, Tanjong Pasir Laba Road (Sinclair
10836).
Thalassia hemprichii (Ehrenb.) Aschers.
Marine plant; rhizome creeping, conspicuously noded; leaves 2-6, in 2 ranks;
blades strap-shaped, 10-40 cm long; flowers dioecious, in few-flowered terminal
inflorescence. In mud at low tide, Pulau Samulun (Sinclair SFN 38581).
Annotated List of Seed Plants in Singapore (XI) 115
>
146. NAJADACEAE
Najas graminea Delile
Fresh water submerged herb; stems slender, branched; leaves opposite or nearly
so, linear lanceolate, 1.5-2.5 cm long, with 3 or more minute teeth on each side;
flowers very small, green, unisexual. In ditches and ponds, Gardens’ Lake (Ridley
8946, Type of N. graminea var. angustifolia Rendle).
Najas Kingii Rendle
Like the above, but leaves larger (2-4 cm long) and with more (6-19) teeth on
each side. In streams, Tanglin (Ridley s.n. in 1889).
147. ZANNICHELLIACEAE
Key to the genera
A. Leaves 7-13 nerved; style 2-fid; anthers inserted at the same level .................. Cymodocea
A. Leaves with a main vein and 2 thin marginal veins; style entire; anthers inserted at different levels
ree ge cas il AW alg E stn o Fa Oy Sorin Rak Evade S op we is 3 Halodule
Cymodocea rotundata Aschers. & Schweinf.
Marine herb, submerged; stems distant, erect, with few leaves at the top, in 2 series;
leaves linear or filiform, with a rounded top, 7-15 cm long and 7-13 nerved. Telok
Paku, (Sinclair SFN 40527).
Cym. isoetifolia Aschers.
Tanah Merah Besar (Sinclair SFN 39509). In sandy, muddy, substratum near low
water.
Cym. serrulata Aschers. & Magnus
Tanah Merah Besar (Sinclair SFN 39508).
Halodule tridentata (Steinh.) F.v.M.
Also called Diplanthera uninervis Aschers. Submerged marine herb; stems distant,
erect, with few leaves at the top, in 2 series; leaves linear, 7-10 cm long, 3-toothed,
the median tooth as long as or slightly longer than the lateral teeth; flowers solitary,
unisexual, naked. In sandy coast, Pulau Senang (Burkill 547), Pulau Pawai.
148. TRIURIDACEAE
Sciaphila maculata Miers.
Slender saprophytic herb, 10-15 cm tall, purplish grey, usually not branched; leaves
minute; flowers unisexual; perianth 6-lobed; stamens 6 in a staminate flower, and
carpels numerous in a carpellate flower. In dense forests, living in decaying
vegetable matter, very rare, Selitar (Ridley s.n. in 1880s). Formerly called S.
affinis Becc.
149. LILIACEAE
Key to the genera
caves mck and succnicnt often spiny-edged ¢.'. 2... 62 ee ae ecw ele re cele a wees Aloe
A. Leaves thin and flexible
B. Leaves ovate or ovate-lanceolate
C. Leaves, especially the upper ones, always with a long tendril-bearing tip; slender stemmed climber
ee lee er an bhp han o Me pes a pem a eae Gloriosa
C. Leaves long-petiolated, erect; underground stem very short.................- Peliosanthes
116 Gard. Bull. Sing. 40(2) (1987)
B. Leaves long and narrow or sometimes scale-like
D. Leaves scale-like, the function replaced by needle-like cladophylls; drooping or climbing
EL eTeT Teer er eee rrerrry Ce errr ier Asparagus
D. Leaves long and narrow, grass-like; erect
E. ‘Stems leafy,' in two rows |. 0.520 62500). 200 sae eee ee Dianella
E. Leaves tufted, foliage plants
F. Plants with prominent runners... 2:;..s4 4. i ee. oe Chlorophytum
F. Plants crowded, with short and thick rhizomes................,-.+ss56 Liriope
Aloe barbadensis Mill.
Also called A. vera (L.) Webb. A short-stemmed, succulent plant; leaves pale green,
sword-shaped, 30-60 cm long, crowded along the top of the stem, with marginal,
dark-coloured spines; flowers yellow, orange or red, in lax panicles. Native of Cape
Verde and Canary Island. This is the true aloe, a well-known medicinal plant;
several other species and cultivated varieties are also introduced. M#
Asparagus plumosus Baker
Slender herbaceous climber, up to 1-2 m long; true leaves minute, dry, scale-like,
replaced by fine, green, needle-like flattened branches (or cladophylls), 0.3-1 cm
long; flowers small, axillary, solitary or in pairs, greenish or white. Native to
S. Africa, often cultivated as an ornamental.
Asp. sprengeri Regel
Roots tuberous; stems climbing or drooping; flattened branches (or cladophylls)
1-2.5 cm long; flowers white, in short racemes. Native to S. Africa, sometimes
planted. The tender shoots of another species, the garden asparagus (Asp. offici-
nalis L.), a native of the Mediterranean, are sold in markets as vegetables.
Chlorophytum comosum (Thunb.) Baker
Herb; leaves radical, sessile, linear, 15-35 cm long, sometimes with white or pale
yellow margins or median band; flowers white, in branched racemes. Native to
S. Africa, often cultivated for its foliage, sometimes hanging from pendulous
runners. mia
Dianella ensifolia (L.) DC.
Rhizomatous tufted herb, 60 cm to 1 m tall; leaves grass-like, in 2 rows, 20-60
cm long, sheathed below; flowers bluish or white, in a términal panicle; filaments
thickened; berry globose, deep blue, 0.7-1 cm across. In open or slightly shady
places, often near the sea, Sungei Buloh (Ridley s.n. in 1891).
Gloriosa superba L.
Climbing plant with underground tubers; stems slender; leaves spirally arranged,
narrowly lanceolate, sessile, 8-25 cm long, with a long-tapering tip, the tip of higher
leaves forming a short, coiled tendril; flowers solitary, long-stalked, placed beside
the leaves; perianth lobes at first pale yellowish green, than partly red with yellow
margins, finally dark red, strongly bent backwards. Native to tropical Africa and
Asia (but not wild in Malaya and Singapore), cultivated as an ornamental. All
parts, especially the tubers, are poisonous to eat.
Liriope spicata Lour.
Herb, with semi-transparent rhizomes; leaves grass-like, linear-lanceolate, tufted,
25-50 cm long, 1-1.8 cm wide, often with white margins; flowers small, pale lilac
or white, in short spikes 10-15 cm long (pot-plants usually producing shorter leaves
and inflorescences). Native to S. China and Japan, commonly cultivated as a pot
plant; the rhizomes are of medicinal value, #f1*% . Another broader-leaved (1-2.5
cm wide) species, L. muscari Bailey, with violet flowers, is less commonly cultivated.
Annotated List of Seed Plants in Singapore (XI) 117
Peliosanthes teta Andr. ssp. humilis (Andr.) Jessp.
Herb; leaves crowded on a horizontal rhizome, erect, ovate-lanceloate, 15-20 cm
long, 5-6 cm wide; with strong longitudinal veins; petioles 12-15 cm long; flowers
globose, violet or purple, in racemes about 10 cm long. Bukit Timah, Reservoir
woods. (Ridley s.n. 1908), Selitar. Formerly called P violacea Wall, and P viridis
Ridl.
150. AGAVACEAE
Key to the genera
A. Ovary superior
TR EIN NNEC ere ore ws Pe Ariat ick > sk ewe wo clu <i os sia enle 6 0a DEBE PRs ee 8% Yucca
B. Periath-segments united at the base
Go@vules 4-iany per OvalyOGihe nus). OUT. ok APPL Uk ee ee Cordyline
C. Ovule solitary
>. Shrubs or trees; stems present, woody... 20.0... 5.800 08. Dracaena (incl. Pleomele)
eaters. Stein Very SiOrt, TOrOUS .!0 20 ess Ferd OA CI ol. Sansevieria
A. Ovary inferior
Po Piowersuceular. milorescegce pamiculate . 2... ssc cons lds Sad a enc wel ny ee Agave
E~ Fiowers zygomorcphic:: inflorescence TACEMOSe = 5%... lac see sees ee Polianthes
Agave americana L.
A large, stemless plant with a rosette of 50-60 thick, sword-shaped leaves; the
blades 0.8-1.5 m long, the marginal spines often recurved; flowers pale yellow,
in large panicles on a culm of 4-8 m tall. Vern. Century plant #45 . A native
of Mexico, sometimes planted. Several other species (e.g. A. sisalana Perr., A.
germinoflora Ker-Gawl.) are also occasionally planted.
Cordyline terminalis (L.) Kunth
Shrubby, 3-4 m tall; stems slender; leaves narrowly oblong, 30-50 cm long,
arranged in close spirals at the tip of branches; petioles 5-15 cm long; flowers lilac,
in terminal panicles. Native of temperate Asia and Australia and the Pacific Islands,
planted in gardens. The leaves are in a wide range of colours. *#. Several other
Australian species, e.g. C. stricta Endl., C. fruticosa A. Chev., are also occasionally
cultivated.
Dracaena aurantiaca Wall.
Shrub, 2-3 m tall; little or not branched; leaves varying from lanceolate to ovate,
0.3-1 m long, green or purple with circular lighter patches, crowded on the ends
of branches; petioles winged; flowers greenish white or purple, in terminal panicles,
0.5 m long. In forests, Bukit Timah, Kranji, Changi, Selitar (Ridley 1650).
Drac. brachystachys Hook. f.
Little branched shrub, 5-6 m tall; leaves crowded at the ends of branches; leaves
sword-shaped, acuminate, to 1 m long. Pulau Serapu (Ridley 10/28).
Drac. elliptica Thunb.
Slender shrub, 1-2 m tall; branches drooping; leaves lanceolate or broadly oblong,
12-15 cm long. Gardens’ Jungle (Ridley 4412), Chua Chu Kang.
Drac. fragrans Ker. (= Pleomele fragrans N. E. Br.).
Tall shrub, to 6-8 m high; leaves oblong-lanceolate, 0.5-1 m long, 5-10 cm wide,
green or with light longitudinal bands, crowded on the ends of branches, without
a petiole; flowers yellowish, clustered on branches of a panicle, fragrant especially
at night. Native of tropical Africa, sometimes planted. Several other African species
118 Gard. Bull. Sing. 40(2) (1987)
are also commonly planted, these include: D. godseffiana Sand. (leaves oblong-
ovate, green marked with light dots) D. sanderiana Sand, ex Masters (leaves long-
lanceolate, white-margined), etc.
Drac. granulata Hook. f.
Tall tree, 15-20 m high, branched above; branches densely leafy; leaves linear,
acuminate, 15-25 cm long. In forests, Bukit Timah, Bukit Mandai (Ridley 3800).
Drac. maingayi Hook. f.
Large tree, 15-20 m tall, with many branches forming a dense crown; leaves oblong-
lanceolate, acuminate, 25-30 cm long. Common in sandy and rocky woods near
the sea, Changi (Ridley 44/3), Bukit Timah, Labrador.
Drac. porteri Bak.
Low, little branched shrub, 1-1.5 m tall; leaves linear or linear lanceolate, 30-40
cm long. In lowland woods, Singapore (Wallich 5148B), Gardens’ Jungle.
Drac. singapurensis Rid.
- Shrub with slender stems, less than 40 cm tall; leaves lanceolate or elliptic, acute,
15-18 cm long. Bukit Timah, Chua Chu Kang (Ridley 6235).
Drac. umbratica Ridl.
Erroneously called D. terniflora Roxb. in earlier literature. Low shrub, little
branched, to | m high; leaves lanceolate to elliptic, 3-5 cm long. Gardens’ Jungle,
Changi (Ridley 4758).
Polianthes tuberosa L.
Herb, 1 m tall, tuberous beneath; basal leaves narrow, red-spotted on the under
surface, 15-50 cm long; upper leaves gradually shorter and then passing into bracts;
flowers white, fragrant, in pairs on terminal spikes. Native to Mexico, sometimes
cultivated. #&£ .
Sansevieria trifasciata Prian
Herb, leaves 2-6 arising from the underground rhizome, the blades erect, sword-
shaped, leathery, cross-banded dark and light green, 0.3-1 m long; flowers greenish
white or greyish, in fascicles on a raceme 40-75 cm long. Native to tropical Africa,
sometimes cultivated. Vern. Bowstring hemp, /é. Another introduced species
is S. cylindrica Bojer which has erect cylindric leaves 0.5-1 m long. Their strong
fibres are made into mats, bowstrings, etc.
Yucca aloifolia L.
Stems simple or branched, 1-3 m tall; leaves sword-shaped, 20-50 cm long, hard
and spiny; flowers white, in panicles of 30-60 cm long. Native to the West Indies
and eastern N. America. A form with white-streaked leaves is cultivated. Another
species with smooth leaf-margins, Y. gloriosa L. is also introduced.
151. SMILACACEAE
Key to the genera
A. Perianth-segments free; stamens.6, free «... ... s+ > 5 +i jess) =e bape « age Smilax
A. Perianth-segments united; stamens 3, fused... 4. ..0iss+» «5 «aes e) mies bp ee Heterosmilax
Heterosmilax indica A. DC.
Climber, with stipular-tendrils; stems slender, unarmed; leaves ovate-lanceolate,
with a round base, 10-20 cm long, 3-8 cm wide; flowers in umbels. Once collected
from Tanjong Katong (Hullett s.n.), now extinct.
Annotated List of Seed Plants in Singapore (XI) 119
Smilax bracteata Pres| var. barbata (Wall. ex DC) Koyama
Woody climber, climbing by stipular-tendrils; stems stout, covered with red and
whitish bristles and conic prickles; leaves thick leathery, ovate, with a round base,
15-20 cm long, 7-15 cm wide, glaucous beneath; petioles thick, 3-4 cm long; flowers
unisexual, in compound umbels. In open country and forest edges, Tanglin, Bukit
Timah (Ridley 4811, 8043).
Smil. calophylla Wall. ex DC
Wiry shrub, erect, unarmed; leaves lanceolate, acuminate, 3-nerved, glaucous
beneath, 10-15 cm long; flowers yellow, unisexual, in small, sessile umbels; fruit
globose, reddish brown. In forests, Bukit Timah, Jurong, Chua Chu Kang (Ridley
s.n. in 1892).
Smil. leucophylla Bl.
Woody climber, stout, sparsely prickly; leaves leathery, ovate-oblong, acuminate,
3-nerved, 15-25 cm long; petioles 1-1.5 cm long, with a large dilated sheath. In
forests, Bukit Timah, Tanglin, Bukit Mandai (Ridley 10403). Vern. Ubi danan.
Smil. megacarpa DC.
Stout woody climber, prickly, with strong stipular-tendrils; leaves leathery, dark
green, oblong to elliptic, 3-nerved, 8-20 cm long; petioles 1.5-4 cm long; thick,
twisted, sheathed below; fruit globose, 1.5 cm across, yellow or dark purple. In
woods, Tanglin (Ridley 4815, 5106), Bukit Mandai.
Smil. myosotiflora DC.
A slender climber; leaves herbaceous, lanceolate or ovate-oblong, caudate,
3-nerved, 10-15 cm long. In forests, Gardens’ Jungle (Ridley 6250). Sinclair
recorded another species, S. woodii Merr. without certainty as occuring in Singapore
based on Sinclair SFN 38570 collected from Jurong Road in 1949.
152. AMARYLLIDACEAE
Key to the genera
CRS Wie i ce ie caine, lassie RK ao ele aur 5 2 Oh ayes ns bok Allium
A. Ovary inferior
ER A IRE te dat oO > Me ect eis ele 2 oe ae ed kee ee eke ees Zephyranthes
B. Flowers in umbels or globose-heads, erect or pendulous
C. Flowers without a crown or cup
D. Flower-stem hollow (leaves long and narrow; flowers large, trumpet-shaped)..........
ee ah Se A Ae tah ale ae ie Sea's g's av Sup sles » Hippeastrum
D. Flower-stem solid
E. Flowers 50-100 in a globose head; leaves oblong, petiolate ........... Haemanthus
E. Flowers 6-12 in a congested umbel; leaves long and narrow, without a petiole .....
Crinum
Pita e ole Bow ole loa S | (el Seles o ©) 0 = jo dia eo @ eS © 6 ©\.e wisn. s) 0.6, 60ule 6 6 G's @ Tye. 6 ee Dee T Een ee te
C. Flowers with a crown or cup inside the perianth-tube, consisting of joined filaments of stamens
F. Leaves board, narrowed to a petiole
is) PaO Wer -SeRINCIMS SPLCdGIN®, ITASTANE .. 6c. os ew em ct ee cease eee Eucharis
Tae mower -scuments ascending, OGOUTIESS 6 ce ce eee ee ee Eurycles
Po Eeaves swore-snaned, without a petiole? .. 5s ee a Hymenocallis
Allium fistulosum L.
Bulbous herb; leaves radical, long-tubular, hollow, 30-50 cm long; flowers whitish,
small, in dense, terminal umbellate-heads; flowering stem 40-60 cm long. Native
of temperate Asia, cultivated for the edible leaves. Vern. Spring onion, Welsh
onion, X# . Several other species are used as vegetables or for cooking, these
120 Gard. Bull. Sing. 40(2) (1987)
include A. cepa L. (the onion, #% ), A. sativum L. (the garlic, # ), A. tuberosum
Rottl. ex Spreng. (or as A. ramosum L., the Chinese leek, 4£4 ), etc.
Crinum asiaticum L.
Bulbous herb; leaves narrowly lanceolate, acuminate, 0.5-1.5 m long; flowering
stem 1-1.2 m, with 6 or more flowers on a congested umbel; flowers white, 12-18
cm long, the segments narrow; filaments reddish; fruit irregularly globose, 4-5
cm across, with one or few large green seeds inside. On sandy shores, Changi,
Sungei Buluh, Sungei Tengah (Ridley s.n. in 1891), Kranji; sometimes also
cultivated. Vern. Bakung, <#& .
Crinum giganteum Andr.
Flowers larger (about 25 cm long) and with broader segments than the above
species. Introduced from Java, thus called ‘Java Lily’.
Eucharis grandiflora Planch. & Link
Bulbous herb, with 2-4 leaves; leaf-blades oblong or ovate, 20-25 cm long; petioles
_as long as the blades; flower stems 50-80 cm long, with 2-5 flowers at the tip;
flowers white, 6-8 cm across, widened part of the perianth-tube pale yellow in-
side. Native of Colombia (‘‘Amazon Lily’’), sometimes cultivated. S#/Eé.
Eurycles amboinensis Loud.
Bulbous herb; leaves broadly heart-shaped, 20-30 cm long; petioles stout, 15-40
cm long; flowering stem 10-50 cm long. Native of N. Malaya, the Philippines to
E. Australia, occasionally cultivated. Also called E. sylvestris Salisb.
Haemanthus multiflorus Martyn
Bulbous herb; leaves 4-7, oblong, with a stout midrib, 20-30 cm long; scape solid,
flattened; flowers bright or pale red, numerous in a globose head 12-16 cm across
(‘‘Blood Lily’’ or ‘‘Powder puff’’). Native to tropical Africa, sometimes cultivated.
PUHETE .
Hippeastrum puniceum Urb.
Bulbous herb; leaves 6-8, strap-shaped, 30-45 cm long, usually not fully grown
before flowering; flowering stem 30-50 cm long, with 2-4 or more flowers at the
top; flowers trumpet-shaped, 9-12 cm long, red, orange-red or in other colours.
Native to tropical America (‘‘Barbados Lily’’), sometimes cultivated; also called
H. equestre Herb. ‘kEK{E .
Hymenocallis littoralis Salisb.
Bulbous herb; leaves radical, strap-shaped, acute, with a narrow base, 40-70 cm
long; scape flattened, 2-edged, 30-70 cm long; staminal cup (or crown) 2.5-3 cm
long. Native to tropical America, sometimes cultivated (“‘Spider Lily’’). Also called
Pancratium littorale Jacq.
Zephyranthes candida Herb.
Bulbous herb; leaves linear, 12-40 cm long, very thick; scapes with only one flower
at the tip; flower white, sometimes rose-tinted outside, 4-5 cm long. Native to
S. America. #i#. Several other species are also occasionally cultivated, these in-
clude: the light-pink flowered Z. carinata Herb, the yellow-flowered Z. carinata
Herb, the yellow-flowered Z. flava Bak. and others.
153. HYPOXIDACEAE
Curculigo orchioides Gaertn.
Rhizomatous herb; leaves narrowly lanceolate, 1-1.2 m long including the petiole,
12-15 cm wide, glabrous; flowers bisexual, several to many in erect, head-like
Annotated List of Seed Plants in Singapore (XI) 121
inflorescence on a short scape (about 10 cm long); bracts lanceolate, 2-3.5 cm
long; perianth 1-1.2 cm across, pale outside and bright yellow inside. In sunny
or slightly shady places, formerly all over the island. Cluny Road (Ridley s.n. in
1895), Bukit Timah, Chua Chu Kang. Called C. Jatifolia Dryand ex W. T. Ait
in literature.
Curc. recurvata Dryand. ex W. T. Ait
Differs from the above species in the nodding head-like inflorescences. (also called
Molineria recurvata Herb. by some authors). Gardens (Ridley s.n. in 1896).
Curc. villosa Wall.
Like C. orchioides, but the undersurface of leaves and petioles are densely hairy,
and bracts are stiff, oblong, 4-5 cm long. Chua Chu Kang (Ridley s.n. in 1895),
Bukit Timah.
154. TACCACEAE
Tacca integrifolia Ker-Gawl.
Rhizomatous herb; leaves simple, entire, lanceolate, 0.8-1 m long including the
petiole; flowers in umbels; involucral bracts leafy, dark purple; scape 45-60 cm
long, stout; floral bracts thread-like, 12-15 cm long; perianth 1.5-2 cm across,
purple; segments in 2 series, the outer ones thinner, paler and smaller than the
inner ones. In forests, Bukit Timah. Chua Chu Kang, (Ridley s.n. in 1894). Formerly
called T. cristata Jack.
Tacca leontopetaloides (L.) O. K.
Leaves tripartite or pinnatifid, 0.5-1 m long; flowers green. On sandy shores, Pulau
Semakau, Sinclair SFN 38925; Pulau Senang.
155. DIOSCOREACEAE
Dioscorea alata L.
Stem slender, 4-angled or -winged, twining in anti-clockwise direction; leaves
opposite, orbiculate-cordate, 5-7-nerved. Native of E. Asia, cultivated for its
underground tubers (‘‘the greater Yam’’). X® .
Dios. bulbifera L.
Stem twining clockwise, glabrous, often with axillary bulbils; leaves cordate,
broadly ovate, 8-40 cm long; 6-30 cm wide, conspicuously cross-veined; petiole
2-13 cm long; flowers seesile or nearly so, white to dark violet; male spikes 2-3.5
cm long, in panicles; female spikes in axillary fascicles, 10-20 cm long. In waste
ground and near villages, Tanglin, Changi, Pulau Merambong (Corner 29960).
Dios. esculenta (Lour.) Burk.
Stem twining clockwise; leaves alternate; tubers 4 to many, close to the soil
surface, anchored by strong thorny roots. Native to Indo-China, cultivated for its
edible tubers (‘‘the lesser yam’’). #/% .
Dios. glabra Roxb.
Stem rather stout, thorny at base; leaves lanceolate to elliptic-ovate, 4-10 cm long,
glabrous; petiole 2-2.5 cm long; male and female spikes 10-30 cm long, in panicles.
Common in forest, Gardens’ Jungle, Tanglin, Bukit Mandai.
Dios, hispida Denns.
Stem spiny, twining closewise; leaves 15-30 cm long, palmately 3-foliolate; Male
panicles 30-50 cm long. Several cultivated varieties; tubers mostly globose, usually
not lobed. Bukit Timah (Goodenough s.n. in 1914).
122 Gard. Bull. Sing. 40(2) (1987)
Dios. laurifolia Wall.
Stem slender, smooth or prickly; leaves lanceolate to oblong, 3-nerved, 5-8 cm
long; male and female spikes in panicles. In edges of forests, Tanglin, Gardens’
Jungle (Ridley s.n. in 1893), Bukit Timah, Changi.
Dios. polyclades Hook. f.
Stem stout, pubescent, thorny near the base; leaves ovate, hairy, 5-7 cm long;
petiole as long as the blade, hairy; male and female spikes in panicles. In forest
edges, Chua Chu Kang, Sungei Pandan, Bukit Timah.
Dios. prainiana R. Kunth
Stem slender, glabrous, unarmed; leaves alternate or opposite, membranous,
elliptic-oblong, 5-7-nerved, 10-15 cm long; petiole 4-6 cm long; male spikes 30-60
cm long. In forests, Bukit Timah, Gardens’ Jungle.
Dios. pyrifolia Kunth
Stem pubescent, the base thorny; leaves opposite, leathery, ovate, cordate,
_ 5-nerved, 6-10 cm long, pubescent below; petiole 2-5 cm long; male spikes 30 cm
or more long; female spikes 8-10 cm long. Common in thickets, Chua Chu Kang,
Changi.
Dios. stenomeriflora Prain & Burk.
Large climber, to 25 m tall; stem purple; leaves leathery, glabrous, elliptic, cor-
date, 3-nerved, 10-25 cm long; male spikes solitary, axillary; flowers long tubular,
about 1 cm long. In forest edges, Changi. Another large climber with large cordate-
hastate leaves, D. sansibarensis Pax, a native of tropical Africa, was once planted
in the Botanic Gardens; now escaped and more or less naturalized in the surroun-
ding areas of the Gardens; propagated by bulbils.
156. PONTEDERIACEAE
Key to the genera
A. Flowers zygomorphic; perianth forming a tube at the base; leaves with a short swollen petiole ...
rear es eee ee Ne eee ee NE ee Eichhornia
A. Flowers regular; perianth segments free; leaves with a long slender petiole .......... Monochoria
Eichhornia crassipes (Mart.) Solms.
Stoloniferous floating herb; roots dark brown, much branched; leaves nearly round-
ed, 5-20 cm long and broad; petiole spongy, 10-30 cm long, much swollen near
the centre; flowers zygomorphic, lilac blue; segments 6, the median one larger,
3-3.5 cm long, with a yellow blotch near the centre. Native of Brazil, in aquaria
and ponds; a serious pest in water catchment areas. Vern. Water hyacinth, JAiki¢ .
Monochoria hastata (L.) Solms.
Aquatic herb; leaves spear-shaped, 6-10 cm long and wide, with 2 pointed lobes
at the base; petiole 20-50 cm long, more or less erect; flowers light blue-violet,
2-2.5 cm across, in head-like inflorescence arising from the axil of a sheath in
leaf-stalk. In ditches and wet places, Changi (Goodenough s.n. in 1890), Gelang.
Monoch. vaginalis (Burm. f.) Presl.
Leaves narrower than the above species, the base rounded or heart-shaped, not
pointed. In ditches and ponds, Ang Mo Kio (Ridley 4598), Chua Chu Kang, Gelam.
en
és renee
Annotated List of Seed Plants in Singapore (XI) 123
157. IRIDACEAE
Trimezia martinicensis Herb.
Bulbous herb; leaves grass-like, 50-60 cm long, in 2 rows; flowering scape 60-70
cm long with 6 or more flowers in succession; flowers bright yellow, the inner
segments oblong, 2-2.5 cm long, with a dark brown blotch at the base inside. Native
to Mexico, cultivated and more or less naturalized. (Several species of Belamcanda,
Moraea etc. are occasionally planted in gardens.)
158. BURMANNIACEAE
Key to the genera
A. Stamens 3
Be Ovary 3-loculate; autotrophic or saprophytic *. 20). eee... Burmannia
eee a ect me sarmopnytiG ) 2 red ee EAS SPR... Gymnosiphon
RR ee tne eee ens oe cISN Swe Tee EI ee ewe wc bees Thismia
Burmannia championii Thw.
Saprophytic herb; stem swollen below and buried in decaying debris, slender above
ground, whitish, 8-20 cm long; leaves scale-like; flowers 3 or more in small clusters,
yellowish white; perianth tube 3-angled, not winged. In forests, Bukit Timah (Ridley
s.n. in 1890), Changi. Formerly called B. tuberosa Becc.
Burm. coelestis D. Don.
Autotrophic herb; stem slender, 10-25 cm tall; leaves small, narrow, to 1.2 cm
long, crowded at the base of stem; flowers 1-4 at the tip of stem, the perianth
tube about 1 cm long, 3-winged, light blue, with 6 small yellow or white perianth
segments. In open grassland or sandy places, locally common. Botanic Gardens,
Changi, Kranji, Bedok (Ridley s.n. in 1903).
Gymnosiphon aphyllus Bl.
Saprophytic herb, 10-17 mm long; leaves acute, 1-2 mm long; flowers white or
lilac, 5-6 mm long, in loose inflorescence. Formerly collected from Bukit Timah
(Ridley s.n. in 1893), now probably extinct.
Thismia aseroe Becc.
Slender, saprophytic herb; stem erect from creeping rhizome; flowers solitary, on
top of the erect branch; perianth-tube obconical, 1-1.2 cm long, the segments
triangular, with bright orange, long tails. In dense forests, formerly collected at
Woodlands, Bukit Timah (Ahmad s.n. in 1932), Kranji, now probably extinct.
Thism. fumida Ridl.
Like the above, but with narrower perianth-segments. Formerly collected from
forests at Chua Chu Kang, now extinct.
159. PHILYDRACEAE
Philydrum lanuginosum Banks ex Gaertn.
Grass-like herb; leaves narrow linear, 30-40 cm long, 1-1.2 cm wide; flowering
stem to 1 m tall, often branched at the top; flowers yellow, in compact heads,
subtended by stiff brown bracts; perianth segments 6, the outer 3 very unequal,
the inner three equal. In wet open places, formerly collected from Bedok (Ridley
5907), now extinct. H# .
124 Gard. Bull. Sing. 40(2) (1987)
160. COMMELINACEAE
Key to the genera
A. Cultivated plants
B. Short-stemmed; leaves sword-shaped, dark green above, crowded in a rosette.......... Rhoea
B. Stem slender, trailing; leaves oblong, alternate
C. Outer and inner perianth-lobes’ frée °°... ON Pa ase ee eee Tradescantia
C. Outer and inner perianth-lobes united below i700 .0..5 0 2.0 ee. foes ee Zebrina
A. Native plants (and weeds)
D. Inflorescence a compact head, breaking out through the base of the leaf-sheath .......
ie es ee pains aa a 08 ileal poate ba cereal Syed a" ita eich ela ce oe ec ne Forrestia
D. Inflorescence terminal or axillary, not breaking out through the base of leaf-sheath
E. Inflorescences congested, enveloped by a spathe or overlapping bracts
F. Inflorescences terminal: stamens 6... . 345... 53.0008 oe Cyanotis
F. Inflorescences terminal and axillary; stamens 3 .................... Commelina
E. Inflorescences more or less branched, not enveloped by a spathe or overlapping bracts
G. Fruit blue, smooth and shining, not dehiscent ..............- a. eee Pollia
G. Fruit whitish, dehiscent
H. Inflorescence densely flowered; flowers hairy, 6 stamens all fertile .......
oad aie aid isa ein ag w tyme Uo hie leah ot ald cdke\ee eee) mata ean Floscopa
H. Inflorescence fewer flowered, the branches slender; flowers not hairy, only
2' or 3.stamens are fertile 00) Pr) ok Murdannia
Commelina attenuata Koen.
Creeping herb, rooting at the nodes, with ascending, upright branches 20-30 cm
high; leaves linear-lanceolate, recurved, 1.5-2 cm long; sheath hairy; flowers in
cymes, enclosed in a green, folded, funnel-shaped spathe; peduncle 1 cm long;
flowers bright blue, stamens 3. In sandy sea-shores, Changi (Ridley 6003), Gelang.
Comm. benghalensis L.
Like the above species, but leaves elliptic or ovate, hairy; the leaf-tip blunt. In
waste places, Gelang (Ridley 3856), Tanglin.
Comm. diffusa Burm. f.
Also called C. nudiflora L. Like C. attenuata, but leaves ovate-lanceolate, glabrous.
Common in waste ground, Changi (Ridley 3935).
Cyanotis cristata D. Don
Prostrate herb, up to 30 cm tall, hairy; leaves oblong-lanceolate, 1.5-2.5 cm long;
sheathed below; inflorescence short, terminal, enclosed by overlapping bracts;
flowers pale blue. In sandy places near the sea, Changi, Gelang (Ridley s.n. in 1893).
Formerly called C. barbata D. Don.
Floscopa scandens Lour.
Slender creeping herb, ascending branches 15-30 cm high; leaves narrowly ellip-
tic, pointed at both ends, 5-8 cm long, shortly petioled; panicle terminal, 3-5 cm
long, with many ascending branches; flowers pink. In muddy ditches or damp
places, Tanglin (Daud 6007).
Forrestia gracilis Rid.
Stout herb, creeping shortly at base; erect stem slender, | m tall, glabrous; leaves
lanceolate, 15-20 cm long, the margin red-hairy; petiole winged; flower-heads small,
with few sessile, white flowers. In forests, Tanglin, Pulau Ubin (Ridley 48/0), Bukit
Timah. Another species, | marginata Hassk., with stout but creeping stems, was
recorded from Bukit Timah by Wallich. (Note: Correct name for the genus is
Amischotolyte).
‘Annotated List of Seed Plants in Singapore (XI) 125
Murdannia nudiflora (L.) Brenan
Formerly called Aneilema nudiflorum Wall. Creeping herb; ascending branches
slender, to 30 cm tall, leaves lanceolate, 2-8 cm long, the tip acute; sheath bearded;
cymes terminal, 3-5 cm long, branched at the tip; flowers small, violet, perfect
stamens 2. In waste ground and dry sandy places.
Murd. vaginata (L.) Brueckn.
Formerly called Aneilema vaginatum (L.) R. Br. Leaves linear, fleshy, 5-7 cm long.
In sandy places, Telok Kurau, Besut (Sinclair & Kiah S.F-N. 40785).
Pollia sorzogonesis Endl.
Stem slender, hairy, erect, 1 m tall; leaves lanceolate, the tip acuminate, 15-20
cm long; panicle terminal, pubescent, 20-30 cm long and wide, the branches
spreading; flowers small, white; perfect stamens 3. In forests, Pulau Ubin (Ridley
4759).
Rhoeo spathacea (Sw.) Stearn
Formerly called R. discolor Hance. Short-stemmed herb; leaves erect or ascend-
ing, sword-shaped, 15-25 cm long, green above and purplish beneath; flowers
_ white, small, in axillary clusters, enveloped by 2 boat-shaped bracts (‘‘Oyster
plant’’). Native of tropical America, cultivated as an ornamental. #Ss .
Tradescantia fluminensis Vell.
Ascending or creeping herb; leaves oblong, 4-8 cm long, green or white-striped
above, purple beneath; flowers white, in clusters. Native of S. America, this and
several other species are sometimes cultivated.
Zebrina pendula Schnizl.
The stem trailing along the ground, rooting at the joints; leaves oblong, 5-7.5 cm
long, alternate, with silvery green and purple stripes in the middle and around
the margin; reddish purple beneath. Native of Mexico, cultivated. fAft® .
161. FLAGELLARIACEAE
Key to the genera
A. Climbers; leaves lanceolate, the tip transformed into tendrils; flowers bisexual ........ Flagellaria
ie Paree erect heros; leaves normal; flowers umisexual 2.2.0.5. 6220 ee eee Hanguana
Flagellaria indica L.
Climber; stem slender but strong, 3-10 m long; leaves linear-oblong, 8-25 cm
long, usually ending in a short coiled tendril; flowers white, bisexual, in a widely
branched terminal panicle, 8-50 cm long. In open thickets near the sea, Kranji,
Bajau (Ridley 3589a), Changi. ## .
Hanguana malayana (Jack) Merr.
Formerly called Susum malayanum (Jack) Hook. f. Large erect herb, aquatic or
terrestrial, highly variable; leaves grass-like, tapering at both ends, 30-90 cm long,
densely nerved, rather thick; flowers unisexual, in a terminal panicle, 40-50 cm
long. In forests and in damp places, Bukit Timah, Seletar, Ponggol, Bukit Pan-
jang (Ridley s.n. 1907).
126 Gard. Bull. Sing. 40(2) (1987)
162. XYRIDACEAE
Xyris complanata R. Br.
Formerly called X. anceps Lam. Tufted herb; leaves linear, stiff, grass-like
reddish, 10-50 cm long; flower-head with many dark brown imbricate bracts, on
a stiff, slender, often twisted scape, 15-60 cm long; flowers yellow. In open sandy
places, Balestier Plain, Pulau Tekong, Changi, Geylang (Ridley 5757).
Xyris pauciflora Willd.
Like the above, but plants smaller and shorter (leaves 7-25 cm long) and the head
with fewer flowers. Balestier Plain (Ridley 5877), Water Catchment Areas.
163. ERIOCAULACEAE
Eriocaulon longifolium Nees
Herb with fibrous roots; leaves linear, grass-like, 8-30 cm long; scapes slender,
erect, 30-45 cm high, with a globose or ovoid, greyish head at the tip; flowers
minute, unisexual. Common in open damp places and ditches, Tanglin, Water
Catchment Areas, Jurong (Ridley s.n. in 1889). Formerly called EF. sexangulare L.
Eriocaulon truncatum Buch. Ham. ex Mart.
Smaller than the above species; leaves 3-5 cm long; scapes 4-10 cm high. Along
damp paths on sandy ground, Tanglin, Bukit Mandai (Ridley 3920), Water Catch-
ment Areas.
164. BROMELIACEAE
Ananas comosus (L.) Merr.
Perennial herb; leaves thick, linear lanceolate, 50-120 cm long, with sharp marginal
spines; flowers in a dense, leafy crowned head; scape stout, 0.5-1 m tall; many
small juicy fruits forming a spurious, broadly ellipsoid fruit (‘‘pine-apple’’
4 ), 10-30 cm long. Native to tropical America, many cultivated varieties, some
as ornamentals, mostly for the edible fruits. Several others of this family (also
from trop. America) are occasionally cultivated as ornamentals. These include:
Aechmea bracteata Griseb. (rosette-forming herb, with red, ribbon-like bracts on
the branched inflorescence), Pitcairnia integrifolia Ker-Gawl. (leaves forming
crowded rosettes; flowers scarlet, on a panicle arising from the leaf centre), and
Tillandsia usneoides L. (a rootless plant, with slender greyish hanging stem and
narrow, curved leaves 3-6 cm long, commonly known as ‘‘Spanish moss’’), etc.
165. PALMAE
Synoptic key to the genera*
A. Leaves palmate or palmately divided......... Licuala, Livistona (native and cult.), Rhapis (cult.)
A. Leaves pinnate or pinnately divided, (twice pinnate in Caryota)
B. Leaf-divisions folded back in bud, trough-like.....................-. Arenga (cult.) Caryota
B. Leaf-divisons infolded in bud, roof-like
C. Female inflorescence head-like, fruit cluster globose ..............-+++cseecsecsere Nypa
C. Inflorescences spike-like, often forming in panicles
D. Fruit covered with scales; leaves usually spiny
E; Erect palit 30 7s eee Metroxylon (cult.), Raphia (cult.) Salacca
* Only the native and commonly cultivated genera are treated.
Annotated List of Seed Plants in Singapore (XI) 127
E. Climbing rattans ...... Calamus, Daemonorops, Korthalsia, Myrialepis Plectocomis
D. Fruit smooth; leaves usually not spiny
TIRE RT UE ME CAO AMINA De ng Ea ina sz wie 5 diy aap Ds wip o nee ue vw eee acces
ie eit aan fe a od Archontophoenix, Areca, Cocos, Elaeis, Roystona (all cultivated)
ie Palms tormimerechmps ..../.2...........%. Chrysalidocarpus (cult.) Cyrtostachys,
Iguanura, Nenga, Oncosperma, Pinanga, Ptychoraphis, Ptychosperma (cult.)
Archontopheonix alexandrae (F. Muell.) Wendle. & Drude
Stem solitary, straight and slender, to 12 m tall (and less than 25 cm in diameter),
ringed; leaves pinnate, 2-3 m long, arched; leaf-divisions whitish beneath; in-
florescence 30 cm long, much branched, drooping. Native of E. Australia,
sometimes cultivated, known as ‘Alexandra palm’’.
Areca catecu L.
Stem solitary, slender and ringed, to 10 m high; leaves pinnate, 1.2-2 m long, the
lower ones usually drooping; inflorescence much branched; male flowers in one
row, with 6 stamens; females at the base of branches; fruit ovoid, 3-6 cm long,
orange or red. Native country uncertain, cultivated in villages. Sliced nut is wrapped
in a betel leaf (Piper betel L.) with a little dash of lime and chewed. Vern. Pinang,
betel nut, 4#% . (Note: the species name often misspelt as ‘catechu’).
Areca triandra Roxb.
Like the above, but male flowers arranged in 2 rows, with 3 stamens in each flower.
Native of tropical Asia, occasionally cultivated as ornamental.
Arenga pinnata Merr.
Stem solitary, to 20 m tall, commonly covered with old leaf-bases; leaves pinnate,
5-6 (— 10) m long, with black fibres at the base; inflorescence arising among the
leaves, pendulous; sugary liquid exuded from the cut of the young inflorescence;
fruit round, 4-5 cm across. Probably a native of eastern India, formerly called
Arenga saccharifera Labill. Vern. Kabong, sugar palm, Pts .
Calamus densiflorus Becc.
Clustering high climbing rattan with stem reaching 40 m tall; pinnate leaves 1.1 m
long, curved; leaflets up to 60 pairs; sheath bright to yellowish green, densely armed
with spines; male and female inflorescences axillary, superficially similar, with
tubular sheath (or spathe); fruit ovoid, 2 by I.2 cm, covered in rows of scales.
In forests, Gardens’ Jungle, Bukit Mandai (Ridley 6280). Vern. Rotan chichi.
Calam. diepenhorstii Miq.
Gardens’ Jungle, Bukit Timah (Ridley s.n. in 1907).
Calam. erinaceous (Becc.) Dransf.
Formerly called C. aquatilis Ridl. Ponggol, Holland Road, Gardens’ Jungle, Changi
(Ridley 6275). Vern. Rotan bakau.
Calam. insignis Griff.
Bukit Timah (Ridley s.n. in 1892), Gardens’ Jungle. Vern. Rotan batu.
Calam. javensis Bl.
Kranji, Bukit Timah (Ridley s.n. in 1896).
Calam. laevigatus Mart. ;
Formerly called C. pallidulus Becc. Jalan Bray (Ridley 628/), Bukit Timah, Bukit
Mandai.
Calam. lobbianus Becc.
Seletar, Bukit Timah (Ridley 896/), Chan Chu Kang, Pulau Damar.
128 Gard. Bull. Sing. 40(2) (1987)
Calam. lucidus Becc.
Formerly called C. laxiflorus Becc. Woodlands (Ridley 12607).
Calam. ornatus Becc.
Bukit Timah (Ridley 11581).
Calam. oxleyanus T. & B.
Bukit Timah (Ridley 11463), Tuas.
Calamus ridleyanus Becc.
Mandai (Ridley 3504, syntype), Gardens’ Jungle.
Caryota mitis Lour.
Clump-forming palm; stems to 4 m tall; leaves twice pinnate, 1-1.75 m long; leaflets
flat, all widening outward to a blunt tip, fish-tail like; male and female flowers
in same inflorescence; inflorescences both terminal and axillary, developing from
top downward, the stem slowly dying after flowering. Common in forests all over
the island. Vern. Zizkas, Fish-tail palm.
Chrysalidocarpus lutescens (Bory) Wendl.
Stems slender, ringed, 5-6 m tall, in clumps; leaves pinnate, 1-1.2 m long, with
yellow rachis; inflorescences among the leaves, male and female flowers in small
clusters; fruit ovoid, 3-4 cm long, yellow. Native to Madagascar; commonly
cultivated in pots or on ground, always remaining in vegetative state. ®BRT .
Cocos nucifera L.
Stem solitary, ringed, often curved and leaning, to 30 m tall; leaves pinnate, 2-6
m long, with numerous pairs of narrow.leaflets; flowers in branched spikes among
the leaves; female flowers near the base of the branches; fruit ovoid globose, more
or less trianglular, 20-30 cm across, edible. A very useful plant, widely cultivated
all over the tropics; true home uncertain, it has been suggested that it is a native
of the Indian Ocean. Vern. Kelapa, Coconut tree, + .
Cyrtostachys renda BI.
Formerly called C. lakka Becc. Clump-forming palm; stem erect, slender to 10
m or more tall, prominently ringed; leaves pinnate, to 1.5 m long; sheath scarlet;
one female flower flanked by 2 males, in panicles below the leaves; fruit small,
ellipsoid, 8 x 4 mm, with a narrowed top. Formerly found in wet places near
the sea, Kranji, Tuas, Chua Chu Kang, now extinct, but commonly cultivated in
gardens. Vern. Pinang raja, Sealing Wax Palm, S#4LeMy .
Daemonorops angustifolia (Griff.) Mart.
Clustering, high climbing rattan, with stem reaching 40 m tall; pinnate leaves
2-3.5 m long; sheath greenish brown, densely covered with triangular, black spines;
male and female inflorescences similar, terminal; spathe boat-shaped; fruit globose,
to 1.8 cm across, covered in rows of reddish brown scales. In forests, Chua Chu
Kang, Chan Chu Kang (Ridley 4622), Bukit Timah, Bukit Mandai. Vern. Rotan
sepat.
Daem. didymophylla Becc.
Mandai, Changi (Ridley 6273), Chan Chu Kang, Seletar, Gardens’ Jungle. Vern.
Rotan hudang.
Daem. grandis (Griff.) Mart.
Seletar, Ang Mo Kio, Bukit Mandai, Bukit Timah, Bukit Arang (Ridley 1660),
Gardens’ Jungle, Changi.
oc
Annotated List of Seed Plants in Singapore (XI) 129
Daem. hystrix (Griff.) Mart.
Bukit Mandai (Ridley 3480) Jurong, Sungei Murai, Gardens’ Jungle, Seletar,
Kranji, Tuas, Yio Chu Kang. Vern. Rotan sabite.
Daem. leptopus (Griff.) Mart.
Kranji, Bukit Mandai, Bukit Panjang, Jurong, Bukit Arang, Ang Mo Kio (Ridley
s.n. in 1894). Vern. Rotan bakau.
Daem. longipes (Griff.) Mart.
Seletar, Bukit Timah (Ridley 6902), Bukit Mandai, Changi, Chan Chu Kang. Vern.
Rotan sepah, Rotan chochor.
Daem. micracantha (Griff.) Becc.
Bukit Timah (Ridley s.n. in 1900).
Daem. periacantha Miq.
Bukit Mandai, Tuas, Bukit Timah (Ridley 3492).
Elaeis guineensis Jacq.
Short trunk of young trees covered with leaf-bases after the old leaves have been
removed; slow-growing old trunk to 3-10 m tall, often dented with leaf-scars; leaves
pinnate, arched, 4-5 m long, the lowest leaflets becoming thorns; cylindric male
and globose female inflorescences borne on the same plant; fruit orange-red, in
globose clusters 30-45 cm across. Oil from the fibrous oily pulp around the seeds
is used for cooking and industrial uses. Native to West Africa. Vern. Kelapa bali,
Oil Palm, ite .
Hyophorbe verschaffeltii Wendl.
Also known as Mascarena verschaffeltii L. H. Bailey. Trunk solitary, ringed,
3-4 m long, swollen above, not at the base (thus called ‘Bottle palm’); leaves pin-
nate, 1-2 m long; inflorescences below the leaves. Native of the Mascarene Islands.
Iguanura ferruginea Becc.
Small solitary or clustering undergrowth palm; stem very short or nearly absent;
leaves pinnate, 1 m long, with a much larger terminal leaflet; male and female
flowers crowded in groups on a simple or branched inflorescence. In forests,
Gardens’ Jungle, Bukit Timah.
Iguan. geonomaeformis Mart.
Stem 1-2 m tall; leaves bifid or 2-9-lobed, very variable. In forests, Bukit Timah,
Chan Chu Kang.
Korthalsia echinometra Becc.
Clustering high-climbing rattan; stem often more than 30 m tall; leaves to 1.8 m
long, with terminal thorny extension to 70 cm long; leaflets up to 25 pairs, narrowly
elongate, greyish white beneath; inflorescences crowded in axils of uppermost
leaves, to 60 cm long; fruit ovoid, 2.5 by 1.5 cm, covered by many vertical rows
of reddish-brown scales. In forests, Bukit Timah, Chan Chu Kang (Ridley 3521).
(Korthalsia is well-known as a genus of ant-plants. Ants inhabit the inflated ochrea
of the leaves).
Korth. grandis Rid.
Seletar (Ridley s.n. in 1894, lectotype), Bukit Panjang.
Korth. rigida BI.
Also called K. polystachya Mart. and K. wallichiaefolia Wendl. Bukit Timah (Ridley
6674).
Korth. scaphigera Griff. ex Mart.
Seletar, Woodlands, Bukit Timah (Ridley 6272). Vern. Rotan simut.
130 Gard. Bull. Sing. 40(2) (1987)
Licuala ferruginea Becc.
Small fan palm; stem very short; leaves divided to the centre; leaflets 10-13, the
central one the largest, to 1 m long; petioles about 1 m long, thorny at base; panicles
50-60 cm long; spathe brown, pubescent. In forests, Gardens’ Jungle. Bukit Timah.
Lic. grandis Wendl.
Slender palm, 1-2 m tall; leaves orbicular, to 1 m across, nearly entire except the
edges; petioles 1 m long, spiny. Native of New Hebrides and New Britain, often
cultivated.
Lic. spinosa Wurmb.
Bushy palm, in clumps, 2-5 m tall; leaves orbicular; leaflets 15-18, linear, 30 cm
long; petioles 1 m long, thorny the entire length; panicles erect or arching. In open
dry places or swampy areas, Pulau Ubin, Pulau Brani.
Lic. triphylla Griff.
Stemless; leaves divided into 3-7 leaflets, 20-22 cm long; petioles slender, 30-40
_ cm long, sparsely thorny. In forests, Changi.
Livistona chinensis R. Br.
Solitary palm; trunk stout, ringed, to 15 m tall; leaves fan-shaped, nearly rounded,
wider than long, 40-50 cm wide, divided to about the middle of the blade; petioles
spiny; flowers bisexual, in panicles; fruit ovoid (2 x 1.5 cm). Native of S. China,
planted. ## .
Liv. kingiana Becc.
Leaves 1.5 m across, with a few divisions down to the central rib, mainly in the
lower quarter; leaflets widening outwards, wedge-shaped; petioles massive, to 1.5
m long, spiny in lower parts; fruit globose, 5-6 cm across. In swampy forests,
Chan Chu Kang, Bukit Mandai. Formerly called Pholidocarpus kingiana Ridl.
Liv. rotundifolia (Lamk.) Mart.
Trunk to 20 m tall; leaves 1.2 m long; tips of leaf-divisions forked halfway down;
fruit globose, 2 cm across. Native of E. Malesia, cultivated.
Liv. saribus (Lour.) Merr. ex Cheval.
Trunk to 30 m tall; leaves to 1.2 m across, leaf-divisions almost reaching the cen-
tral rib; fruit globose, 1.5-2 cm across. Native of Indo-China and several parts
of Malesia, cultivated.
Metroxylon sagu Rottb.
Stout tree, forming big clumps, to 10 m tall; leaves pinnate, massive, to 5 m long,
with about 60 pairs of strap-shaped leaflets; flowers in large terminal, wide-
spreading panicle; male and female flowers mixed. Native to Moluccas and
W. New Guinea, formerly cultivated for the starch.
Mpyrialepis scortechinii Becc.
Climbing thicket-forming rattan, to 40 m tall; young stems with horizontal rows
of spines; whole leaf 3-5 m long, with terminal thorny extensions to 1.5 m long;
male and female inflorescences on separate plants, arising from the nodes of
uppermost reduced leaves; fruit depressed globose (3 x 2.5 cm), tipped with black
stigmas and covered with myriads of tiny scales. Bukit Mandai (Ridley 5860), Chan
Chu Kang, Kranji, Bukit Timah. Vern. Rotan kertong. Called Plectocomiopsis
annulatus Ridl. and P. scortechinii (Becc.) Ridl. in Ridley’s Flora.
Nenga pumila (Mart.) Wendl.
Also called N. wendlandiana Scheff. Tufted dwarf palm, 3-5 m tall; leaves pin-
nate, to 1.5 m long; leaflets linear, petiole 20 cm long, with a purplish sheath;
Annotated List of Seed Plants in Singapore (XI) 131
inflorescence usually 3-branched; fruit ovoid, 2-2.5 cm long, yellow to bright red.
In forests; Changi, Seletar. (Note: Nenga is very similar to Pinanga, but differs
from it in : 1) inflorescence with one spathe; 2) flowers always spirally arranged;
3) seeds laterally arranged).
Nypa fruticans Wurmb.
Rhizome very stout, creeping in mud; aerial stem very short; leaves pinnate,
5-6 m long, erect; inflorescence subterminal, erect, stalk 1 m long; fruits chestnut
brown, in a globose cluster, 20-25 cm across. In tidal river, often forming large
colonies. A very useful plant, leaves are used for roofing; sugar and alcohol can
be obtained from the flower spikes; and the endosperm of the seed is edible. Vern.
Nipah, *#% . (The generic name usually incorrectly spelt as Nipa).
Oncosperma horridum (Griff.) Scheff.
Tall slender palm, in clusters, with very dense crowns; stems to 20 m tall, covered
with massive downward directed black spines; leaves pinnate; leaflets horizontal
or nearly so; inflorescence large, below the leaves; flowers unisexual, spirally
arranged; fruit round, ripening black, 1.5 cm across, waxy. In inland forests, Bukit
Timah, Water Catchment Areas, Changi. Vern. Bayas.
Oncosp. tigillarium (Jack) Ridl.
Formerly called O. filamentosum BI. Like the above species, but (1) leaflets droop-
ing, and (2) often near the coast, standing in or near salt or brackish water. Tanglin,
Changi, Kranji, Vern. Nibong. (Note: There is relic stand of Nibong palms near
the main gate of the Istana).
Pinanga disticha Bl. ex Wendl.
Small forest undergrowth palm, often forming large clumps; stem very slender,
to 1.3 m tall; leaves V-shaped, 20-30 cm long, simple or occasionally pinnately
divided into broad or narrow leaflets; flowers in threes (with a female and 2 males),
on a simple spike, 10 cm long; fruit ellipsoid, about 1 cm long; ripening red. In
forests, Seletar.
Pin. limosa Ridl.
Usually solitary, 1-2 m tall; leaves oblong in outline, with deeply forked tip, about
30-40 cm long, entire or divided into irregular broad leaflets with curving sides.
In muddy spots in dense forests, Bukit Timah, Seletar.
Pin. malaiana (Mart.) Scheff.
In small clumps; stem 30-50 cm tall; leaves broad, spreading, 30 cm long, pin-
nate; leaflets with 2 main nerves. In forests, Chan Chu Kang.
Pin. simplicifrons (Miq.) Becc.
Tiny palm, often forming large clumps; stem slender, winding, usually less than
1 m tall; leaves entire, oblong, deeply forked, 20-25 cm long. In forest.
Pin. singaporensis Ridl.
Stem tufted, 2-5 m tall; leaves pinnate, 1 m long; leaflets about 13, sigmoid.
Formerly found in Bukit Timah, Bukit Mandai and Seletar, probably extinct
sometime ago.
Pin. subruminata Becc.
Slender palm, to 1.3 m tall; leaves simple, V-shaped. In forest, formerly collected
in Kranji.
Plectocomia griffithii Becc.
Solitary climbing rattan, to 50 m tall; whole leaf to 7 m long, including a terminal
thorny extension to 3 m long; sheath green, armed with combs of brown spines;
132 Gard. Bull. Sing. 40(2) (1987)
leaflets to about 60 on each sides; male and female inflorescences similar, terminal;
fruit globose, to 1.5 cm across, covered with rows of red-brown scales. Seletar,
Kranji, Bukit Mandai (Ridley 3470). Vern. Rotan dahan.
Ptychoraphis singaporensis Becc.
Slender palm; stems 2-3.5 m tall; 2-3 in small clumps; leaves pinnate, 1-1.7 m
long, with numerous leaflets; inflorescences 1-3, in leaf axils, 30-40 cm long; fruit
fleshy, red, ellipsoid, 1-2 cm long. In forests, Tanglin, Sungei Buluh, Tuas. Vern.
Kerintin, Rintin.
Ptychosperma macarthurii (Wendl.) Nichols
Trunks bamboo-like, to 6 m tall; leaves once pinnate, with many rather broad
leaflets (7-20 cm long, blunt). Panicles below the leaves, 40-45 cm long; female
flowers flanked by the males; fruit ellipsoid, orange, 1.2-1.5 cm long; seeds
5-grooved. Native of New Guinea, commonly cultivated in gardens. Seeds dispersed
by birds, naturalized. Another species, P sanderianum Ridl., with much narrower
leaflets, is occasionally planted.
Raphia farinifera (Gaertn) Hylander
Stem annulate, unarmed, robust, to 15 m tall; leaves pinnate, very large, up to
15 m long, leaflets of variable sizes, the larger ones 2.4-3.6 m long. Native of
Madagascar, formerly called R. ruffia (Jacq.) Mart.
Rhapis excelsa (Thunb.) Henry (= R. flabelliformis L. Hér.)
Trunks slender, straight, 2-4 m tall, in clumps, usually covered with fibrous leaf-
sheaths; leaves fan-shaped, about 30 cm across, dark green, shining; leaflets 5-11,
the ends blunt and toothed, rarely flowering under our climate. Native of S. China,
planted in gardens or treated as ‘dwarf plants’ in pots. tr . Another cultivated
species, R. humilis Bl., is a smaller plant, with smaller leaves of only about 6 leaflets.
Roystonea oleracea (Jacq.) Cook (= Oreodoxa oleracea Matt.)
Like R. regia, but much taller and the trunk is swollen only at the base, the crown
of leaves also spreads in all directions. Less commonly cultivated than R. regia.
Native of Barbados, called ‘Cabbage Palm’.
Royst. regia (H.B.K.) Cook (= Oreodoxa regia H.B.K.)
Trunk columnar, greyish, ringed, to 20 m tall, slightly swollen at the base and
middle; leaves pinnate, 3 m long, arched; male and female flowers growing along
spikes in a large panicle. Native of Cuba, often planted in rows as avenue trees,
noted for its stateliness and elegance, thus called ‘Royal Palm’, S#p+ .
Salacca affinis Griff.
Stems stout and short, sometimes appearing stemless, in clumps; leaves pinnate,
in big, dense rosettes, 3-4 m long; leaflets flat, in one plane, oblong, 30-40 cm
long; flowers unisexual, dioecious, in erect spikes subtended by large spathes; fruit
globose, about 2.5 cm across, beaked, scaly; seeds 1-3, enbedded in a juicy pulp.
In wet spots in forests, Bukit Timah, Chan Chu Kang (Ridley 4421). Vern. Salac.
(The generic name sometimes incorrectly spelt as Zalacca). Imported fruits from
Indonesia of a related species, S. edulis Bl., are sometimes sold in markets.
Sal. conferta Griff
Differs from S. affinis in having linear leaflets arranged in 2 planes, and male and
female flowers on the same inflorescence, forming a massive terminal head. In
swampy forests, often forming thickets, Tanglin, Bukit Timah, Changi, Chan
Chu Kang (Ridley 3143) Jurong. Vern. Asam payah, Salak hutan. (This plant dif-
fers from other species of the genus in having terminal inflorescences thus re-named
as Eleiodoxa conferta (Griff.) Burret. in some literature.)
Resolving Iron Deficiency in Wrightia religiosa
by Foliar Analysis and its Amelioration Using
an Iron Chelate as a Soil Additive
THAI WU FOONG and CHENG Nol YANG
Botanic Gardens, Parks and Recreation Department, Singapore.
Abstract
Iron deficiency in Wrightia religiosa was associated with a suboptimal level of ‘‘active Fe’’ in the young
leaves. Soil application of Nervanaid Fe 132, a carrier of FEEDTA, at 10 g per plant and particularly
at 20 g per plant helped chlorotic plants to regain their healthy vigour. Treatment brought the ‘‘active
Fe’’ to levels comparable to or above those of the control.
Introduction
Wrightia religiosa is a popular ornamental shrub in Singapore because of its
fragrant flowers and ease of propagation. It is also a popular choice for bonsai.
Recently, a noticeable population of Wrightia established in parks and along road-
sides has been affected by iron deficiency. Iron deficiency was confirmed by partial
recovery of deficient plants upon foliar treatment with FeSO,.
Iron deficiency symptoms manifest themselves initially as interveinal chlorosis of
the young leaves as iron does not move readily from old leaves to the young flushes.
Iron is required for chlorophyll synthesis and when it is present at suboptimal level,
insufficient chlorophyll is synthesized thus causing chlorosis (Plate 1). As deficiency
becomes more advanced and acute, the affected young leaves grow to become the
older affected leaves, while the newly emerged flushes remain chlorotic. Eventually,
the entire plant assumes an overall chlorotic appearance (Plate 2). In very severe cases,
the leaves become almost bleached of colour and have random necrotic spots. Dieback
of growing tips is common (Plate 3).
A spectrum of physical and chemical soil properties has been identified as con-
ducive to iron deficiency in plants (Chaney, 1984; Lindsay, 1984; Lindsay and Schwab,
1982; Mortvedt et al., 1977 Vejsadova et al., 1982). The iron chelate, FEEDTA, is
known to be an effective soil additive in overcoming iron chlorosis where the soil
PH is below 6.70 (Lindsay and Schwab, 1982). This investigation critically examined
the efficacy of this fertilizer in resolving iron chlorosis in Wrightia thriving on acid soil.
Materials and Methods
The trial area was located in a park where there was a concentration of iron-
deficient Wrightia plants. The sites were characterized by acidic soil pH values rang-
ing from 4.00 to 6.70.
Mention of a trademark or proprietary product does not constitute a guarantee or warranty of the
product by the Parks & Recreation Department, and does not imply its approval to the exclusion of other
products that may be equally suitable.
133
134 Gard. Bull. Sing. 40(2) (1987)
The iron chelate, Nervanaid Fe 132 (a.i. FEEDTA, 12.7% Fe) was tested at the rates
of 10 g per shrub and 20 g per shrub respectively. Five trial plants were used for
each treatment and control. The chelate was dissolved in 5 1 of tap water and then
poured evenly around the area spanned by the plant. In the case of the control, only
tap water was used. Treatment began at zero time after the first foliar sampling and
was thereafter maintained at weekly intervals for 4 weeks. Where Wrightia plants
existed in groups, each trial plant was selected in such a manner that it was separated
from the next by at least one guard plant so that the risk of spillover of treatment
effect was minimized.
From preliminary analyses, no relationship was discernible between total foliar
Fe content and iron chlorosis. In many cases, chlorotic plants were found to have
significantly more total Fe than their healthy counterparts. This was consistent with
the findings of Bennett (1945) and DeKock et al. (1979). However, the ‘‘active Fe’’
fraction extractable by 1N HC1 showed distinct differences between healthy and defi-
cient young leaves. This parameter was monitored during the course of treatment.
Foliar sampling was done three times i.e. at zero time, week 2 and week 4. Only
the first two pairs of fully-developed young leaves were sampled for analysis. These
were washed with Teepol (a non-ionic detergent), 0.1N HC1 and finally rinsed with
deionized water. Fresh subsamples were cut into small pieces with a pair of stainless
steel scissors, dried between filter papers and weighed out in duplicates. Each 1 g
sample was extracted with 20 ml. of IN HC1 for 24 hours with occasional agitation
(Takkar and Kaur, 1984). ‘‘Active Fe’’ was analysed by the PU9000 AA spectrometer
immediately after filtration.
Results and Discussions
Results were subjected to the Duncan Multiple Range Test for significance and
are presented in Table 1 and Histogram 1.
Table 1
Changes in the ‘‘active Fe’’ fraction in ppm in fresh young leaves
upon treatment with Nervanaid Fe 132
For each column: Values with the same letter are not significantly different at P < 0.01.
Before treatment at zero time, iron-deficient plants had significantly lower 1N
HC1 - extractable Fe or ‘‘active Fe’’ than the control plants (Table 1 and Histogram 1).
A typical iron-deficient Wrightia plant is depicted on Plate 1.
After two applications of the iron chelate, foliar analysis revealed that the
‘*active Fe’’ fractions in treated plants became comparable to those in the controls
at week 2. The significant-increase in ‘‘active Fe’’ in treated plants concurred with
a pronounced recovery from iron chlorosis. This was especially remarkable with the
20 g treatment. At this stage, the new flushes assumed the healthy green colouration.
Iron Deficiency in Wrightia religiosa 135
However, the older leaves remained chlorotic. The recovering process appeared to
take effect over the first and second week (Table 1 and Histogram 1).
At week 4, the treated plants had attained satisfactory recovery. With the 20 g
chelate treatment, the ‘‘active Fe’’ had significantly risen to 35.9% above that of
the control. ‘‘Active Fe’’ contents in the controls and plants treated with 10 g chelate
were similar although somewhat higher in the latter. (Table 1 and Histogram 1).
The longevity of treatment effect was monitored after the cessation of treatment
at week 4. Plants treated with 20 g chelate remained healthy for at least 6 months
whereas those treated with 10 g chelate reverted to the original state of iron chlorosis
after 2-3 months.
HISTOGRAM 1:
% Changes over control in the “active Fe
component in treated plants with time
50
o
>
2
225
Xe
100% : 2 ite
control sie Ba 4 week
|
Plate 1: Mild iron deficiency
in Wrightia religiosa
Plate 2: Advanced iron deficiency
in Wrightia religiosa
Plate 3: Dieback of growing tips
in advanced iron chlorosis
in Wrightia religiosa
136
Iron Deficiency in Wrightia religiosa 137
Conclusion
In agreement with the findings of Hellin et al. (1987) and Wallace et al. (1984),
the present study showed positive results in the amelioration of iron chlorosis in
Wrightia religiosa with FEEDTA as a soil additive where the soil pH was acidic. Both
levels of Nervanaid Fe 132 tested were found to be effective in correcting iron defi-
ciency. The higher dosage, however, should be used as its ameliorative effect was
more persistent.
It was possible to resolve iron chorosis in Wrightia by determining the tN HC1 -
extractable Fe or ‘‘active Fe’’. The likely sufficient level of this component that is
compatible with normal growth in Wrightia is about 20 ppm on a fresh weight basis
(Table 1).
Further investigations are in progress to resolve iron chlorosis in Wrightia by soil
analysis. The ultimate solution for iron chlorosis in Wrightia lies in determining
whether the deficiency is true or induced. If it is induced deficiency, the cause needs
to be ascertained so that suitable corrective measures could be administered
accordingly.
References
Bennet, J.P. (1945). Iron in leaves. Soil Sci., 60, 91-105.
Chaney, R.L. (1984). Diagnostic practices to identify iron deficiency in higher plants.
J. Plant. Nutr., 7(1-5), 47-67.
DeKock, P.C., Hall, A. and Inkson, R.H.E. (1979). Active iron in plant leaves. Ann.
Bot., 43, 737-740.
Hellin, E., Ureha, R., Sevilla, F and Alcaraz, C.F. (1987). Comparative study on
the effectiveness of several iron compounds in iron chlorosis correction in citrus
plants. J. Plant Nutr., 10(4), 411-421.
Lindsay, W.L. (1984). Soil and plant relationships associated with iron deficiency
with emphasis on nutrient interactions. J. Plant Nutr., 7(1-5), 489-500.
Lindsay, W.L. and Schwab, A.P. (1982). The chemistry of iron in soils and its
availability to plants. J. Plant Nutr., 5(4-7), 821-840.
Mortvedt, J.J., Wallace, A. and Carley, R.D. (1977). Iron, the elusive microelement.
Fertilizer Solutions, 21(1), 26, 28, 30, 32, 34, and 36.
Takkar, P.N. and Kaur, N.P. (1984). HC1 method for Fe*+ estimation to resolve
iron chlorosis in plants. J. Plant Nutr., 7(1-5), 81-90.
Vejsadova, H. and Lastuvka, Z. (1982). The importance of FeEkU 1 ior reversibility
and phosphate-induced chlorosis in maize (Zea mays L.). Biologia Plantarum
(Praha), 24(6), 401-406.
Wallace, A., Wallace, G.A., Wood, R.A. and Abou-Zamzam, A.M. (1984). Uptake
by plants of iron from iron chelates. J. Plant Nutr., 7(1-5S), 695-698.
{bRe1) jg % sopftev?
nit te re
Chemical Growth Retardation of Baphia nitida with PP333
LEONG CHEE CHIEW*
Parks & Recreation Department
Botanic Gardens, Singapore 1025
Abstract
PP333 (common name paclobutrazol), [(2RS, 3RS)-1- (4-chlorophenyl)-4, 4-dimethyl-2-(1 ,2,4-triazol-
1-yl-) pentan-3-01)] was tested on Baphia nitida hedges to evaluate its effects on growth. Results show
that growth of B. nitida was retarded without phytotoxicity symptoms at the rates of PP333 applied. Growth
retardation of up to six months was achievable at the PP333 rates tested, making chemical growth control
a useful tool for the maintenance of such hedges at reduced manual pruning and related labour costs.
Introduction
PP333 is a broad spectrum growth retardant which acts by inhibiting gibberellin
biosynthesis and reducing cell division and extension (Lever et al. 1982; Sugavanam
1984). It is xylem mobile and its effects include overall stunting of the plant, with
shorter internodes and darker green leaves (Sugavanam 1984); it also has good
fungicidal activity.
PP333 has been used on a variety of crops including graminaceous crops like wheat,
barley and rye (Froggatt et al. 1982) in stem shortening properties to reduce lodging,
and in fruit trees to reduce pruning requirements. In apples, growth retardation
became apparent about 1% months after treatment and shoot growth of treated plants
was about 48% that of the untreated (Lever et al. 1982). Ornamental greenhouse
crops like Begonia, Chrysanthemum, Hibiscus, Coleus etc were dwarfed by PP333
without any phytotoxicity symptoms (Shanks 1980).
Baphia nitida is widely planted in Singapore, being used for screening purposes.
Much labour is expended in manual pruning to maintain the desired height and lateral
spread of the hedges. This paper reports a study conducted in the Singapore Botanic
Gardens into the use of chemical growth retardation to control the vegetative growth of
B. nitida with the aim of reducing the frequency of manual pruning.
Materials and Methods
PP333 was tested at rates of 0 g (Control), 0.5 g, 1.0 g, and 2.0 g active ingredient
per plant. A randomised complete block design consisting of three blocks was used.
Each block contained four plots which were four stretches of established hedge, each
about 10 m long and containing 30 plants. Plots within each block were spaced
2 m apart. The treatments were randomly assigned to the four plots in each block.
*The author is currently with the Strategic Planning Division of the Ministry of National Development,
Singapore.
139
140 Gard. Bull. Sing. 40(2) (1987)
The retardant was applied as a soil drench. A shallow trench, 3-5 cm deep, was
dug around the base of each plant. PP333 at its desired rate, diluted in water, was
poured into the trench which was then refilled with soil. Each plant received 150 ml
of solution.
Only one application of retardant was carried out. The hedges were then left to
grow and were pruned at regular intervals by the Singapore Botanic Gardens main-
tenance workers regularly deployed for hedge pruning operations. Pruned foliage
were collected for dry weight determination. This was measured after the plant
materials were dried at 100°C until constant weight.
Results and Discussion
Pruning of B. nitida hedges was carried out 1, 3, 6 and 9 months after application
of retardant. Dry weights of foliage pruned from all plots in the three blocks are
shown in Table 1. As seen in Table 2 and Figure 1 there was no significant difference
between all treatment means at the first pruning one month after PP333 treatment.
Table 1
No. of Dry weight, kg
Sequence months
of after Block 1 Block 2 Block 3
pruning PP333
application
The trial started with the application of PP333 as a soil drench. Blocks 1, 2 and 3 were hedges in three
different lawns at the Botanic Gardens. T1, T2, T3 and T4 refer to treatments of 0.5g, 1.0g, 2.0g and
Og (Control) active ingredient per plant respectively. Pruning of hedges was carried out 1, 3, 6 and 9 months
after the trial started. Foliar material from individual plots were collected at each pruning for dry weight
determination.
Table 2
No. of Paclobutrazol, g/plant
Sequence months
of after PP333 2.0
pruning application enna
Dry weight readings in the 3 blocks/replicates were averaged. Going across the table, readings (at each
pruning) with the same alphabet are not significantly different at the 1% level.
MEAN DRY WEIGHT (kg)
] 3 5 7 9
NUMBER OF MONTHS AFTER PP333 APPLICATION
Fig. 1. Mean dry weight of foliage pruned during the duration of the trial; X - 0.5 g PP333/plant,
O - 1.0 g PP333/plant, @ - 2.0 g PP333/plant, A - control.
14]
142 Gard. Bull. Sing. 40(2) (1987)
In the second pruning though, three months after PP333 treatment, a sharp and signifi-
cant decrease in dry weights of foliage pruned from retardant-treated plots was
observed. Growth retardation was equally effective at all three rates of PP333 tested.
Subsequent prunings also yielded dry weight readings which were not significantly
different for all three PP333 rates but were significantly different from the control
readings. Visually a clear difference was seen between plant growth in control and
treated plots (Figure 2).
Plants treated with retardant began to show signs of breaking out of retardation
six months after the start of the trial. Although still compact in growth compared
to the control plants, new actively growing shoots began to be produced. An increase
in dry weight readings of foliage pruned from PP333 treated plots was found nine
months after the start of the trial (Table 2 and Figure 1). Again the dry weights of
foliage pruned from retardant-treated plots were not significantly different at all three
PP333 rates but were significantly different from control readings. Nevertheless even
the retardant-treated plots were growing considerably at this stage and were losing
their compact and neat appearance.
‘Statistically the results show that all three rates of PP333 tested were equally
effective in retarding B. nitida growth. However, visually it was seen that the higher
rates of 1.0 and 2.0 g active ingredient per plant maintained a more compact and
neater appearance of the plants for a longer period. To reduce chemical costs, the
lower rate of 0.5 g per plant, found to effectively control growth for up to about
six months, should be favoured.
To demonstrate the practicability of the 0.5 g per plant application rate, a 100 m
long stretch of B. nitida hedge consisting of 300 plants was treated with this rate of
Fig. 2. Differences in vegetative growth of retardant-treated and control plants. The length of hedge
on the right was given 0 g PP333/plant (control) while that on the left was given 0.5 g PP333/plant.
The picture was taken just before the second pruning, 3 months after the start of the trial.
*
Growth Retardation of Baphia nitida 143
PP333. The plants were pruned one month after retardant application and no fur-
ther pruning was subsequently done. Retardant application at the same rate was
repeated six months after the initial application. Figure 3 shows the condition of the
hedge one year after the trial began. The plants were compact in growth, had shorter
internodes, darker green leaves and required a light pruning at most to restore uniform
height for the hedge.
Fig. 3. PP333-treated hedge one year after retardant application. The hedges were treated with 0.5 g
PP333/plant applied as a soil drench; application was repeated six months after the start of the
trial. Except for one pruning one month after the first retardant application, no other pruning
was done.
No phytotoxicity symptoms were encountered throughout the trial following retar-
dant treatment. Application by soil drench was chosen because a preliminary trial
showed that it effected greater retardation than foliar spray. McDaniel (1983), and
Barrett and Bartuska (1982) also reported that soil drench was more effective than
leaf application for Chrysanthemum and Phaseolus. It appears that the reduced
effectiveness of PP333 applied to mature leaves suggests that it is less effectively
translocated through the phloem compared to the xylem.
Conclusion
The results reported here show that chemical growth retardation of PP333 is an
effective tool for reducing the need for manual pruning of B. nitida hedges. This
is expected to make maintenance of such hedges less manpower intensive, and when
compared to costs for manual pruning, less expensive.
144 Gard. Bull. Sing. 40(2) 987)
Acknowledgements
I wish to thank Mr Rehan bin Yusoff and Mr Ang Peter both of whom contributed
to this project through their technical assistance, and ICI (Singapore) Pte Ltd for
their gift of PP333 samples.
References
Barrett, J.E. and C.A. Bartuska (1982). PP333 effects on stem elongation depen-
dent on site of application. HortScience 17(5): 737-738.
Froggatt, P.J., W.D. Thomas and J.J. Batch (1982). The value of lodging control
in winter wheat as exemplified by the growth regulator PP333. In: Opportunities
for manipulation of cereal productivity. A.F. Hawkins and J. Jeffcoat (Eds).
BPGRG Monograph 7: 71-87.
Lever, B.G., S.J. Shearing and J.J. Batch (1982). PP333 — A new broad spectrum
growth retardant. Proc. 1982 British Crop Protection Conference — Weeds: 3-10.
McDaniel, G.L. (1983). Growth retardation activity of paclobutrazol on Chrysan-
themum. HortScience 18(2): 199-200.
Shanks, J.B. (1980). Chemical dwarfing of several ornamental greenhouse crops with
PP333. Proc. 7th Ann. Mtg. Plant Growth Regulator Working Group, 1980: 46-51.
Sugavanam, B. (1984). Diastereoisomers and enantiomers of paclobutrazol: their
preparation and biological activity. Pestic. Sci. 15: 296-302.
Growth and Yield of Mango cv Golek
in Java Over 25 Years
E.W.M. VERHEIJ
Department of Tropical Crop Science, Agricultural University, Wageningen.
Abstract
Trunk cross sections of selected ‘Golek’ clones increased almost linearly from the age of 5 to 25 years
at the high rates of 90 cm’ and 70 cm? per year for nucellar seedlings and grafted trees respectively.
Yields were extremely low and erratic and did not seem to affect tree growth. The highest yield levels
occasionally attained by individual trees amounted to 1 fruit per cm? trunk cross section.
The 2 best seedling trees were far superior to the rest and about as fruitful as the best clone; the latter
attained an aggregate crop of 3320 fruit and a trunk size of 1560 cm’, against average values of 1765 fruit
and 1580 cm? for the other grafted clones. Differences between clones were large but not sufficiently con-
sistent to substantiate them. The results suggest that a similar analysis over the full 45-year experimental
period (!) and including the clones of other cultivars in the trial, could be very worthwhile.
Introduction
The mango collection at the Chukurgondang experimental farm near Pasuruan
comprises about 200 cultivars: 72 from East Java, 64 from Central and West Java
and 66 from other countries. The collection is unique because of the long period of
observation (from 1941) and because tree growth has been recorded as well as yield.
At the time several fruit tree collections were established by the Agricultural Exten-
sion Service (Terra, 1948), to test the suitability of different species and cultivars.
Chukurgondang is situated in the centre of mango cultivation in East Java and the
mango collection goes down to the clonal level in order to establish whether the tree
population of a so-called cultivar is indeed uniform or not, and if not, whether clonal
selection within the cultivar is worthwhile.
At Chukurgondang the original record books from 1941 are still in use. To find
out whether a comprehensive analysis of this wealth of information might be worth
the effort, such an analysis was carried out for the clones of a single cultivar, “Golek’,
and over a 25-year period only. The results are presented in this paper.
Materials and Methods
Pitoyo et al (1982) gave a general description of Chukurgondang Station. The farm
is about 50 m above sea level. The area has a monsoon climate with a prominent
dry season: average rainfall (1960-1981) from May to November amounts to only
220 mm, against 1140 mm during the remaining five months.
The trees are planted at 14 m square and now form a closed stand. The soil is
sandy and deep, infertile, and with low water retention. Mango growers in the area
prefer somewhat better soils. There is no irrigation. The orchard is clean-cultivated;
fertilizers and manure have been applied mainly in the early years. The staff list several
pests and diseases which cause serious fruit losses in some years (Pitoyo et al, 1982):
145
146 Gard. Bull. Sing. 40(2) (1987)
mango hoppers, fruit flies, mealy bug, bats and birds, anthracnose and sooty mould.
The trees are too big for effective crop protection.
The selected ‘Golek’ clones in Chukurgondang are listed in Table 1. They are iden-
tified by a number and have all been propagated from a mother tree collection in
nearby Pasuruan.
Table 1
Number of trees of the 8 clonal selections of cv Golek,
year of planting and rootstock
Clone Number
Trees on ‘Madu’ stock, planted in 1941
Trees on ‘Madu’ stock, planted in 1956
Nucellar seedling trees, planted in 1956
In addition to the ‘Golek’ selections, the collection comprises somewhat smaller
numbers of ‘Madu’, ‘Arumanis’ and ‘Manalagi’ trees; numerous other cultivars are
represented by pairs of trees of one or two selections only.
Annual records for each tree include:
e trunk girth, for the grafted trees at 10 cm above and below the union and for the
seedling trees at the same height as the scion measurement;
e tree height and tree spread in the E-W and N-S directions;
¢ number and weight of fruit.
For this paper only the trunk girth and number of fruit per tree have been analysed,
over a 25-year period from planting. Girth was converted to trunk cross section (“basal
area’), assuming a circular cross section. Basal area brings out differences in size
between trees better than girth or diameter (Verheij, 1972); differences in tree weight
are generally somewhat greater than differences in basal area.
The trees of each clone are planted side by side; this arrangement and the low
tree numbers per clone precludes statistical analysis.
Results
Figure 1 depicts basal area growth and annual crop load for the three groups of
‘Golek’ trees listed in Table 1. As the trees grow older and bigger they can bear larger
numbers of fruit. To be able to compare crop loads irrespective of tree size, the crop
load has been expressed as the number of fruit per cm’ basal area. Only crop loads
exceeding | fruit per 8 cm? basal area are shown in Fig. 1.
The growth curves in Fig. 1 are rather straight. After a slight upswing in the
first few years, the seedling trees grow at a rate of about 90 cm’ per year against
around 70 cm? for the grafted trees. Initially all trees planted in 1956 grow equally
well, but after 5 years the superior vigour of the seedling trees becomes manifest.
The trees are extremely poor yielders. Clone 35 planted in 1956 comes out best,
but even for this clone in half the years the crop is too light to be shown in Fig. 1.
The seedling trees only produce a crop in excess of one fruit per 8 cm’ basal area
once in 25 years. It should be borne in mind that even a yield of one fruit per 2 cm?
basal area is not at all high. In apples for instance, yields exceeding | kg/cm? trunk
cross section are not uncommon (Verheij, 1972; Verheij and Verwer, 1972).
There is no indication in Fig. 1 of a relationship between yield and growth in the
sense that in a fruitful year basal area growth is reduced. Nor is there evidence of
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148 Gard. Bull. Sing. 40(2) (1987)
biennial bearing. Apparently the yield levels are too low to slow down growth. This
even applies to the most fruitful individual ‘Golek’ trees, which in some years pro-
duced about | fruit per cm’ basz: area. Unfortunately trunk girth was measured in
August, that is, between bloom and harvest. Consequently the effect of a heavy crop
on trunk growth is spread over 2 years and therefore hard to trace.
In Fig. 2 the cumulated numbers of fruit over 15 and 25 years are plotted against
basal area at these ages for all ‘Golek’ trees planted in 1956. The most fruitful trees
are those which produce the highest number of fruit in relation to tree size. These
trees have been marked by lines linking the values at 15 to 25 years.
This brings five outstanding trees to light: the three trees of clone 35 which stood
out in Fig. 1 already, along with two nucellar seedlings of clone 229. These two deviate
so much from the unproductive remainder that it is highly improbable that they belong
to the same population; they may for instance be sexual instead of nucellar seedlings.
The two superior seedling trees are in league with the grafted trees of clone 35,
which between them vary a great deal in size and yield, but form a trio through the
parallel course of the lines, indicating that from age 15 to 25 all three trees were equally
fruitful. The smallest grafted tree (< 1000 cm? basal area at 25 years) and the one
but largest seedling tree (> 2500 cm? basal area) are the two most fruitful trees! After
25 years both trees have borne an aggregate crop of 2.75 fruit per cm?’ basal area
cumulative
nr. of fruit
6000
best seedlings
2000
e e
e
% *e
a 4, low ~~ * yielding seedlings
>
- cm? trunk cross section
0 1000 2000 3000
Fig. 2. Cumulative number of fruit against tree size, 15 (4) and 25 (©) years from planting, for each
‘Golek’ tree planted in 1956. Lines link values at 15 and 25 years for the most fruitful trees.
Mango cv Golek in Java 149
(although up to the age of 15 years the small tree had done much better than the
large one in relation to its size).Planting density of the small tree would have to be
nearly 2% times as high to match the cumulated yield per ha of the large tree at 25
years. The difference in basal area suggests that a 2'2-fold increase in number of
trees per ha can indeed be accommodated. Higher yields per ha in the early years
and easier management of the small trees no doubt would compensate for the cost
of the extra trees.
In Fig. 3, growth and yield of all ‘Golek’ clones on ‘Madu’ stock are compared
at intervals of 6 years.
Since yields in the early years are most important for the grower, the clones in
Fig. 3 have been arranged in order of declining yield over the first 18 years; after
12 years the order was almost the same. For all clones planted in 1941 yields during
the 13th to 18th year — when the trees should have been at their best — were
particularly poor. Consequently the yield picture is largely determined by the crops
produced during the last six-year period. Only clone 35 planted in 1956 was most
productive in the 13th to 18th year. These trees have borne the largest aggregate crop,
not only over 24 years, but throughout. On the other hand, trees of the same clone
planted in 1941 produced less than half as much. Because of this large variability
which was also found between trees within some of the clones, it is impossible to
substantiate the observed differences between the clones.
Looking at the lower half of Fig. 3 it is clear that the differences between clones
in the rate of growth were quite large. During the first six years, trees of the most
vigorous clone grew three times as fast as trees of the weakest clone; the same holds
for the last 6-year interval. However, the clones are not very consistent in their growth
rates, so that the differences in tree size after 24 years are not so large. Clone 177
had the largest trees (of similar size as the seedling trees), clone 229 the smallest.
Fig. 3 also shows that there may, after all, be some interaction between tree growth
and yield. This is revealed by the drop in tree size ranking with increasing age for
the more productive clones on the left hand side, as against rank gains for the least
productive clones on the right side. In other words: tree growth of the better yielding
clones gradually lags further behind that of the lowest yielding clones. This trend
is remarkably consistent. The logical explanation is that fruiting moderates growth,
but the reverse — moderate growth promotes fruiting — cannot be ruled out.
Discussion
In retrospect, the experiment was of course premature. The confusion over varia-
tions between and within mango cultivars was — and still is — a real problem.
However, the role of virus diseases and other endemic infections has to be clarified
before clonal selection can make headway. In this respect little progress has been
made in the 45 years since the trial started.
The experiment is also much too weak to yield convincing conclusions in respect
of the clones. So what is left is a unique long-term record of growth and yield of
‘Golek’ trees. Unfortunately the yields have been very low, in fact too low to affect
growth sufficiently to study the relationships between growth and yield. Administrative
problems during wartime and early independence may have left room for pilferage
of fruit or casual recording. However, the consistency of the data per tree from year
to year suggests that the records are generally sound and that the low yields are real.
The one serious omission is the lack of information on flowering, fruit set and
early fruit drop, so that after all these years one can only speculate on the reasons
for the poor crops. Probably paucity of bloom and destruction of inflorescences by
pests (mango hoppers, tip borers) and diseases (anthracnose) are the main reasons
nr of fruit
cumulative 28 Fane
3000 ,
NY"
\
2000 \
\
N
N
\
1000 .. -
“aN ®
Clone nr: 35/5631 133 33 177, 35/41 229 «195 = 255
om | fee Bol ee om
NN :
1000 \ \ \
PS EZZARNNEE
ee tea
2000
Fig. 3. Growth and yield of ‘Golek’ clones on ‘Madu’ stock.
Top: Mean number of fruit per tree, cumulated over 6-year periods.
Bottom: Mean trunk cross section per tree at 6-year intervals; numbers in the columns indicate
ranking for tree size: 1 = smallest; 9 = largest.
150
Mango cv Golek in Java 151
for crop failure. While gathering the data for this paper in 1983, most inflorescences
withered away owing to tip borer damage.
There are indications, at least in some cultivars, that twigs which have flowered
will not flower the following year, irrespective of fruit set (Lal Singh and Abdul Aziz
Khan, 1939). If this is the case, destruction of a heavy bloom by pests or diseases
may therefore decimate the next crop as well as the current crop. Moreover crop failure
invites untimely flushing, so that the terminal buds may not be receptive at the time
of flower induction in the following year. Verheij and Snijders (1986) describe a similar
sequence of events to explain the poor and erratic cropping of clove trees; picking
the clove buds also deprives the tree from the stabilizing effect of a load of fruit on
the growth rhythm. Current research in Australia aims at solving the riddles posed
by erratic flowering in mango (Scholefield, 1983).
The seemingly straight growth curves of the trees probably should be interpreted
as having the shape of a flattened S, because that is the common shape of trunk growth
curves in forestry. Extending the curves up to the present day (45 years for the 1941
planting!) may settle this point. Westgarth and Buttery (1965) report the results of
a spacing experiment with rubber trees over a 30-year period. Converting their girth
data into basal area also produces flattened S-curves. The highest growth rates were
attained at 5-15 years and ranged from 66 to 14 cm? per tree per year for the widest
and closest spacings respectively. In comparison the mango growth rates of 90 and
70 cm? basal area increments are quite high.
It is concluded that the results of fast-growing, poorly cropping ‘Golek’ trees form
a useful basis of reference, against which improvements in the yield : growth rela-
tionship, obtained under more intensive husbandry, can be measured. A further
analysis of the Chukurgondang data is recommended, to cover the full 45-year period
of recording and to compare growth and yield of ‘Golek’ with that of other leading
cultivars.
Acknowledgements
The Director of the Malang Research Institute for Food Crops, Dr. Soetarjo
Brotonegoro, kindly granted permission to use the field records and the Chukurgon-
dang staff have been very hospitable and helpful.
References
Lal Singh and Abdul Aziz Khan, (1939). Relation of growth to fruit bearing in
mangoes. Indian J. Agric. Sci. 9 (6): 835-867.
Pitoyo, M., Muljono, S. and Schild, D.J., (1982). Survey-inventory and proposals
on the upgrading of the sub-station Chukurgondang. Mimeo report, Malang Res.
Inst. for Food Crops: 36 p.
Scholefield, P.B., (1983). Mango research in the northern part of Australia. Paper
presented at International Workshop on Promoting Research on Tropical Fruits,
Jakarta, May 30 - June 6. Mimeo. 13 p.
Terra, G.J.A., (1948). Horticulture in: Agriculture in the Indonesian Archipelago
Vol. IT. Ed. Hall, C.J.J. van and Koppel, C. van de. A. van Hoeve, the Hague:
622-746. Dutch.
Verheij, E.W.M., (1972). Competition in apple, as influenced by Alar sprays, fruiting,
pruning and tree spacing. Meded. Landbouwhoge-school Wageningen, 72-4: 54 p.
152 Gard. Bull. Sing. 40(2) (1987)
Verheij, E.W.M. and C.H.A. Snijders, (1986). Syzygium aromaticum (L) Merril and
Perry in Plant resources of South East Asia; Proposal for a handbook. Eds.
Westphal, E. and Jansen, P.C.M. PUDOC, Wageningen: 61-66.
Verheij, E.W.M. and Verwer, F.L.J.A.W. (1972). Yield-density relations for apple
trees on a dwarfing and a semi-dwarfing rootstock. Neth. J. Agric. Sci. 20: 58-66.
Westgarth, D.R. and Buttery, B.R. (1965). The effect of density of planting on the
growth, yield and economic exploitation of Hevea brasiliensis. Part 1: The ef-
fect on growth and yield. J. Rubber Res. Inst. Malaysia 19 (1): 62-73.
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Items 1-4 can be purchased from the Commissioner, Parks & Recreation Department,
Botanic Gardens, Cluny Road, Singaore 1025; tel. no. 4709917.
For overseas orders, payment should be by bank draft or International Money Order and made
L ayable to the Commissioner of Parks & Recreation, Singapore.
Items 5-7 can be purchased from Singapore National Printers (Pte) Ltd, 303 Upper Serangoon
Road, Singapore 1334, tel. no. 2820611 ext. 125, 105 or 118.
*
All prices quoted are in Singapore Dollars
Overseas postage is extra
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