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THE

GARDENS’ BULLETIN

SINGAPORE

Volume XXVII

1974

A periodical reflecting the interests

and activities of the Botanic Gardens

Singapore

EDITOR

Chang Kiaw Lan

To be purchased at the Botanic Gardens

Cluny Road, Singapore 10

Price: $$26.50

Published by Botanic Gardens

Parks & Recreation Division

Public Works Department

Printed by the Singapore National Printers (Pte) Ltd

)

Digitized by the Internet Archive ©

in 2014

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httos://archive.org/details/gardensbulletins427unse

CONTENTS

Vol. XXVII

PART I — 15th February, 1974 Pages

CORNER, E. J. H.: Boletus and d Phylloporas i in ee further notes and

descriptions - - - - - - 1-— 16

ANTHONY, Ming: Contribution to the knowledge of Cecidia of Singapore - - 17-— 65

KENG, Hsuan: Annotated list of seed plants of Singapore (II)_ - - - - 67— 83

CHOW, K. H: Morphology and ecology of some introduced herbaceous legumes - 85— 94

CHU, F. Fei-Tan: Anatomical Features of the Dipterocarp Timbers of Sarawak - 95-119

REVIEW: Palms of Malaya (Hsuan Keng) - - - : - - - 121-122

KUEH Tiong-Kheng: New Plant Disease Records for 1972 - - - - 123-129

REVIEW: Animal Life and Nature in Singapore (Lee Chu San) _ - - - - 131-132

SINCLAIR J.: The genus Horsfieldia (Myristicaceae) in and ees Malesia I:

H. sabulosa and H. whitmorei J. Sinclair spp. nov. - . - - 133-141

PART II — 30th December, 1974

HOLTTUM, R. E.: Asplenium Linn., sect. Thamnopteris Presl - - - 143-154

—: A Commentary on Comparative Morphology in Zingiberaceae- - - - 155-165

—: The Tree-ferns of the genus Cyathea in Borneo - ets - : - - 167-182

GEH, Siew Yin et KENG — protynologsal catia on some inland |

Rhizophoraceae - - - - a oe 183—220

Report of the Botanic Gardens for the year 1973 - : - . - - 221-245

KENG, Hsuan: Annotated list of seed plants of Singapore (III) - - - - 247-266

INDEX a: : : : : : é : E ee Bed ITT

INDEX, VOL. XXVII

Basionyms and synonyms appear in italics, new taxa, names and status in

boldface. Plates are expressed by page numbers with letters.

orchioides 159

Abrus precatorius 254 pterocalyx 159

Acacia 123

auriculiformis 254 Alseodaphne bancana 78

inci Alsophila 168

cincinnata 254 ae he

farnesiana 254 australia |

burbidgei 180

mangium 254

pseudo-intsea 255

Acalypha wilkesiana 34

Achasma 156, 157, 159, 160-164

macrocheilos 162 ,

megalocheilos 163 frichodesma 179

pauciflorum 162 vexans 174

spaerocephalum 162 Alternaria sp. 128

ahieetincum 162 brassicicola 124, 127

Actinodaphne sp. 20, 24, 38, 39, 60, 65G Sct

glomerata 78 Alysicarpus vaginalis 255

contaminans 174

latebrosa 173

var. ornata 172

ramispina 173

hullettii 78 :

: 4 Amanita 1, 2

ee 18 . Amherstia nobilis 255

malaccensis 78 + 158. 159

pruinosa 78 mmomum i nee 161

Adenanthera bicolor 255 ae ,

pavonina 255 nial 162

Adinobotrys erianthus 262 ie

Aeschynomene indica 255 Retiscen GR

Afzelia bijuga 262 Nate =

retusa 262 es oe

Agathis alba ssp. borneensis 135 enorme

Agelaea borneensis 252 Andira Bee 255

hullettii 252 Angelesia splendens 250

macrophylia 252 Anisophyllea 183, 186, 190, 192, 193, 194

trinervis 252 corneri 187, 193

vestita 252 distiha 44, 65L, 184-186, 203, 204, 209, 211, 212

wallichii 252 Anisoptera 98, 99, 100, 102, 103, 104, 106, 110, 111

Ageratum 123 grossivenia 98, 115

Agromyza phaseoli 92 laevis 98, 99, 115

Albizzia falcata 255 marginata 98, 115

falcataria 255 reticulata 115

ittoralis 255 Annona 68

moluccana 255 muricata 70

pedicellata 255 reticulata 70

retusa 255 squamosa 70

Allium 123 Annonaceae, key to genera 68-69

Allospori 8 Aporosa benthamiana 34, 53, 65K

Alphonsea 68 Arachnis hypogaea 255

maingayi 69 Aralia triloba 81

Alpinia 158, 160, 161 Areca 123

alliance 163 Aromadendron 67

allied genera 157 elegans 67

allughas 163 Artabotrys 69

galanga 163 costatus 70

267

268

maingayi 70

odoratissimus 70

suaveolens 28, 65A, 70

unicinatus 70

wrayi 70

Artocarpus 124

Asbolisia sp. 125

Ascochyta phaseoiorum 128

Asparagus 124

Asplemium, sect. Thamnopteris 143, 144, 150

key 145

amboinense 154

antiquum 146, 151

antrophyoides 147, 154

australasicum 144, 145, 147, 148, 149

carinatum 146, 151

colobrinum 146, 151

var. taeniophyllum 152

curtisorum 148

cymbifolium 146, 150, 151

forma lingganum 151

ficifolium 147

grevillel 147, 153

humbertii 147, 154

longum 144, 146, 150

marinum 143

musifollum 140, 149, 151

nidus 143-145, 147, 149

var. australasicum 144

var. biseriale 146, 148

var. curtisorum 146, 148

var. multilobium 146, 149

var. musifolium 144, 149

var. nidus 143, 146, 148

var. nidiforme 149-150

var. plicatum 146, 149

var. sinuosianum sensu Chr. 153

oblanceolatum 153

pachyphyllum 154

pacificium 145, 147

perlongum 150

phyllitidis 144-146, 150, 152

ssp. malesicum 147, 153

ssp. phyllitidus 146, 152

phyllitidis sensu Holtt. 153

simonsianum 146, 152

simplex 154

simplex sensu Kunze 153

spirale 144, 146, 150

subspathulatum 154

squamulatum 154

taeniosum 154

tenerum 143

trichomanes 143

Atuna excelsa 250

Baccaurea griffithii 34

Barclaya motleyi 83

Barringtonia 186

Bassia 124

Bauhinia acuminata 256

blakeana 256

flammifera 256

griffithiana 256

monandra 256

Gardens’ Bulletin, Singapore — XX VII (1974)

purpurea 256

semibifida 256

tomentosa 256

variegata 256

Begonia sp. 124

Beilschmiedia kunstieri 78

malaccensis 78

Bixa 124

Boesenbergia 157, 161, 164

Boletellus purpurascens 8

Boletinus squarrosoides 15

Boletus 2, 4, 5, 8

subgen. Austroboletus 2, 8

subgen. Boletellus 3, 8, 9

subgen. Boletus 3

subgen. Leccinum 2, 6

subgen. Punctispora 8

subgen. Tylopilus 9, 10, 14

subgen. Xerocomus 11, 15

aculifer 14

albipurpureus 11

albo-ater 10

ananas 3

anripes 6

ballouii var. fuscatus 9

betula 2, 8

borneensis 2, 7, 8

cervicolor 10

chlamydosporus 4

coccineinanus 9

destitutus 11

dictyotus var. morchellipes 2

ferruginospora 11

funerarius 10

graveolens 7

hastulifer 14

hibiscus 2, 3, 4

junghuhnii 12

longicollis 3

longipes 9, 10

longipes var. alba 10

var. latiporus 9

lubricus 5, 8

malaccensis 2

mucosus 2

nanus 10, 11

nugatorius 12

olivaceiluteus 14

peronatus 6, 7, 13

portentosus 2, 4

punctisporus 8, 13, 14

raphanolens 11

rubiicolor 3

rubroglutinosus 4

rufo-aureus 5

satisfactus 11

schichianus 8

setigerus 2, 12, 13, 14

sinapicolor 5, 6

squarrosipes 7, 8, 13

subreticulatus 5

sylvestris 7

thibetanus 6, 11, 12

tristior 10

Index

tropicalis 4

variisporus 11

viridis 10

xylophilus 6

Borreria 124

Botryodiplodia theobromae 123, 125, 126, 128

Bougainvillea 122

Brassica 124

alboglabra 248

chinensis 248

juncea 248

oleracea 248

pekinensis 248

Brownea ariza 256

Bruguiera 4, 183, 184, 186, 187, 194, 195

Bryophyllum calycinum 249

Caesalpinia bonduc 256

crista 256

globulorum 256

Nuga 256

pulcherrima 257

sappan 257

Calliandra brevipes 257

haemacephala 257

Calophyllum ferrugineum 23, 36, 38, 58, 60, 65E

inophylloide 24, 59

var. singaporense 37

inophyllum 37

javanicum 37

pulcherrimum 27, 37, 38, 57, 58

Cananga 69

odorata 70

Canangium 124

odoratum var. fruticosa 70

Canarium 29

hispidum 29

pilosum 29, 30, 65E

Canavalia catharica 257

ensiformis 257

gladiata 257

maritima 257

obtusifolia 257

turgida 257

Cantharellus 1

Capnodium moniliforme 125

Carallia 183, 192, 194, 196

brachiata 187, 188, 190, 192, 195, 202A, 202B,

213, 205

eugenioidea 187, 193

Carenophila 164

Carex 164

Carica 124

Cassiaolata 257

fistula 257

fructicosa 257

hirsuta 258

mimosoides 258

nodusa 257, 258

obtusifolia 258

occidentalis 258

siamea 257, 258

tora 258

Cassytha filiformis 78

Castanopsis wallichii 35, 65F

269

Catimbium 157, 161, 163

Celosia 124

Cephalcuros virescens 124, 125, 126, 127

Ceratophyllum demersum 83

Cercospora sp. 125, 127

celosiarum 124

citrullina 124, 125

erechtitis 125

gerberae 126

hydrangeae 126

kikuchii 126

mori 127

phyllanthicola 128

Ceriops 183

candolleana 195

cucurbitarum 123, 126, 127, 128

Chrysanthemum 124

Cinnamomum camphora 79

deschampsii 79

iners 79

javanicum 79

ridleyi 79

verum 79

zeylanicum 79

Citrullus 124

Citrus 125

Cladosporium sp. 125

cladosporioides 126

oxysporum 124

Clematis dioscoreifolia var. robusta 82

Clerodendron 45 |

deflexum 45

Cnestis platantha 252

Coccomelia nitida 250

Cocculus trilobus 82

ovalifolium 82

Cochliobolus geniculatus 124, 128

Cochliobolus lunatus 126, 128

Cocos 125

Coelodepas glanduligerum 34, 65K

Colletotrichum sp. 125

capsici 127

circinans 123

graminicola 126

lindemuthianum 128

Combretocarpus 183, 190, 194, 196

rotundatus 184, 185, 187, 196, 202B, 206, 214

Conamumum 164

Coniothyrium fuckelii 127

Connarus

ferrugineus 252

grandis 252

monocarpus ssp. malayensis 252

oliogophyllus 252

semidecandrus 252

Corticium rolfsii 125, 127, 128

salmonicolor 123

solani 125, 128

Corynespora cassiicola 126, 128

Coscinium wallichianum 82

Cotylelobium 115

Cotylelobium spp. 99, 100, 102, 103, 104, 106, 110,

TK FEZ

burckii 115

270

malavanum 11S

melanoxyvlon LS

Crotalaria alata 258

bialata 258

mucronata 258

quinquetolia 258

retusa 258

sultiana 258

Cryptocarya sp. 39

caesia 79

ferrea 79

griffithiana 79

impressa 79

kurzii 79

Cucurbita 124

Curcuma 125, 158, 159, 164

Curvularia leonensis 127, 128

borreriae 124

eragrostidis 123, 127, 129

penniseti 123

senegalesis 124, 129

Cyathea 167-169

subgen. Alsophila 169

sect. Alsophila 170

key 170, 181

sect. Gymnosphaera 170

key 173

subgen. Cyathea 168, 169

subgen. Sphaeropteris 169, 170

sect. Schizocaena 170

key 175

sect. Sphaeropteris 168, 169, 170

key 174

subsect. Fouriniera 170

subsect. Sphaeropteris 170

key 174

acanthophora 167, 170, 171

agatheti 176, 180

alternans 175, 177

ampla 180

angustipinna 175, 177

arborea 168

arthropoda 175, 177

assimilis 176, 178

borneensis 171, 172

brachyphylla 181

brunois 177

burbidgei sensu Chr. 179, 180

capitata 175, 177

contaminans 174

discophora 167, 176, 179

elliptica 178, 180

incisoserrata 170, 172, 173

fuscopaleata 177

glabra 173, 174

havilandii 167, 170, 171, 182

Kembarangana 180

kinabaluensis 177

korthalsii sensu Chr. 172

latebrosa 171, 173

leucothricha 174

loheri 168, 170, 172, 181, 182

longipes 167, 170, 171

megalosa 167

Gardens’ Bulletin, Singapore — X XVII (197-4)

megalosora 176, 179

moluccana 177, 178

obtusata 172

oosora 170, 171, 181, 182

polypoda 176, 180

pscudobrunonis 177

ramispina 173, 174

recommutata 173

Squamulata 176, 178, 180

stipitipinnula 167, 176, 178

trichodesma 176, 179, 180

trichophora 175, 176, 178

tripinnata 174, 175

vexans 174

wallacei 180

Cyathocalyx 69

maingayi 73

ramuliflorus 28, 70

ridleyi 71

sumatranus 71

virgatus 72

Cyathostemma 68

hookeri 71

Viridiflorum 71

Cyclea laxiflora 82

Cynometra cauliflora 259

ramiflora 259

Dalbergia candenatensis 259

hulletii 259

junghuhnii 259

parviflora 259

rostrata 40, 651, 259

scortechinii 259

torta 259

velutina 259

Delonix regia 259

Derris amoena 259

elliptica 259

heptaphylla 260

heterophylla 260

scandens 260

sinuata 260

thyrsiflora 260

Desmos 69

chinensis 71

dasymaschala 68, 71

dumosus 71

Dehaasia incrassata 79

microcarpa 79

nigrescens 79

Desmodium 85, 87, 125

aparines 88

canum 85, 86, 88A

heterocarpon 260

var. ovalifolium 260

heterophyllum 86, 260

intortum 85, 87, 88, 88A, 89, 90

polycarpum 260

sandwicense 85, 88, 88B, 89

triflorum 86, 260

umbellatum 260

uncinatum 85, 88, 88B, 89, 90

Dialium laurinum 260

maingayi 40, 260

Index 271

wallichii 261 indica 261

Dichapetalum sordidum 250 orientalis 261

Dicksonia mollis 167 parcelli 261

Dieffenbackia 122, 125 subumbrans 261

Dillenia suffruticosa 4 Mtlingera 163

Digitaria 125 Hugenia 27, 42, 126

Dipterocarpaceae cumingiana 42, 43

anatomical data 113, 114 malaccensis 43

density of timbers 115—118 rugosa 43, 64

Dipterocarpus 101, 115 subdecussata 43, 65J

Dipterocarpus spp. 99, 100, 101, 102, 103, 104,106, — |agraca fragrans 40, 65J

110,111 libraurea chloroleuca 82, 83

acutangulus 115 licus 126

applanatus 115 Ficus spp. 42

borneensis 101, 115 pumila 41, 65

caudiferus 115 recurva 41

confertus 115 retusa 41

conformis 115 sinuata ssp. sinuata 42, 65H

coriaceus 115 l‘issistigma 69

costulatus 115 fulgens 71

crinitus 115 lanuginosum 71

cuspidatus 115 latifolium 71

eurynchus 115 manubriatum 72

exalatus 115 Flemingia strobilifera 263

fagineus 115 Fomes lignosus 126, 129

geniculatus 115 Fragaria indica 250

globosus 115 Friesodielsia 68

?gracilis 24, 115 biglandulosa 72

humeratus 115 . glauca 72

lowii 115 latifolia 72

mundus 115 Fusarium coccophilum 125

nudus 115 oxysporum 128

oblongifolius 115 solani 127

pachyphyllus 115 Gall-makers

palembanicus 115 Amorphococcus sp. 37

penangianus 115 Beesonia 32

rigidus 115 Bruggmanniella 37, 40

sarawakensis 115 Eriophyes gyrographus 39

stellatus 115 Eriophyes sp. 29, 38, 42

tempehes 115 reijnvannae 29

verrucosus 115 Gallacoccus 23

Dolichos lablab 261 anthonyae 23, 33

Dracaena 125 secundus 23, 24, 30

Drepananthus ramuliflorus 70 Leptynoptera sulphurea 37

Dryobalanops 115 Megatrioza vitiensis 43

Dryobalanops spp. 100, 102, 106, 110 Nothopoda pauropus 46

aromatica 102, 115 Pediobius 32

beccarii 115 Platygaster 38

fusca 103, 115 Gastrochilus 164

lanceolata 102, 115 Geonthus 163

rappa 102, 103, 115 coccineus 163

Duchesnea indica 250 Geostachys 159, 164

Durio 125 Gerbera 126

Elaeis 125 Gliricida sepium 261

Elettaria 125 Globba 158, 161

Ellipanthus griffithii 252 Glomerella cingulata 123, 125, 126, 127, 128, 129

tomentosus 252 Glycine 126

Ellipeia cuneifolia 71 wightii 85,91, 92A

Endiadra sp. 39, 61 Goniothalamus 68

Entada spiralis 261 malayanus 72

Enterolobium saman 265 tidleyi 72

Entoloma 1 tapis 72

Erechtites 125 Grewia blattaefolia 27, 45, 65L

Erythrina fusca 261 Gymnacranthera 75

baneana 75

eugenifolia 75

var. cugeniifolia 42

torbesii 75

Gvinnosphaera glabra 174

Gynotroches 183.190, 192, 193, 194

a\ilaris’ 184,188, 187, £96. 202) 207.9200, 215

Gyrodon 4

Gyroporus I4

castaneus 14

Heimieclla 8, 14

japonica 14

mandarina 5

retispora 9, 14

subretispora 9

Helicotylenchus dihystera 124

Hemitelia alternans 177

Hernandria cordigera 81

nymphaeifolia 81

ovigera 81

peltata 81

Hibiscus 126

tiliaceus 4

Hopea 115, 116

Hopea spp. 100, 103, 104, 106, 110

aequalis 115

altocollina 115

andersoni 115

argentea 115

beccariana 103, 115

bracteata 115

centipeda 116

cernua 116

dasyrrhachis 116

dryobalanoides 116

dyeri 116

enicosanthoides 116

fluvialis 116

eriffithii 116

latifolia 116

megacarpa 116

mesuoides 116

micrantha 116

nervosa 103, 116

nutans 103, 116

pachycarpa 116

pedicellata 116

pentanervia 103, 116

pterygota 116

sangal 103, 116

semicuneata 103, 116

tenuinervula 116

treubii 116

Hornstedtia 157, 158, 159, 161, 163

leonurus 160, 163

Horsfieldia 75, 133, 134, 139, 140, 141

glabra 140

irya 75, 133, 135, 136, 139, 140

macrocoma 133

macrocoma Var. canarioides 75

novoguineensis (non Warb.) 135, 140

reticulata 140

ridleyana 135

sabulosa 133

Gardens’ Bulletin, Singapore — XXV11 (1974)

spicata 135, 139, 140, 141

subglobosa 76, 140

sucosa 76

superba 76

palauensis 140, 141

palewensis (non Kanchira) Whitmore 135

parviflora 139

polyspherula 140

tomentosa 76

wallichii 76, 133, 135, 134

whitmorei 135, 139, 140, 141

Hoya diversifolia 29

Hydrangea 126

macrophylla 249

Hymenaea courbaril 261

Hypserpa cuspidata 24, 41, 65M, 82

Illigera appendiculata 81

trifoliata 81

Impatiens 126

Imperata 126

Indigofera hirsuta 261

tinctoria 261

Intsia bijuga 262

Ischaemum 126

Johannesteijsmannia 122

Kadsura cauliflora 68

scandens 68

Kaempferia 158, 159, 161, 164

Kalanchoe laciniata 249

pinnata 249

Kandelia 183, 184

Knema 75

communis 42

conferta 76

curtisii 76

furfuracea 76

glaucescens 76

globularia 76

hookeriana 76

intermedia 76

latericia 76

laurina 76

malayana 76

Koompassia malaccensis 262

Kunstleria ridleyi 262

Languas 160, 163

Lasianthus maingayi 44

Lauraceae

key to the genera 77—78

Leguminosae

key to subfamilies 253

tribes 253-254

Leptosphaeria suffulta 127

Leptosphaerulina trifolii 124

Leucaena glauca 262

leucocephala 262

Licaria splendens 250

Limacia velutina 83

Lindera lucida 79

malaccensis 79

Lithocarpus conocarpus 20, 35, 54, 65F

sundaicus 24, 36, 55

Litsea 38

accedens 80

Index

cordata 80

costalis 80

elliptica 20, 39, 65H

erectinervia 80

ferruginea 80

firma 80

garciae 80

grandis 39, 80

griffithii 80

lanceifolia 80

lanceolata 80

machilifolia 80

megacarpa 80

myristicaefolia 80

perakensis 80

petiolata 80

ridleyi 81

robusta 81

sebifera 80

singaporensis 80

Lycopersicum 126

Macaranga triloba 34

Macrophomina phaseolina 125

Magnolia 67

coco 67

maingayi 67

Magnoliaceae

key to the genera 67

Mallotus penangensis 35

Manghania strobilifera 263

Mangifera 126, 127

indica 27, 28

Maranthes corymbosa 250

Marasmius pulcher 123, 126

Marasmiellus inoderma 123, 126

scandens 126, 128

Matthaea sancta 77

Meiogyne 69

virgata 72

Melastoma 127

Meliola canangae 124

Melodorum elegans 72

fulgens 71

prisomaticum 74

Meloidogyne incognita 124, 126, 127

javanica 124

Menispermaceae

key to the genera 82

Mesembryanthemum 122

Mezaneuron sumatranum 262

Mezzettia leptopoda 72

Michelia 67, 127

alba 67

champaca 67

figo 67

Micropora 81

curtisii 81

?Microxyphium sp. 125

Miliusa

longipes 72

Millettia atropurpurea 262

eriantha 262

maingeyi 262

Mimosa invisa 262

pudica 262

sepiaria 263

Mitrella kentii 72

Monocarpia marginalis 73

Moringa pterygosperma 249

Morus 127

Mucuna bennetti 263

gigantea 263

Mycospharella sp. 129

Myristicaceae, key to genera 75

Myristica 75

cinnamomea 77

crassa 77

elliptica 77

fragrans 77

maingayi 77

maxima 77

Myrothecium gramineum 128

Nasturtium indicum 248

officinale 248

Negrospora sphaerica 126

Nelumbo nucifera 83

Neottopteris antiqua 151

australasica 147

curtisora 148

cymbifolia 151

elliptica 148

grevillei 153

humbertii 154

mauritiana 148

musifolia 150

nidus 147

ovata 154

pachyphylla 154

phyllitidis 152

rigida 148

salwinensis 148

simonsiana 152

simplex 154

squamulata 154

stenocarpa 148

stipitata 154

taeniosa 154

vulgaris 143, 147

Nephelium 127

Nephrolepis biserrata 46, 65M

Neptunia natans 263

oleracea 263

Nicolaia 156, 157, 159-162, 164

hemisphaerica 161

solaris 161, 162

Nigrospora sacchari 123

sphaerica 127

Northaphoebe coriacea 81

kingiana var. malvescens 20, 39, 40, 62, 65H

umbelliflora 81

Nymphaea capensis 83

lotus 83

nouchali 83

pubescens 83

stellata 83

Nymphaeaceae, key to genera 83

Oncosperma 4, 6

Orchidantha 164

Ormosia banca 263

273

274

macrodisca 263

sumatrana 263

Oryza 127

Oxymitra biglandulosa 72

Pachyrhizus erosus 263

Padebruggea maingayi 262

Palaquium obovatum var. obovatum 44

semaram 44, 65L

Pandanus odoratissimus 4

Parashorea spp. 100, 103, 105, 106, 110

macrophylla 105, 106, 116

malaanonan 105, 106, 116

parvifolia 105, 116

smythiesii 105, 106, 116

Parastemon urophyllus 250

Parinari asperulum 250

corymbosa 250

excelsa 250

oblongifolia 251

sumatranum 251

Parinarium costatum 251

griffithianum 250

Parkia speciosa 263

Passiflora 127

’ Paxillus squarrosus 15

Pellacalyx 183, 186, 190, 193, 194

axillaris 184, 187, 202A

saccardianus 184, 187, 208, 216

Peltophorum ferrugineum 263

pterocarpum 263

Pennisetum 127

Peronospora parasitica 124

Persea americana 81

Pestalotia psidii 127

Pestalotiopsis sp. 127

cruenta 127

Phaenthus nutans 73

ophthalmicus 73

Phaeomeria 156, 162, 163

Phaeosaccardinuta ? tenuis 124

Phaeoseptoria sp. 126

Phaeotrichoconis crotalariae 125

Phanera flammifera 256

griffithiana 256

semibifida 256

Phaseolus 127

angularis 264

atropurpureus 85, 91, 92,92A

aureus 264

lathyroides 85, 92, 92B

lunatus 264

vulgaris 264

Phlebopus 4

Phoebe cuneata 81

declinata 81

grandis 81

opaca 81

Phoma eupyrena 128

herbarum 124

Phomopsis sp. 123, 124

mangiferae 127

Phylloporus 14, 15

bellus 15

brunneolus 15

Gardens’ Bulletin, Singapore — X XVII (1974)

ochraceobrunneus 1S.

rhodoxanthus 15

squamosus 14

stenosporus 15

Phyllitis 149

Phyllosticta celosiae 124

Phytophthora nicotianae var. parasitica 125

palmivora 127

Piper 127

Pithecellobium affine 264

clypearia 40, 63, 651, 264

confertum 264

contortum 264

dulce 264

ellipticum 264

globosum 264

jiringa 264

lobatum 264

Pittosporum ferrugineum 249

Plagiostachys 160

Planchonella 4

Podoxyphium sp. 128

Poga oleosa 188

Pogostemon 127

Poinciana pulcherrima 257

Polyalthia 69

angustissima 73

cauliflora 73

var. beccarii

var. desmantha

clavigera 73

glauca 73

hookeriana 73

hypoleuca 73

lateriflora 73

macropoda 73

rumphii 73

sclerophylla 73

scortechinii 73

sumatrana 73

Polyosma conocarpa 249

fragrans 249

ridleyi 249

Pongamia pinnata 264

Popowia 69

fusca 74

pisocarpa 74

ramosissima 74

tomentosa 28, 60, 65K, 74

Porogramme rauenalae 125

Porphyrellus subflavidus 8

Practylenchus brachyurus 126

Prunus arborea 251

grisea var. tomentosa 251

polystachya 251

?Pseudocercosporella sp. 128

Pseudoepicoccum cocos 125

Psidium 127

Psophocarpus tetragonolobus 265

Psoralea corylifolia 265

Psychotria ovoidea 44, 65L

Pterocarpus indicus 265

Pygeum parviflorum 251

persimile 251

ee ee ee eee

Index

Pyramidanthe prismatica 74

Pyricularia grisea 125

Raphanus 127

sativus 248

Rhizophora 4, 183, 184, 186, 194, 195

mucronata 195

Rhizophoraceae

anatomy 187

leafblade 187-189, 217

petiole 190, 218

stem 190, 218

wood 190, 191, 219, 220

tribal characters 192

classification 197, 198, 201

by Hooker 197, 201

by Melchior 198, 201

by Schimper 197

flower and pollen 194-196

fruit 184

leaf tissues 189

morphology of tribes 198—199

pollen grains 195

seed 185

seedling 186

tribe Anisophylleeae 198—200

tribe Gynotrocheae 198—200

tribe Rhizophoreae 198—199

Rhizopus artocarpi 124

Rhodamnia cinerea 43, 65J

trinervia 43, 44

Ricinus 128

Rorippa indica 248

Rosa 128

chinensis 251

indica 251

Rourea acuminata 253

fulgens 252

minor 253

mimosoides 253

rugosa 253

Rubus angulosus 251

glomeratus 251

moluccanus 251

Russula 1

Salacca 4

Salacia korthalsiana 30

Samanea saman 265

Sanseviera 128

Saraca indica 265

thaipingensis 265

Sauropus 128

Saxifraga sarmentosa 249

stolonifera 249

Scaphochlamys 158, 159, 160, 164

Scolopendrium 144

Semecarpus sp. 28

Septobasidium lepidosaphis 125

Phaeogyroporus 4

portentosus 2, 4

tropicus 4

Serianthus dilmyi 265

grandiflora 265

Sesbania grandiflora 265

Shorea sp. 27, 33, 95, 100, 103, 105, 106, 107, 110

275

acuminatissima 118

acuta 116

agami 118

albida 116

almon 116

amplexicaulis 116

andulensis 116

angustifolia 116

argentifolia 116

atrinervosa 117

beccariana 116

belangeran 34

biawak 117

bracteolata 118

brunnescens 117

bullata 116

carapae 116

cordata 118

collaris 118

coriacea 116

crassa 117

cristata 116

curtisii 19, 20, 24, 30, 33, 50, 51, 52, 65B, 65C.

65D, 116

cuspidata 118

dasyphylla 116

dolichocarpa 118

domatiosa 117

elliptica 116

faguetiana 118

faguetioides 118

fallax 116

ferruginea 116

flava 117

flaviflora 116

flemmichii 116

foxworthyi 117

geniculata 117

gibbosa 118

glaucescens 117

havilandii 117

hopeifolia 118

iliasii 118

inaequilateralis 116

inappendiculata 117

induplicata 118

isoptera 117

kunstleri 116

ladiana 117

laevis 117

lamellata 118

leprosula 108, 116

leptoclados 116

longiflora 118

macrantha 116

macrobalanos 118

macrophylla 117

macroptera 117

maxwelliana 117

mecistopteryx 117

monticola 117

mujongensis 118

multiflora 118

myrionerva 117

276

obscura 117

obovoidea 118

ochracea 107, 118

ovalis 117

ovata 108, 117

pachyphylla 117

palembanica 117

pallidifolia 117

parvifolia 117

parvistipulata 108, 117

patoiensis 118

pauciflora 23, 41, 117

pilosa 117

pinanga 117

platycarpa 117

platyclados 117

praestans 117

pubistyla 117

quadrinervis 117

resina-nigra 118

resinosa 118

retusa 117

rubella 117

rubra 117

rugosa 117

scaberrima 117

scabrida 108, 117

scrobiculata 117

seminis 117

slooteni 117

smithiana 117

stenoptera 117

subcylindrica 118

superba 117

uliginosa 117

teysmanniana 108, 117

venulosa 117

virescens 118

xanthophylila 118

Sindora wallichii 265

Sonneratia 4

Spatholobus ferrugineus 265

maingayi 266

rideyi 266

Sphacelotheca hainanae 126

Sphaerostilbe repens 127

Stemphylum lycopersici 124, 125

Stenolobium 128

Strobilomyces 16

velutipes 16

Strongylodon macrobotrys 266

Strychnos sp. 40, 65K

Stylosanthes 128

humilis 85, 92B, 93

Talauma 67

Lanigeru 68

singaporensis 68

villosa 68

Tamarindus indica

Tephrosia noctiflora 266

Terminalia catappa 30

Tetraploa aristata B. & Br. 127

Thamnopteris australasia 147

antiqua 151

Gardens’ Bulletin, Singapore — X XVII (1974)

grevillei 153

nidus 147

var. musifolia 149

var. phyllitidis 152

orientalis 152

pachyphylla 154

phyllitidis 152

simonsiana 152

simplex 154

simplex sensu Pres] 153

squamulata 154

Theobroma 128

Tinomiscium petiolare 83

Tinospora crispa 83

Trichomerium sp. 127, 128

?crotonis 124

Trigonobalanus 5

Tripospermum sp. 124

gardneri 128

Upuna borneensis 99, 100, 102, 103, 105, 106, 110,

111, 112, 418

Uraria crinita 266

Urophyllum hirsutum 44, 65L

Uvaria 68

cordata 74

curtisii 74

grandiflora 74

hirsuta 74

leptopoda 74

lobbiana 74

macrophylla 74

pauci-ovulata 74

purpurea 74

rufa 74

Uvariella leptopoda 74

Vanilla 128

Vatica spp. 99, 100, 102, 103, 104, 110, 111, 112

badiifolia 118

borneensis 118

brunigi 118

coriacea 118

dulitensis 118

endertii 118

granulata 112, 118

globbosa 118

havilandii 112, 118

maingayi 118

mangachapoi 112, 118

micrantha 118

nitens 118

oblongifolia 118

odorata 118

papuana 118

rynchocarpa 118

sarawakensis 112, 118

umbonata 112, 118

venulosa 118

vinosa 118

Vigna 128

marima 266

retusa 266

uniquiculata 266

ssp. sesquipedalis 266

ssp. sinensis 266

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Index

277

Vitex pubescens 45, 65M magna 75

vestita 45 malayana 24, 29, 59, 75

Vitis gracilis 45, 65M ridleyi 71

Voandzeia 128 Zingiber 129, 157, 158, 159, 164

Willughbeia coriacea 29 clarkei 157

Xanthophyllum 128 Zingiberaceae

Xylopia 69 growth-habit 156

caudata 74 inflorescence-structure 157, 164

curtisii Zornia diphylla 266

ferruginea 75

Typographica! Corrections in the Index

page

267 alter (Albizzia) ittoralis to littoralis

267 alter (Anisophyllea) distiha to disticha

268 alter (Boletus) anripes to auripes

269 alter (Caesalpinia) Nuga to nuga

270 alter (Cyathea) Squamulata to squamulata

213 alter Ormosia banca to bancana

274 alter (Phylloporus) squamosus to squamosus

276 alter (Spatholobus) rideyi to ridleyi

276 alter (Talauma) Lanigeru to lanigeru

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GARDENS’ Bee

SINGAPORE

Vol. XXVII, Part I

I5th February, 1974

CONTENTS

Corner, E. J. H.: Boletus and Phylloporus in Malaysia : further notes and

descriptions - - - -

ANTHONY, MING: Contribution to the knowledge of Cecidia of Singapore -

KENG, Hsuan: Annotated list of seed plants of Singapore (II) - - -

Cuow, K. H.: Morphology and ecology of some introduced herbaceous legumes

CuHu, F. Fei-TaAN: Anatomical Features of the Dipterocarp Timbers of Sarawak

REVIEW: Palms of Maiaya (Hsuan Keng) - . - - - -

KUEH, TIONG-KHENG: New Plant Disease Records for Sarawak for 1972 - -

REVIEW: Animal Life and Nature in Singapore (Lee Chu San) - - -

J. SINCLAIR: The genus Horsfieldia (Myristicaceae) in and outside Malesia I:

H. sabulosa and H. whitmorei J, Sinclair spp. nov. - - - -

To be purchased at the Botanic Gardens, Singapore

Price: S $18

Published by Authority

Printed by the Singapore National Printers (Pte) Ltd

1974

PAGE

l- 16

17— 65

67— 83

85— 94

95-119

4Z1—122

123-129

131-132

133-141

MELOY OOYO_OYIE EPO DYE YEE LE

| i

KE FED

VEN EOL ERE IE VIO

fib iy 2

GARDENS’ BULLETIN

SINGAPORE

Vol. XXVII, Part I 15th February, 1974

Boletus and Phyiloporus in Malaysia:

further notes and descriptions

E. J. H. CORNER

Botany School, University of Cambridge

New taxa:— Boletus subgen. Boletus, B. hibiscus sp. nov. (Sarawak), B. rubroglutinosus

sp. nov. (Sarawak); subgen. Leccinum, B. peronatus sp. nov. (Sarawak), B. squarrosipes sp.

nov. (Malaya); subgen. Punctispora subgen. nov., B. punctisporus sp. nov. (Sarawak); subgen.

Tylopilus, B. longipes Mass. var. albus var. nov. (Sarawak), B. tristior sp. nov. (Sarawak);

subgen. Xerocomus, B. albipurpureus sp. nov. (Sarawak), B. setigerus sp. nov. (Java). —

Phylloporus squamosus sp. nov. (Sarawak), P. stenoporus sp. nov. (Java).

In the week of 21 August 1917 C. F. Baker and N. Patouillard had the luck

to encounter a run of boleti in the Gardens’ Jungle in Singapore. They described

16 new species but, as many of these had already been collected by H. N. Ridley

and described by Massee, merely five of their names have survived (Corner 1972).

During the week 20-27 August 1972 I had the luck to encounter a similar run of

boleti in Sarawak about the neighbourhood of Kuching, especially in Bako

National Park. I record 20 species of which six appear to be new. These and a

few others from Malaya and Java are here described. So it is that every region

of south-east Asia adds to our knowledge of boleti.

The weather in Malaysia had been exceptionally dry from May or June 1972

until the beginning of August; in places there had been water-rationing. Heavy

showers then induced this first crop of agarics which began around Kuching on

19 August with boleti and Cantharellus and passed on to Amanita, Russula and

Entoloma. In early September I had to leave on my homeward journey via Ceylon

where a similar dry period had extended until about 12 September. The monsoon

broke at this date when I was in Polunarawa. Unfortunately I could not delay,

but I wondered what fungi would come up in the extensive, if mainly secondary,

forests that are around this ancient town; the region is unexplored mycologically.

Many trees at Polunarawa had already begun to develop new leaves in anticipation,

as it were, of the rains which had ceased in the middle of March; the parched

scene was set for the mycologist, and it struck me as a good occasion for

gasteromycetes.

2 Gardens’ Bulletin, Singapore — XXVII (1974)

In Sarawak I was particularly interested in the fungi of Bako National Park

because it consists largely of the xerophilous kerangas forest on sandstone with

a few small valleys of typical lowland dipterocarp forest. Yet I could find no

marked difference between the fungous floras of these very different kinds of

forest. The species of Boletus and Amanita seemed to occur indiscriminately.

Many were widely distributed species of lowland West Malaysia and Singapore

which are without kerangas forest.

I draw attention to Boletus hibiscus as a massive ally of B. (Phaeogyroporus)

portentosus which has not been reported from Malaysia, to B. punctisporus as an

ally of the North American B. betula, to B. setigerus as yet another small species

with thick-walled cystidia, and to the two new species which I assign to subgen.

Leccinum for they relate with B. borneensis which I have described from Kinabalu

(Corner 1972).

BOLETUS Fr.

subgen. Austroboletus Corner

B. dictyotus (Boedijn) Corner

var. morchellipes Corner (1972) p. 85.

Pileus 7-17 cm wide, conical then convex to more or less plane, fawn ochraceous

felted, dry, becoming cracked; margin at first extensively appendiculate and in

unopened fruit-bodies concrescent with the upper ridges of the stem-reticulum,

but sometimes detaching from the pileus as a more or less broken ring. Stem

8-15 cm x 7-25 mm (excluding the reticulum), subcylindric, white, coarsely

alveolate-reticulate downwards, the meshes towards the base of the stem 5-15 mm

deep. Tubes 12—20 mm long, white then vinaceous pink, soft; pores 0.5-2 mm

wide, white then vinaceous pink. Flesh 20-35 mm thick in the centre of the pileus,

white, unchanging, but yellowish at the base of the stem with age. Spores 12—15.5

x 5—6u (spore-body), with small warts 0.5—1p, high, both ends of the spore smooth.

Sarawak, Bako National Park, scattered in the humus, rarely two together,

in kerangas and lowland dipterocarp forest, Corner P-147, 25 Aug. 1972.—Malaya,

Pahang, Fraser’s Hill, Corner s.n., 6 Sept. 1972.

This with its massive spongy stem is one of the more striking species of

Malaysia. It was common at Bako and the fruit-bodies were eaten by squirrels.

B. malaccensis (Pat. et Baker) Corner (1972) p. 85.

Pileus without marginal veil. Stem —9 cm high, attenuate upwards from the

fusiform base, in the lower third with shallow, very elongate meshes -25 x 4 mm

and 1—-1.5 mm deep, wholly minutely yellowish pruinose. Spores 10.5-13(C-14) x

4.7-5.5. Basidia 27-34 x 10-11; sterigmata 4, 6-8» long. Tube-trama be-

coming sloppy-gelatinous in alcohol-formalin.

Sarawak, Bako National Park, solitary in humus, Corner P—-158, 26 Aug. 1972.

This collection agreed closely with those from Singapore.

B. mucosus Corner (1972) p. 77.

Sarawak, Bako National Park, common in all inland parts whether in kerangas

forest or not.

Boletus and Phylloporus in Malaysia 3

The young fruit-bodies had the margin of the pileus extended into a flange,

as in B. longicollis, and disrupting to make a broken annulus on the reticulate stem.

B. rubiicolor Corner (1972) p. 86.

Pileus —-6 cm wide, minutely rufous madder or purple madder squamulose;

margin appendiculate -15 mm long, disrupting into fragments or collapsing on

to the stem. Stem 13-15 cm x 7-8 mm at the apex, 10-15 mm at the subclavate

base, white, finely reticulate at the apex, coarsely and rather shallowly costato-

reticulate downwards with elongate meshes 10-17 x 2-4 mm, 1-3.5 mm deep,

most pronounced about the middle of the stem. Tubes —20 mm long; pores

1-1.3 mm wide. Flesh —13 mm thick in the centre of the pileus, soft, sappy, easily

putrescent, unchanging.

Except for the much larger fruit-bodies with pronounced marginal flange

to the pileus and rather strongly lacunoso-reticulate stem, the collections agreed

with previous ones from Singapore. This appears to be one of the very early

species to fructify at the beginning of a fungous season.

Sarawak, Kuching, Semanggoh Forest, Corner P-91, 19 Aug. 1972; Bako

National Park s.n., 26 Aug. 1972, in kerangas-humus; solitary, scattered.

subgen. Boletellus Murr.

B. ananas Curtis

Sarawak, Bako National Park, in kerangas-humus, s.n. 26 Aug. 1972.

B. longicollis Ces.

Sarawak, Bako National Park, in kerangas-humus and that of lowland diptero-

carp forest, frequent, solitary, 26 Aug. 1972.

subgen. Boletus

Boletus hibiscus sp. nov.

Pileus 9-26 cm latus, convexus dein planus et concavus, laevis, siccus, laete cervino-

luteus dein (? de sporis depositis) cinnamomeo-ferrugineus; margine flavido, primo ut mem-

brana 3-4 mm lata tubulos excedenti. Stipes 7-12 cm x 16-45 mm, cylindricus, firmus dein

subcompressus intusque spongiosus, cortice dura, luteus dein e basi fusco-brunnescens, ex

integro minute pruinoso-furfuraceus, haud reticulatus; mycelio albo. Tubi 9-12 mm longi,

sinuato-ventricosi, pallide cervino-cinnamomei dein cinnamomeo-ferruginei; poris 0.8-1.5 mm

latis, angulatis, rugulis longitudinalibus subcompositis, marginem pilei versus saepe radiato-

elongatis, concoloribus. Caro in centro pilei 16-35 mm crassa, firma dein spongiosa, flavo-alba,

basim stipitis versus dura et intense ochraceo-lutea, sub superficie pilei flava, fractu vix

flavescens vel in stipite subbrunnescens, nec cyanescens nec rubescens. Odor subacidus.

Sporae 8-9.5 x 6.3-7y, in cumulo laete ferrugineae (nec olivaceae nec incarnatae), ovoideae,

laeves, inamyloideae, |-guttatae, tunica in KOH subincrassata, apiculo 0.2, longo.

Ad terram arenosam maritimam in Hibisceto tiliaceo, gregaria, saepe in circulis latis.

Sarawak, Bako National Park, Telok Pandan et Telok Bako, 28-29 Sept. 1972, leg. E. J. H.

Corner P-181 (typus, CGE).

Pileus bright brownish yellow, becoming cinnamon-ferruginous from the

spores; margin exceeding the tubes as a flange 3-4 mm wide. Stem yellow, then

fuscous brown from the base upwards, minutely pruinose, firm then somewhat

baggy with spongy interior and firm rind. Tubes sinuate, pale brownish cinnamon

then ferruginous; pores angular, becoming subcompound with internal longitudinal

ridges, concolorous with the tubes. Flesh thick, yellowish especially in the stem.

unchanging.

4 Gardens’ Bulletin, Singapore — X XVII (1974)

Spores bright ferruginous in the mass. Basidia 30-42 x 8.5—-10p, 4-spored.

Cheilocystidia 30-45 x 4-8, more or less narrowly ventricose, often with pro-

longed or bifurcate obtuse apex 2-3y wide, thin-walled, colourless, as a sterile

edge to the pores. Pleurocystidia not seen. Surface of pileus consisting of a dis-

rupted and flattened pile of hyphal ends —100, long, with submoniliform clamped

cells 20-40 x 7-15, not encrusted, the end-cells subclavate. Surface of the stem

covered by a more or less disrupted and (?) mainly sterile hymenium of immature

basidia and cystidia as the cheilocystidia. Tube-trama phylloporoid with long-

celled mucilaginous hyphae 4-1lp wide, clamped, becoming rather toughly gela-

tinous in alcohol-formalin. Hyphae clamped, inflating, with scattered oleaginous

hyphae in the stem and pileus; in the stem mainly longitudinal with the cells

70-400 x 5-35, cylindric or fusiform, thin-walled, with some uninflated inter-

weaving hyphae as laterals of the inflated longitudinal hyphae, but with narrower

and more compact hyphae in the firmer cortex; without chlamydospores at the

base of the stem.

This striking species is recognised from its large size, bright brownish yellow

colour, unchanging flesh, copious ferruginous spore-deposit, and habitat. With

clamped hyphae it comes near to B. portentosus which differs in the olivaceous

fruit-body and spores. The ferruginous spores place the species in Gyrodon which,

however, has decurrent tubes. Phaeogyroporus with olivaceous brown spores has

the sinuate-ventricose tubes. To both genera a boletoid tube-trama is ascribed;

that of B. hibiscus is phylloporoid. According to a collection that I have from

Brazil, Phaeogyroporus tropicus (Rick) Singer has yellow-brown (not olivaceous)

spores, phylloporoid trama, and adnexed or tapered-adnate tubes. Therefore, while

the distinction of these imperfectly characterised genera (including Phlebopus) is

so uncertain, I retain the species in Boletus near to B. portentosus and B. tropicalis.

I found about 150 fruit-bodies of B. hibiscus in two forested bays of Bako

National Park. They grew in more or less complete circles up to 5 m in diameter

in thickets of Hibiscus tiliaceus, Dillenia suffruticosa, Planchonella and Pandanus

odoratissimus, not directly on the sea-front but where this vegetation borders the

outlets of mangrove-streams. It was impossible to associate these circles with any

particular tree or, indeed, to trace the mycelium beyond the lowest level of the

humus where it mixed with the sand. The fungus did not grow with Rhizophora,

Bruguiera or Sonneratia or in the muddy ground of the palms Oncosperma and

Salacca.

B. hibiscus comes in the artificial key to Malaysian boleti (Corner 1972, p. 49),

next to B. chlamydosporus.

B. rubroglutinosus sp. nov.

Pileus 4.5-6 cm latus, sanguineo-ruber, glutinosus. Stipes 6-7 cm xX 9 mm, cylindricus,

concolor, rubro-pruinosus, ad basim mycelio ochraceo-aureo vestitus. Tubi 5 mm _ longi,

sinuato-adnati, flavi; poris minutis, sanguineo- vel aurantiaco-rubris, dein sordide aurantiaco-

incarnatis. Caro —9 mm crassa, flava, intense cyanescens ut tubi porique. Sporae 9.5-12 x

3.7-4.54,, in cumulo fusco-purpureae, leniter olivaceotinctae, laeves.

Ad terram in silva. Sarawak, Bako National Park, Corner P-145 (typus, CGE), P-145a.

25-26 Aug. 1972.

Pileus 4.5-6 cm wide, convex (immature), evenly blood-red, smeary-viscid

pelliculose. Stem 6-7 cm x 9 mm, cylindric, dry, blood-red, minutely red scurfy-

pruinose, not reticulate, base slightly tapered and thinly villous with the ochraceous

golden-yellow mycelium. Tubes 5 mm long, adnate then sinuate, yellow; pores

minute, blood-red or orange-red, then dull orange-pink. Flesh -9 mm thick in

Boletus and Phylloporus in Malaysia 5

the centre of the pileus, yellow, quickly and intensely cyanescent on bruising or

cutting as the tubes and pores. Smell none.

Spores in the mass fuscous purplish tinged olivaceous. Basidia 25-32 x 9-10p:

sterigmata 4. Cheilocystidia 20-40 x 8-14y, clavate to subventricose or subfusoid,

thin-walled, as a sterile edge to the pores. Pleurocystidia —75 x 14y, ventricose,

thin-walled, with a short appendage 2» wide, sparse. Surface of the pileus com-

posed of appressed, narrow, colourless mucilage-hyphae 2—3.54 wide, with cylin-

dric ends, neither as a pile nor as a palisade, without clamps. Surface of the

stem with a disrupted and mainly sterile hymenium of clavate sterile basidia,

subglobose cells 14-21 wide, and more or less ventricose cystidia, some with an

appendage -60 x 2-3y; stem-hyphae 5—l6y wide, without clamps, many secon-

darily septate. Tube-trama boletoid, swelling and firmly gelatinous in alcohol-

formalin, with a narrow medulla of narrow hyphae 3-5» wide and a cortex of

gradually divergent hyphae —94 wide with strongly mucilaginous walls.

This species is distinguished by the glutinous red pileus, the dry red fur-

furaceous stem, the minute red pores, the yellow cyanescent flesh, and the yellow

mycelium. With boletoid trama and viscid pileus it should go into Suillus of modern

authors but, among Malaysian boleti, it seems to me related with B. subreticulatus

and B. lubricus, neither of which have cyanescent flesh. It may be rare for I found

but two specimens. In the artificial key to Malaysian Boletus (Corner 1972, p. 41)

it should be entered in Group C after Heimiella mandarina.

B. rufo-aureus Mass.

Sarawak, Bako National Park, in kerangas-humus, Corner P—182, 28 Aug.

1972.—Malaya, Pahang, Fraser’s Hill, Corner s.n., 6 Sept. 1972.

Though large, with the pileus 8-12 cm wide and the stem 11 cm x 27 mm,

the specimens were immature; it is the wont of this species to mature late in

development. The few spores that I found were smaller than usual and 9.5-11 x

3.5—-4.2; cystidia, most basidia, and the hyphal ends on the pileus had not reached

their adult size and shape.

B. sinapicolor Corner (1972) p. 124.

Pileus -—8 cm wide. Stem poroid-reticulate at the apex, the reticulations

elongate and deeper downwards, —0.5 mm deep, but absent from the lower third

of the stem. Tubes 7-8 mm long, sinuate; pores 0.5 mm wide. Flesh hard and

firm in the stem.

Spores 10.5-12.7 x 44.5u. Basidia 30-37 x 9-10; sterigmata (2-3-) 4.

Pleurocystidia -65 x 9-14y, ventricose with narrow apex, thin-walled, inconspi-

cuous, sparse. Cheilocystidia similar, sparse. Surface of pileus with cells 40-80 x

7-14. in the hyphae of the pile, the end-cells clavate or ventricoso-attenuate.

Stem-surface with a disrupted sterile hymenium, along the ridges with excrescent

2—3-septate hyphae -170 x 7-9u. Stem-hyphae with the cells -170 x 7-16n.

Tube-trama subboletoid with slight medulla of narrow hyphae, the cortical hyphae

5-13 wide, becoming somewhat swollen but firm in alcohol-formalin.

Malaya, Pahang, Fraser’s Hill, Corner P—201, 6 Sept. 1972, in humus in

Trigonobalanus-forest.

This collection, from the same valley where I first found the species forty

years ago, has enabled me to add some details to the description. The spores were

slightly longer than I had previously found.

6 Gardens’ Bulletin, Singapore — XXVII (1974)

Recently B. auripes Pk. has been recorded from Japan by Hongo (1970)

whose description of the species, as it occurs in Japan, reads extremely like that

of B. sinapicolor. He describes the yellow tomentose base of the stem but the

yellow colour of the fruit-body does not seem to be the uniform and striking

mustard-yellow of B. sinapicolor. In contrast, for North American specimens of

B. auripes, the mycelium at the base of the stem is given as white (Snell and

Dick 1970) and it is not figured or described as yeliow by Coker and Beers (1943).

But these authors describe, and Hongo finds in the Japanese material, the immature

pores as stuffed and covered, as it were, by a membrane. This is the nature of the

young pores in B. phaeocephalus to which I considered B. sinapicolor to be allied

(Corner 1972, p. 125), but I have not seen this membrane in B. sinapicolor, though

I have had young specimens. Here is evidently a close alliance in which the

Japanese B. auripes approaches B. sinapicolor.

B. thibetanus Pat.

Pileus —8 cm wide. Stem —10 cm high. Pores 0.5 mm wide, golden citrine.

Spores 10.5—13¢-14) x 4.5-5.5u, olive brown turning dark fuscous brown in

potash.

Sarawak, Bako National Park, solitary in humus, Corner P—157, 26 Aug. 1972.

This characteristic species was recognised at once in the forest but microscopic

examination of the dried material some months later revealed a point which may

be peculiar or, hitherto, overlooked. The spores of the Sarawak collection, which

were slightly larger than the Malayan, turned rather dark fuscous brown in potash.

Dried spores of the Malayan material (now thirty years old) are very pale in potash.

B. xylophilus Petch

Malaya, Kuala Lumpur, Lake Garden, on earth in the centre of an isolated

clump of the palm Oncosperma, Corner s.n. 3 Sept. 1972.

subgen. Leccinum S. F. Gray

B. peronatus sp. nov. — Figure lc.

Pileus 6-9 cm, siccus, obscure brunneus villoso-subtomentosus, dein ochraceo-brunneus.

Stipes 7-8.5 cm xX 9-12 mm, subcylindricus v. subfusiformis, basim versus attenuatus, albidus

dein subochraceus, annulis floccosis in squamulis 1-2 mm projicientibus fuscescentibus irre-

gulariter peronatus, haud reticulatus; mycelio albo, Tubi 6-9 mm longi, adnexi, subliberi,

cremeo-albi dein olivaceo-virides; poris minutis, tubis concoloribus, aetate brunnescentibus.

Caro 10-16 mm crassa, albida dein subflavida, aetate vel fractu tarde ochraceo-brunnescens,

haud cyanescens. Odor subacidus. Sporae 8.5-11 (12) x 3.7-4.7, in cumulo olivaceae, laeves.

Ad humum in silva, solitarius v. caespitosus. Sarawak, Bako National Park, Corner

P-142 (typus, CGE) P-144, 25 Aug. 1972.

Pileus 6-9 cm wide, convex then plane, dry, at first dark fawn brown and

villoso-subtomentose, then brownish ochraceous with fuscous brown centre, paler

to the margin, tomentose, becoming minutely cracked, especially in the centre;

without veil. Stem 7-8.5 cm x 9-12 mm, subcylindric or subfusiform —15 mm

wide in the middle, usually attenuate at the base, whitish then pallid ochraceous

buff, irregularly peronate with more or Jess interrupted, divergent, floccose bands

projecting 1-2 mm in the middle of the stem and often appearing as connate squa-

mules, the bands becoming fuscous brownish, stem-apex smooth, not reticulate;

mycelium white, fibrillose. Tubes 6-9 mm long, adnexed, nearly free, subventricose,

pale cream-white then pale greenish olivaceous; pores minute, concolorous with

the tubes then brownish with age. Flesh 10-16 mm thick in the centre of the pileus,

Boletus and Phylloporus in Malaysia |

white then pale yellowish white, becoming pale brownish subochraceous in the

stem and pale brownish or pinkish brown with age or on exposure in the pileus,

not cyanescent or truly rufescent. Smell slightly sour.

Spores olive in the mass. Basidia 26-30 x 9-10; sterigmata 4, 4—Sp long.

Cystidia not seen; pore-edges more or less gelatinised without cellular structure.

Surface of the pileus composed of more or less divergent (—70u) or appressed,

laxly interwoven, uninflated hyphae 3-6y wide with slightly thickened walls, not

encrusted, not as a pile. Surface of the stem sterile except the stem-apex, composed

of narrow longitudinal hyphae 3—6y wide divergent in massive array to form the

peronate scales or annuli; hyphal ends in the scales uninflated or as small clavate

end-cells -40 x 12, as vesicular cells —16y wide, or subfusiform but not set in

a regular layer; internal hyphae of the stem —14, wide, longitudinal, without

clamps, often secondarily septate, at the base of the stem 3-8 wide with slightly

thickened walls, very compact, without chlamydospores. Tube-trama phylloporoid,

composed of longitudinal hyphae 4-8» wide, gradually divergent, rather firmly

gelatinous and somewhat swollen in alcohol-formalin.

This species is distinguished by the peronate scales of the stem. In the collec-

tion P—144 it seemed that they are formed by the cracking of the stem-surface,

but in P—142 they were firm as massive outgrowths of hyphae though evidently

spaced peronately by the elongation of the stem. In this respect the species resem-

bles B. borneensis Corner (1972, p. 106), which differs in the richer colour,

cyanescent flesh, longer spores and boletoid tube-trama. Possibly B. peronatus is

related with B. graveolens Corner. On stem-character the species belongs in subgen.

Leccinum. It will go in the artificial key to Malasian boleti in Group G (Corner

1972, p. 50) after B. sylvestris. Compare the following species with yet shorter

spores.

B. squarrosipes sp. nov. — Figure ld.

Pileus 7 cm, viscidus, pallide cervino-brunneus. Stipes 6 cm x 12 mm, siccus, concolor,

squamulis brunneolis furfuraceo-squarrosus; mycelio albo. Tubi 6 mm longi, sinuati, pallide

flavo-virides; poris 0.5—0.8 mm, tubis concoloribus. Caro alba immutabilis. Sporae 6—7.5 (-8)

x 5—6yu, in cumulo olivaceo-brunneae, laeves.

Ad humum sub arboribus. Malaya, Selangor, Kepong, Corner P-191 (CGE) 1 Sept. 1972.

Pileus 7 cm wide, plane, smeary-viscid, pale fawn brown. Stem 6 cm x 12 mm,

rather stout, fibrous, dry, pale fawn brown, in the upper two-thirds coarsely

squarroso-furfuraceous with pale brown particles or squamules; mycelium white.

Tubes —6 mm long, sinuate, pale greenish yellow: pores 0.5—0.8 mm, pale greenish

yellow. Flesh white, yellowish above the tubes, in the stem brownish, unchanging.

Spores olive brown in the mass, 1-guttate, smooth. Basidia 25-30 x 9.5-11p,

4-spored. Cystidia -46 x 7-lly, fusiform subventricose with the apex 3—4y wide,

thin-walled, sparse on the pore-edges and on the tube-surfaces. Surface of the

pileus composed of appressed hyphae 3-8» wide with mucilaginous walls and

brown sap, with scattered larger hyphal ends 9-18, wide (as if relics of a pile of

hyphal ends). Surface of the stem with the squarrose particles consisting of the

narrow superficial hyphae of the stem divergent in tufts and bearing a dense

patch of sterile hymenium composed of clavate cells and subventricose cystidia

-S5 x Ip, rarely with fertile basidia, and with occasional larger and shortly

septate hyphal ends with the cells 20-45 x 9-13, (as the larger hyphal ends on

the pileus). Hyphae of the stem -—18, wide, longitudinal, without clamps. Tube-

trama subboletoid, rather soft and sloppy in alcohol-formalin, with longitudinal

hyphae —9y» wide.

8 Gardens’ Bulletin, Singapore — XXVII (1974)

This fungus, like the preceding, recalls B. borneensis. Compare, however,

B. lubricus with viscid umber pileus, merely pruinose stem, and larger spores.

B. squarrosipes will go in Group C of the artificial key to Malaysian boleti (Corner

1972, p. 42) next to B. lubricus.

subgen. Punctispora subgen. nov.

Statu ut in subgen. Boletello. Sporae in cumulo olivaceo-brunneae, elongato-ellipsoideae,

verrucis v. echinulis in exosporio hyalino praeditae. Tubi adnexi ventricosi v. subliberi,

olivacei v. flavidi dein olivacei; trama plus minus phylloporoidea. Stipes reticulatus v.

sublacunoso-reticulatus. Hyphae sine fibulis. Typus, B. punctisporus sp. nov.

This subgenus may be described as Boletellus with internally verrucose, not

striate, spores. It is based on the following species B. punctisporus from Sarawak,

which turns out to be an unexpected ally of the North American Boletus betula

Schw. The fruit-bodies of B. punctisporus combine the characters of several sub-

genera or genera. They have the long, red, reticulate stems of Heimiella, the

marginal flange or veil of the pileus so conspicuous in subgen. Austroboletus and

Boletellus, and the yellow to olivaceous tubes with olive-brown spores of Heimiella,

Boletellus and Boletus s. str.

Microscopically the spores have distinctive markings, caused by internal

papillae in the exospore, such as have been described for B. betula by Singer

(1945), Snell and Dick (1970), and Grand and Moore (1971). The same markings

occur in subgen. Austroboletus but its spores are purple-brown or vinaceous and

its tubes, which lack the yellow and olive colours, have the mucilaginous boletoid

trama.

For B. betula, now usually referred to Boletellus, there is already the special

section Allospori Singer. The discovery of B. punctisporus shows that the two

species are part of an alliance equivalent to Boletellus (striate spores) or Austro-

boletus, within which sect. Allospori is distinguished by the viscid pileus without

marginal veil. Two other species may belong. Boletellus purpurascens Heinem.

of the Congo and B. schichianus (Teng et Ling) Teng of China have verrucose

spores. The illustration of the spore of Porphyrellus subflavidus (Murr.) Singer,

given by Grand and Moore (1971), is like that of B. betula but the end of the

spore is smooth as in subgen. Austroboletus, and the tubes and spores of this

species evidently lack the olive colour; hence I retained it in subgen. Austroboletus

where it fits closely (Corner 1972).

B. punctisporus sp. nov. — Figure 1b.

Pileus —6 cm latus, convexus, siccus, villosus, fusco-brunneus, dein in squamulis cuti-

fractus: margine primo ut membrana floccosa -10 mm lata tubos excedenti, dein e pileo

exscisso et ad stipitem annuliformi collapso. Stipes -11.5 cm x 12 mm, ad basim 7 mm

latum attenuatus v. basi subdiscoideo, plerumque elongato-reticulatus, furfuraceus, purpureo-

ruber, basim albovillosum versum laevis brunneus. Tubi -17 mm longi, adnexi, ventricosi,

olivacei; poris 0.5-0.8 mm, olivaceis. Caro 9-10 mm crassa, flavidula, sub superficie pilei

stipitisque rubra, supra tubos subcyanescens. Sporae 14-19 x 7-8u, in cumulo olivaceo-

brunneae, membrana hyalina punctata inclusae, ad apicem porosae, endosporio 0.74 crasso

colorato, nec alatae nec striatae, apiculo 0.54 longo.

Sarawak, Bako National Park, leg. E. J. H. Corner P-154, 25 Aug. 1972, P—154A (typus

CGE) 28 Aug. 1972.

Pileus -6 cm wide, convex, dry, villous, snuff-brown or fuscous brown, then

cracked into small squamules; margin exceeding the tubes as a floccoso-membranous

flange -10 mm wide, splitting from the pileus and collapsing as a ring on the stem.

Stem —11.5 cm x 12 mm, attenuate to the base 7 mm wide or the base slightly

Boletus and Phylloporus in Malaysia 9

dilated and subdiscoid, rather dull purplish red, with a concolorous reticulum

of shallow and narrow, elongate meshes, rough with scurfy purple-red flecks

scattered on the reticulum, smooth and brownish near the white villous base:

mycelium white. Tubes -17 mm long, adnexed, ventricose, dull olive: pores

0.5—0.8 mm wide, dull olive. Flesh 9-10 mm thick in the centre of the pileus, pale

yellow, over the tubes deeper yellow and slightly cyanescent on exposure, dull red

below the surface of the stem and pileus, not otherwise cyanescent or rufescent.

Spores olive-brown in the mass, ellipsoid, obtuse, the hyaline exospore sur-

rounding the brown ovoid endospore (11-15 x 6-7) and pierced by fine processes

appearing as hyaline dots on the surface of the exospore, the endospore-wall 0.71

thick, 1—2-guttate, apiculus 0.5. Basidia 14-16u wide, pyriform clavate; sterig-

mata 4. Cystidia ? lanceolate-ventricose, as the caulocystidia, but sparse. Surface

of the pileus with a thick pile —200p high, eventually disrupted into squamules,

composed of more or less moniliform hyphae with cells 25-75 x 8-19y, the

end-cells subclavate or subventricose. Stem with a more or less disrupted hymenium

of fertile and, mostly, sterile basidia, with scattered projecting, lanceolate or sub-

ventricose, thin-walled cystidia -60 x 7-10». Hyphae of the stem -15y wide,

secondarily septate, without clamps. Tube-trama more or less phylloporoid.

In the forest this species may be mistaken for Heimiella retispora or H.

subretispora, which lack the marginal flange or veil of the pileus. In the collection

P-145A the base of the stem was subdiscoid as in various species of Boletellus.

subgen. Tylopilus Karst.

B. ballouti Pk. var. fuscatus Corner (1972) p. 194.

Sarawak, Bako National Park, Corner P—173, 27 Aug. 1972.

B. coccineinanus Corner (1972) p. 152.

Java, Sukabumi district, Tjiletuh, Corner J-21, 29 April 1972, scattered

singly along an earth-bank at the edge of the forest, but not infrequent.

These specimens differed from the Malaysian merely in the red-brown, rather

than crimson, pileus, the wider stem (1.5-2 mm), the slightly redder pores (crimson

when young, fading on expansion), and the yellow (not orange-yellow) mycelium.

The spores under the microscope were pale ochraceous without olive tint. The

tubes were white, then pink, without a yellow or olive tint.

B. longipes Mass.; Corner (1972) p. 165.

Sarawak, Bako National Park, Corner P—143, P—143A, P-143B, P—143C,

25-28 Aug. 1972.

This was a common species, either solitary or occasionally in small troops,

both in the dipterocarp forest and in the kerangas-forest of the plateau. The spores

had in all the collections the characteristic blood-red colour in the mass, and the

blackening with potash, but there were differences in size; in P—143B the spores

were typical, 11.5-16 x 4.5-5.3u; in P-143A and P-143C they were 12-14 x

4.5-5.7; in P-143 they were 10-13 x 4.5—-5.2u. I did not find var. /atisporus.

The reticulations of the stem become more pronounced in older specimens.

On most the longer ventricose caulocystidia become slightly but distinctly thick-

walled at the base.

10 Gardens’ Bulletin, Singapore — X XVII (1974)

var. alba var. nov.

Receptacula alba; tubis porisque maturis pallide olivaceis, dein subroseis. Sporae

12.5-16 x 4.2-4. Sp, in cumulo purpureo-vinaceae, in KOH nigrescentes.

Sarawak, Bako National Park, Corner P—184 (CGE) 28 Aug. 1972, ad humum in silva

kerangas.

Pileus 6-9.5 cm. Stem 4-6.5 cm x 7-10 mm above, 12-16 mm at the base,

wholly shallowly and irregularly reticulate, the meshes elongate in the lower part

of the stem. Tubes 7-12 mm long; pores 1—1.5 mm wide.

About 30 fruit-bodies were found scattered in one patch of the kerangas

forest. They differed from typical B. longipes only in being pure white, except

for the maturing tubes and pores. The tube-trama was boletoid with narrow,

compact, medullary hyphae 2.5-5 wide, and more mucilaginous cortical hyphae

4-8. wide. The trama remained firmly gelatinous in alcohol-formalin.

B. nanus Mass.; Corner (1972) p. 150.

Sarawak, Bako National Park, scattered in small groups in kerangas humus,

26 Aug. 1972.

B. tristior sp. nov.

Pileus 5-8 cm latus, convexus dein planus v. concavus, siccus, fuligineo-umbrinus dein

sordide brunneo-umbrinus, minute pruinoso-velutinatus et cutifractus. Stipes 3-5 cm x

10-15 mm, subventricosus, subconcolor, ex integro minute fuscobrunneo-pruinosus, ad basim

mycelio albo villosus, haud reticulatus. Tubi -11 mm longi, adnexi, ventricosi, fere liberi,

pallide griseo-rosei; poris 0.5-0.8 mm latis, pallide roseis. Caro 8-10 mm crassa, alba firma

immutabilis. Odor subnullus. Sporae 9.5-12.5 (-16) x 3.5-4.2u, in cumulo pallide roseae,

laeves.

Mh humum, solitarius, sparsus. Sarawak, Bako National Park, Corner P-156, 26 Aug.

Basidia 28-33 x 9-10u; sterigmata 4. Cystidia 40-85 x 6-18u, more or less

ventricose, obtuse, not distinctly appendaged, thin-walled, multiguttulate, not red-

brown in potash, frequent on the tube-surface and pore-edges. Surface of the

pileus with a very short and compact pile 45-65, high, composed of 1—3-septate

hyphae 4-12, wide, the cells 7-33 long, with brown walls, cylindric or submoni-

liform. Stem-surface with an interrupted and more or less sterile hymenium of

clavate cells 16-28 x 4.5-12u and subventricose cystidia -60 x 7—l0u. Hyphae

of the stem —12y wide, secondarily septate, without clamps; at the surface of the

stem 2.5-5 wide, very compact, with slightly thickened walls. Tube-trama sub-

boletoid, with a medulla of narrow hyphae, the larger hyphae —10p wide and

scarcely divergent, little swollen and rather firm in alcohol-formalin.

This species strongly recalls B. albo-ater Schw. but the tubes are strongly

ventricose and the tissue is not rufescent-nigrescent. It comes near to B. funerarius

Mass. which has a rich brown pileus and stem, shorter spores, and a strong smell:

both have the firm flesh, particularly at the surface of the stem. In the key to

Tylopilus for Malaysian boleti (Corner 1972, p. 147) the species comes after

B. cervicolor.

B. viridis (Heinem. et Gooss.) Corner (1972) p. 197.

Pileus with small, pale olivaceous grey squamules under the gluten, crowded

over the centre. Stem 3-4 cm x 5—7 mm above, 5 mm at the base, cylindric or

attenuate downwards, at first white and pale fuscous olivaceous pruinose, then

viscid; mycelium white. Tubes 6-8 mm long, sinuate-adnate, pale pinkish ochra-

Boletus and Phylloporus in Malaysia 1]

ceous; pores pale pinkish ochraceous, Spores 11.5-15 (-17) 44.7, —5.Sp wide in

P—153B, in the mass fuscous ferruginous or tinted chocholate, golden brown under

the microscope. Cystidia not seen. Trama boletoid.

Sarawak, Bako National Park, solitary in humus, Corner P—153, 26 Aug.

1972, and P—153A, 28 Aug. 1972. — Malaya, Pahang, Fraser’s Hill, Corner

P-153B, 6 Sept. 1972.

There are slight differences from previous Malayan collections, such as the

squamules under the viscid layer of the pileus, the stem at first pruinose then

viscid (as in B. thibetanus), and the colour of the tubes and pores. The species

has some similarity with B. nanus Mass.

subgen. Xerocomus (Quél.) Konrad

B. albipurpureus sp. nov.

Pileus 7 cm latus, convexus (vix maturus), siccus, minute villoso-subtomentosus, pur-

pureo-brunneus. Stipes -11 cm x 15 mm, subcylindricus, in parte superiori longitudinaliter

rugulosus, haud reticulatus, ex integro minute furfuraceo-pruinosus, concolor vel basim

versus fuscobrunneus; mycelio albo. Tubi —10 mm longi, sinuato-adnati, pallide olivacei:

poris 0.5-1 mm latis, pallide flavido-olivaceis. Caro alba, sub superficie pilei purpureo-

brunnea, immutabilis. Sporae 14-20 x 5.5-6y, in cumulo olivaceo-brunneae, boletoideae,

laeves, (an laevissime striatae ?). f

Ad humum in silva, solitarius. Sarawak, Bako National Park, Corner P-155, 26 Aug.

1972 (CGE).

Basidia 32-43 x 11-12.5y; sterigmata 4. Pleurocystidia 67-105 x 16-25y,

ventricose with a rather short and obtuse or subcapitate appendage -25 x 7-10p,

thin-walled, copious. Surface of the pileus with a more or less interrupted pile

of cylindric or submoniliform hyphal ends —270, long, the cells 20-90 x 5-Il1p,

with slightly thickened and thinly encrusted walls. Surface of the stem with a

disrupted hymenium of fertile and sterile basidia, scattered ventricose cystidia

often larger than the pleurocystidia with the apex —18 wide, and numerous filiform

hyphal ends -80 x 3—5y with the tips 2 wide. Hyphae of the stem —l6p wide,

often secondarily septate, without clamps. Tube-trama subboletoid with a narrow

medulla of narrow hyphae, the wider hyphae (-12u wide) scarcely divergent, little

swollen and rather firm in alcohol-formalin.

This species comes in the close affinity of B. ferruginospora Corner, B.

raphanolens Corner and B. variisporus Corner. It differs in the larger fruit-body

with purple-brown pileus and stem, the rugulose stem, and the stout pleurocystidia.

The spores agree best with those of B. raphanolens which has yellowish flesh and

mycelium and rugulose-reticulate pileus. It keys out in Group G (Corner 1972,

p. 49) with B. variisporus.

B. destitutus Corner (1972) p. 227.

The following collection of a single specimen | refer to this species for

convenience. It keys out to B. destitutus but the wider spores and richer colouring

of the pileus, though not the small fruit-body, suggest B. satisfactus. In the study

of these small species it is difficult to know, as yet, what are the more critical

points, whether the size of the fruit-body, the length or width of the spores, or

the cyanescence of the flesh are the more reliable characters than, say, colour or

attachment of the tubes. It may be that this collection is a new species with small,

richly coloured fruit-body and short wide spores.

Gardens’ Bulletin, Singapore-— X XVII (1974)

Malaya, Penang Hill, 300 m alt., Corner P-68, 29 July 1972, solitary on a

bank in the forest. — Pileus 22 mm wide, convex, villous to minutely scurfy

velutinate, ferruginous fawn or fulvous cinnamon. Stem 12 x 3.5 mm, attenuate

downwards, pale brownish, apex slightly yellowish, minutely pruinose; mycelium

white. Tubes sinuato-adnate, pale dingy olivaceous buff; pores small concolorous.

Flesh 3 mm thick in the centre of the pileus, pale yellowish white; no part

cyanescent.

Spores 6.7-8 (-9) x 5.5-6.2 (—6.7)u, fuscous olivaceous under the micro-

scope, ovoid, smooth, 1-guttate. Basidia c. 30 x 10-11, 4-spored. Cheilocystidia

as sterile basidia. Pleurocystidia ? (not seen). Surface of the pileus with a pile

of submoniliform hyphal ends —250p long, some —400p long, with the cells 18-60

x 8-18, the end-cells obtuse and subclavate. Surface of the stem with a disrupted

sterile hymenium of clavate cells 20-34 x 7—-12n, with scattered narrow excrescent

hyphal ends 3—5u wide. Hyphae —23, wide in the stem, without clamps. Trama

phylloporoid, with hyphae 3-14, wide.

B. nugatorius Corner (1972) p. 224.

I refer here with doubt the following Javanese collection. It has the spores

and wide superficial hyphae of the pileus of B. junghuhnii but lacks the red colour

in the stem and the cyanescence of the pores. In shape and colour of the fruit-

body, but neither in its size or viscid pileus, it recalls B. thibetanus.

Java, Tjibodas, Corner J—6, 11 April 1972, solitary on the ground in the

forest. Pileus 3 cm wide, convexo-plane, dry, finely tomentoso-villous, purplish

brown, then minutely cracked with the yellow flesh showing. Stem 4 cm x 4 mm,

attenuate upwards, pale brownish yellowish, minutely yellowish scurfy; mycelium

white. Tubes adnexed, olive-yellow; pores 0.3-0.5 mm wide, angular, unequal,

yellow to olive. Flesh pale yellow, no part cyanescent.

Spores 10.5-12 x 4.5—5y, pale ochraceous under the microscope. Cystidia

ventricose, not appendaged, 11—20n wide, thin-walled, scattered in the tubes.

Surface of the pileus with a pile —180» high, composed of moniliform hyphal ends

with cells 20-50 x 12-30, the end-cells subclavate and obtuse. Surface of the

stem with a disrupted sterile hymenium, no ventricose cystidia seen. Tube-trama

phylloporoid.

B. setigerus sp. nov. — Figure la.

Pileus 15 mm latus, siccus, tomentosus, livido-ochraceus. Stipes 15 x 2 mm, fibrillosus,

haud reticulatus, pallide ochraceous: mycelio albo. Tubi 3 mm longi, sinuati, albi dein

flavidi; poris 0.5 mm latis, flavidis. Caro 4 mm crassa, alba immutabilis. Sporae 6.88.3 x

4.5-5,,, ochraceo-incarnatae s.m., laeves. Pleurocystidia -110 x 11-25y, ventricosa, obtusa,

crassitunicata. Superficies pilei ex hyphis angustis appressis instructa.

Ad ripam juxta silvam, solitarius. Java, Sukabumi distr., Tjiletuh, Corner J-29 (CGE),

30 April 1972.

Pileus convex, dry, opaque, thinly subtomentose, pale livid ochraceous. Stem

slightly thickened to the base 3.5 mm wide, pallid ochraceous buff, then pale

fuscous from the base upwards. Flesh slightly yellowish in the stem.

Spores pale pinkish subochraceous under the microscope, ellipsoid obtuse,

even subreniform, smooth, 1—2-guttate, slightly darker in Melzer’s iodine. Basidia

33-42 x 9.5-11u, 4-spored. Cheilocystidia 40-65 x 7-12p, fusiform to subven-

tricose with obtuse, not prolonged, apex 3-6y wide, the walls 0.5-1.5 thick in

the free part of the cystidium but not at the apex. Pleurocystidia -110 x 11-25pn,

ventricose with obtuse, not prolonged, apex 6-9 wide, varying subfusiform but

Boletus and Phylloporus in Malaysia

Se ae sila iia.

Lat “agen

y+ im is

b i

Ae : l

| “2 h

Litt NM v

iene

headed CN me me aes ome aes ween,

————

ot a

mm,

Ty, i lg

Bb he

a pe ee

Pa ” annuus

= Pwo

)

3 Benet ae sae.

‘a « ‘ a ee oe

sors ah aie

eae fa

err. ne a

G ~

Figure 1. a, Boletus setigerus, with pleurocystidia (very thick-walled) and cheilocystidia. —

b, B. punctisporus, in section and with the annulus in surface-view on the reticulate stem. —

¢, B. peronatus. — d, B. squarrosipes. Fruit-bodies x 1, those of B. squarrosipes Xx 4; spores

x 1000, the coloured wall in black; cystidia x 500.

_—_—__)

SSS ee

14 Gardens’ Bulletin, Singapore — XXVIII (1974)

always obtuse, the wall very strongly thickened, the lumen narrow, projecting, the

subhymenial base 5-7 wide and thin-walled but not dilated, with transitions to

the cheilocystidia near the pore-edges. Surface of the pileus composed of appressed,

interwoven, uninflated hyphae 3-6u wide, without a pile of hyphal ends. Tube-

trama with rather toughly mucilaginous longitudinal hyphae -12p» wide. Hyphae

without clamps.

This small species of pallid colour seems near to B. aculifer Corner and B.

hastulifer Corner, and to the much larger B. olivaceiluteus Corner (with acute

cystidia), but all these have a conspicuous pile of hyphal ends on the pileus. The

colour of the spores suggest that B. setigerus may belong in subgen. Tylopilus.

With these doubts, the species comes at the end of the key to Group I of Malaysian

boleti (Corner 1972, p. 52).

GYROPORUS Quél.

G. castaneus (Fr.) Quél.

Java, Sukabumi district, Lengkong, Corner s.n. 29 April 1972, solitary in the

forest.

HEIMIELLA Boedijn

H. retispora (Pat. et Baker) Boedijn

Pileus rose-red, then paler and brownish red on expansion. Flesh lurid yellow,

orange-yellow in the stem, not cyanescent.

In humus of dipterocarp forest, gregarious in compact troops. Sarawak, Bako

National Park, Corner P—160, 26 Aug. 1972, and P—160A, 28 Aug. 1972.

These specimens were typical except for the lack of cyanescence. I think that

H. japonica is at most a variety. One has to be careful now not to mistake this in

the forest for Boletus punctisporus with marginal flange to the pileus and pale

yellow flesh.

PHYLLOPORUS Quél.

P. squamosus sp. nov.

Pileus 6-8 cm, convexo-planus, siccus, tomentosus, purpureo-brunneus dein fusco-

cervinus et squamulis fibrilloso-fasciculatus. Stipes 5-6 cm xX 7-9 mm, subcylindricus,

cervino-brunneus, ex integro brunneo-pruinosus; mycelio albo. Lamellae alte decurrentes,

confertae, alte et late poroideo-anastomosae, albidae dein pallide brunneae. Caro 6-8 mm

crassa, albida dein flavidula, in stipite brunneola, immutabilis. Sporae 8.3-10.3 x 4.5-5.3y,

in cumulo ochraceo-brunneae, laeves. Cystidia 50-128 x 14-20u, copiosa, tenuiter tunicata.

Hyphae sine fibulis.

ae humum sub Quercu. Sarawak, Bako National Park, Corner P-141 (CGE), 25 Aug.

Pileus 6-8 cm wide, convex then plane, not umbonate, dry, brown tomentose

tinged purple, then fuscous fawn and splitting into fibrilloso-fasciculate squamules

larger in the centre and smaller towards the subtomentose margin. Stem 5-6 cm

x 7-9 mm, subcylindric or slightly tapered downwards, or flattened —11 mm wide,

fawn brown, wholly minutely brown pruinoso-subfurfuraceous, weathering smooth,

base thinly white villous. Gills deeply decurrent, rather crowded, 27-30 primaries

Boletus and Phylloporus in Malaysia 15

6-8 mm wide, 3-4 ranks, not dichotomous, becoming deeply and widely poroid

from the stem to the outer part of the limb with transverse and oblique ridges

forming meshes 2-4 mm wide radially, the reticulations not reaching the gill-edges,

thus persistently lamellate with deeply poroid-reticulate interstices, whitish then

rather pale fawn brown. Flesh 6-8 mm thick in the centre of the pileus, spongy,

pallid white tinged yellowish, in the stem pale brownish, not changing colour on

exposure or bruising.

Spores not darkening in potash. Basidia 34-40 x 9.5-l1y; sterigmata 4, 4.5,

long. Cystidia 50-128 x 14-20n, cylindric to subventricose, apex obtuse, pro-

jecting, thin-walled, abundant on the gill-surface and along the fertile edge. Surface

of the pileus with a disrupted pile of hyphal ends —800. long, forming the fibrillose

squamules, composed of subcylindric rows of cells 23-85 x 8-18, the end-cells

obtuse or more or less attenuate (even subappendaged), with brown contents and

thin brown incrustation. Surface of the stem covered with a disrupted sterile

hymenium of subclavate and immature basidia and rather scattered cystidia (as

on the gills). Hyphae of the flesh 3-18» wide, without clamps; longitudinal in the

stem with the cells 25-120u long, 3—6u wide at the surface of the stem, and with

somewhat thickened walls in the interior of the stem. Gill- trama with mucilaginous

longitudinal hyphae 5-15, wide.

This species with such deeply alveolate-reticulate gills comes in size of fruit-

body between the Malayan P. ochraceobrunneus Corner (1970) and the north

temperate P. rhodoxanthus (Shw.) Bres.; it differs from both in the pallid (not

yellow) gills, the squamulose pileus, and the short spores. It may be closer to the

Malayan P. bellus (Mass.) Corner which has the small spores, also pale under the

microscope, a merely villous-subtomentose pileus, and yellow gills reticulate only

at the base. The North American Boletinus squarrosoides Snell et Dick (Mycologia

28, 1936, 468), referred later to Phylloporus and now to Xerocomus, may also

be related; it differs in the more or less reticulate yellow stem, yellow gills poroid

in the outer part of the pileus, and the narrow spores (7-10 x 3.5-4.5y). Com-

pare, also, Paxillus squarrosus McNabb (1969) with dichotomous, not poroid, gills

and rufescent flesh.

P. stenosporus sp. nov.

Pileus 4 cm latus, siccus, subtomentosus v. fere laevis, pallide cervino-brunneus, aetate

subgriseus. Stipes 22 x 7 mm, basim versus attenuatus, brunneo-albidus, ex integro albido-

pruinosus; mycelio albo. Lamellae decurrentes, subdistantes, primariae c. 30, 3-4 mm latae,

ordinibus 3-4 instructae, nec reticulatae nec ‘dichotomae, flavidae. Caro crassa, albida, in

stipite brunneola, immutabilis, Sporae 9.5-11.5 (-12. 5) x 3.7-4.2y, ochraceae s.m., laeves,

2—3-guttatae. Cystidia 60-120 x 14-20u, tenuiter tunicata, copiosa.

Ad ripam juxta silvam, solitarius. Java, Sukabumi distr., Tjiletuh, Corner J-23 (CGE),

29 April 1972.

Spores narrowly boletoid, not amyloid, not darkening in potash. Basidia 9-10p

wide, 4-spored. Cystidia 60-120 x 14-20u, ventricose with slightly prolonged

obtuse apex 6-9 wide, varying clavate or subcylindric, tapered into the narrow

subhymenial base, abundant on the gill-surface and edge. Surface of the pileus

evidently with a pile of inflated hyphal ends -170. long when young, but the

pile becoming flattened and disorganised with age, the hyphae cylindric or, if

short-celled, then submoniliform, the cells 18-120 x 8-17, the end-cells obtuse.

Stem with a disrupted sterile hymenium. Hyphae 8—20u wide, without clamps.

The pale narrow spores distinguish this species which seems near to the

slender P. brunneolus Corner (1970) which has rufescent flesh and slightly cyanes-

cent gills.

16 Gardens’ Bulletin, Singapore — X XVII (1974)

STROBILOMYCES Berk.

S. velutipes Cke et Mass.

Pileus with appressed scales. Stem reticulate only at the apex. Flesh slowly

turning red then black.

Sarawak, Bako National Park, Corner P-159, 26 Aug. 1972; Semanggoh

Forest, Corner s.n., 19 Aug. 1972.

References

Coker, W. C. and Beers, A. H. (1943) — The Boletaceae of North Carolina.

Corner, E. J. H. (1970) — Phylloporus Quél. and Paxillus Fr. in Malaya and

Borneo. Nova Hedwigia 20, 793-822.

(1972) — Boletus in Malaysia. Government Printing Office, Singapore.

Grand, L. F. and Moore, R. T. (1971) — Scanning electron microscopy of basidio-

spores of species of Strobilomycetaceae. Can. J. Bot. 49, 1259-1261.

Hongo, T. (1970) — Notulae mycologicae (9). Mem. Shiga Univ. 20, 49-54.

McNabb, F. R. (1969) — The Paxillaceae of New Zealand. N. Z. J. Bot. 7, 349-362.

Singer, R. (1945) — The Boletineae of Florida, with notes on extralimital species.

Farlowia 2, 138.

Snell, W. H. and Dick E. A. (1970) — The Boleti of northeastern North America.

Contribution to the Knowledge of Cecidia of Singapore

MING ANTHONY

Laboratoire de Botanique Tropicale, Université de Paris

This report is based on the author’s thesis for the “Diplome d’Etudes

Approfondies”’ presented before the Faculty of Science, University of Paris, in

July 1970. The work was done partly at the Department of Botany of the

Singapore University in 1968-69, which is under the direction of Professor A. N.

Rao, and partly in Paris in the Laboratory of Tropical Botany in 1969-70 under

the direction of Professor R. Schnell.

I should like to acknowledge with gratitude the help from many taxonomists

who assisted me constantly throughout my research. In particular, | am indebted

to Dr. H. Keng and Mr. D. H. Murphy of Singapore University who spent so much

time in the determination of plants and gall-makers respectively and for their

advice; to Dr. G. Gusset of Paris University for his help in the naming of plants

and on morphological questions; to Dr. W. L. Chew then Botanist at Singapore

Botanic Gardens and Dr. M. A. Rifai, Director of the Herbarium of Bogor Botanic

Gardens, for their help in taxonomy.

I am grateful to: the Directors of the Singapore Botanic Gardens, who gave

me all facilities to collect throughout the Nature Reserves; Dr. J. Meyer,

Director of the Strasbourg Laboratory of Cecidiology and his team of research

workers for their aid and their advice in cecidiology; Dr. W. Nijveldt (Instituut

voor Plantenziektenkundig Onderzoek, Nederland) for the determination of gall-

midges; Mr. R. D. Eady, Dr. G. J. Kerrich, Mr. C. E. J. Nixon of the British

Museum for the determination of the parasitic wasps; Dr. J. W. Beardsley

(University of Hawaii), for the naming and description of coccids; and Dr. H. H.

Keifer (Sacramento, USA) for the determination of mites.

Finally I thank Mr. S. Murphy, Mr. C. Anthony, and the editor who helped

me with the translation of my thesis.

INTRODUCTION

I began my study in cecidiology in Singapore where material was collected

during the academic year 1968-69, and where subsequently I have had the

opportunity to enrich my collection and to make new observations during two

study trips in January and April 1970.

Singapore is a small island in the Malesian Region, situated at a latitude of 1°

21’ North. It is exposed, at the same time, to an equatorial climate and a monsoon

climate, uniformly hot and humid, nonetheless temperate because of the presence

of the sea( Bib. 23; fig. la); the average annual precipitation is over 2.41 m. with

seasonal maximums staggered from November to January.

18 Gardens’ Bulletin, Singapore -— X XVII (1974)

tchalle de i:19000009

0 _ AG 40 590

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S2=ELoUR 275. 2,

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nut 4

Fig. 1. a, location of Singapore adapted after the “Nouvel Atlas Général” (Bordas ed.);

b, map of Singapore adapted after “Modern Singapore” (1969. Ooi Jin-Bee,

Chiang Hai Ding ed., 285 pp.); ¢. part of the Bukit Timah Nature Reserve,

adapted after an unnumbered plan. The stippled area represents my field of

study. Contour numbers are in ft.

Meanwhile, in the course of ‘‘winter’’ 1968-69 rains were unusually low. This

climatic anomaly certainly influenced the flora and its pathology, and therefore

must have affected my results. I unfortunately cannot assess the importance,

having made my observations over such a short period. Material was collected

leisurely during walks in private gardens and along the wayside. A limited area

in the Bukit Timah Nature Reserve was chosen for systematic collecting and

study (fig. 1, b & c). This Reserve is according to H. M. Burkill*, a block of 184

*H. M. Burkill: A Survey of Nature Conservation on Singapore Island. The Proceedings

of the Ninth Pacific Science Congress, 1957. Vol. 7, 1959.-pp. 34-37, 1 map.

Cecidia of Singapore 19

acres of lowland forest clothing Bukit Timah (Malay for “Hill of Tin’’). It is

granite and without any tin deposits. With its two peaks both slightly over 525 ft

the Reserve is the highest ground on Singapore Island. It is one of the first

Forest Reserves made after Cantley’s report* and has never been exploited for

timber.

One should note that a large number of Shorea curtisii King** saplings

which compose the undergrowth in my field area, is the result of the germination

of anemochorous and heliophilic seeds. This testifies to a recent thinning of

the forest.

Under ‘‘Material and Technique”’ is a description of the method I have used

and perfected for my study. The cecidia that I have collected are listed with notes.

This list follows the style of those of Houard (Bib. 28), and of W. and J. Docters

van Leeuwen-Reijnvaan (hereafter D. v. L. R.) (Bib. 22), and consists of a

succint morphological description of my specimens, illustrated with line drawings

and plates. As far as possible, a few morphological and physiological interpretations

relative to the process of the cecidiogenesis are given. They are based on the

recent works of Strasbourg Laboratory (Bib. 39). Adopting the ideas of Kloft

(Bib. 34), Maresquelle and Meyer (Bib. 38), I have included in my list all

developmental anomalies of parasitic origin, where the anomaly has bearing on

the positive or negative growth, or also on the cellular differentiation.

HISTORICAL REVIEW

The only works of importance concerning cecidia in the Malesian Region are

those of D. v. L. R. (Bib. 6 to 22), and of Houard (Bib. 28). There are also

several previous Indonesian studies generally relevant to the pathology of economic

and agricultural plants, unfortunately disseminated in journals of local interest

and unavailable even in Singapore. The multiple studies of D. v. L. R. on

Indonesian cecidia, complemented by the works of Nalepa (Bib. 40 to 44), on

Acarina, and of Karny (Bib. 30 to 32) on the Thysanoptera are of fundamental

interest. The systematic list of Houard relevant to the zoocecidia of the whole

Malesian Region are of invaluable interest. Nonetheless one must know that the

galls of this immense Region fractioned in a pleiad of islands and islets, are

far from being well known. No previous cecidiological study in Singapore has

been published.

MATERIAL AND TECHNIQUE

GALL COLLECTING AND ITS PROBLEMS

The complete study of each cecidium necessitates much material; several

duplicates are required for fixing, making of herbarium specimens, and separately

for extricating gall-makers for the purposes of culture and despatch to various

specialists for the determination of the host plants and gall-makers.

In the list, gall specimens are numbered in Arabic, identically as the host

plant. Where different types of galls are found on the same host, each cecidium

is labelled with an additional proper number in Roman.

*N. Cantley: Report an the Forests of the Straits Settlements. 1883, fide H.M. Burkill.

**All host-plants here published under this name are recently determined, thanks to

Dr. K. M. Kochummen, Forest Botanist at the Forest Research Institute in Kepong. They

appear under the erroneous name of Shorea pauciflora in earlier publications.

20 Gardens’ Bulletin, Singapore — X XVII (1974)

In order to make a complete study of a gall, various stages of its development

are needed. With some species, these could be found during a single collecting

trip, e.g. the mite-galls on Litsea elliptica (Bl.) Boerl. and Actinodaphne sp. Often,

repeated collecting was necessary as is the case of the psyllid-gall on the leaf of

Notaphoebe kingiana var. malvescens Gamble where juvenile stages were found

in December-January and late stages only in April. Similarly were the coccid-galls

on Shorea curtisii King (gall 231 and gall 231Va) and the wasp-gall (gall 23I1Vb)

on the same host: in December only a few juvenile stages of gall 231 whereas in

April a considerable number of young stages were available. On the other hand,

in the same interval, juvenile stages of 231Va and 23IVb were totally absent. The

wasp-gall (gall 120) on Lithocarpus conocarpus (Oudem.) Rehd. was first collected

end March 1969 when it detached itself from the leaf, taking with it the larva in

the fall. Despite methodic collecting March-September 1969, and in January 1970,

that type of gall was not seen again. Finally in April 1970, I found leaves that bore

marks of recent drop of galls. These observations lead me to conclude that galls

120 have a rapid development in February and March.

The observations above seem to confirm those made in temperate climates

(Mani — Bib. 36. p. 197) to the effect that the majority of cecidia would develop

strictly during a definite moment of vegetative growth of the host plant. Previous

publications on the development cycle of galls refer to work in temperate climates

(ibid. Chap. X, pp. 196-212) where the seasonal alternation fundamentally

influences the vegetative rhythm of the host plant. No study has yet been made

in the humid tropics where inspite of the absence of prominant seasons certain

galls seem to have their own rhythm of development.

These observations suggest that a statistical correlative study in the tropics

would be interesting. It would perhaps be possible to establish a calender of gall

development in Singapore in relation to the climate cycles, the internal rhythm of

growth of the host plant and the cycle of the development of the gall-makers.

PRESERVATION OF THE COLLECTION

The most complete botanical specimen, carrying flowers and fruit if available.

and the one with the least parasites are reserved for determination. Several parts

of which the galls are well formed, ripe or already opened, are kept for the

herbarium. One important part is put aside for obtaining or rearing the gall-makers.

Juvenile or intermediate stages are fixed with F.A.A. for a morphological and

anatomical study of the development.

Herbarium. For each type of gall two sets of herbarium sheets were made. One

set is now at the Laboratory of Tropical Botany of Paris, the other at the

University of Singapore, Botany Department.

The conventional techniques including the drying under pressure used for

preservation in the herbarium are not to be recommended in many cases. Certain

galls have a characteristic relief and a disposition which are useful to preserve.

One can, for this reason, dry out the plant-part carrying the galls in a crystal

powder of CuSO4 dehydrated by heat before use. However this method presents

the inconvenience of crowding and material prepared this way is very fragile.

Fixing in F.A.A. After drying, the fleshy galls shrink and become unrecognisable.

It is better to fix them with F.A.A. I intend to make more precise anatomical and

morphological study of the development of the galls subsequently. In this study

Cecidia of Singapore 21

a series of permanent hand-sections were made by using a method taken from

Johansen (Bib. 29): (a) technique of fixation, sectioning, staining (p. 102); (b)

schedule of dehydration by T.B.A. (pp. 110-113). From these sections it was

possible to make anatomical observations on certain cecidia which are listed.

Using the methods described above, more than one hundred galls were

collected and preserved. This collection represents only a fragment of the galls in

Singapore. Based on the 1536 numbers of galls of Indonesia, as given by D. v. L.

R. (Bib. 22, p. 31), it is anticipated that the present collection will grow considerably

in the years to come.

OBTAINING AND NAMING GALL-MAKERS AND

INIQUILINES

This study necessitates much time and patience and can only be achieved

by a resident worker in the area. Adult gall-makers are difficult to obtain in

sufficient numbers to permit a determination. Thus my conclusions are very

fragmentary and incomplete. There is no general method for obtaining parasites:

the technique to be employed depends largely on the type of the gall-maker.

MITES

With the help of Mr. D. H. Murphy, I have tried out and perfected several

methods to obtain mites: —

(a) Showing the gall-mites in situ. The numerous morphological convergences

between mycocecidia and acarocecidia (yellowish or orange swelling of the lamina,

hypertrophy of the epidermic cells into hairs, parenchymatous, fleshy growths)

had caused some confusion to early cecidiologists. (Mani-Bib. 36, p. 157). A

method which can distinguish the presence of acarocecidia from mycocecidia and

which stains both mites and fungal mycelium alike (if present) is adapted here

from one in use for research on nematodes. So far. fungi are absent in the galls

examined. The process is: —

— Section the fresh and dry galls.

— Place them in a test tube and cover with a mixture composed of 5 parts of H. O.

(20 vol.) and 1 part of ammonia (880) for 24 hours. This is for decolourization of

plant tissues.

— The mixture is discarded and replaced with a solution of Aman’s lactophenol

mixed with 0.05 gm cotton blue stain.

— After boiling the test tube in a double boiler for | hour, leave galls in stain for

at least 24 hours.

-— The liquid is discarded, specimens are washed with pure lactophenol which is

also used as a liquid for preservation.

— Mount specimen in drop of lactophenol.

The cuticle of mites absorb the blue stain but its opacity renders the method

unsuitable for specific identifications of acarocecidia. For this the most classic

method is one which is perfected by Nalepa (Bib. 40: vol. V: p. 49).

(b) Collecting mites by Nalepa’s method. After driving out the mites from the galls

by gradual drying, they should be fixed with heated chloropicric acid. The 70%

alcohol stage used by Nalepa is here bypassed because according to Keifer’s work

22 Gardens’ Bulletin, Singapore — X XVII (1974)

cited by Owen Evans, Sheals and Mac Farlane (Bib. 45), alcohol is a bad medium

for preserving mites, and these authors preferred to preserve them in the liquid

of fixation, With this method a large number of mites can be obtained, but it

necessitates much gall-material. On the other hand, I have noticed as did D. v. L.

R. (Bib. 22; p. 17), that the success of this experiment depends on the humidity

of the air: if the atmosphere is too humid, the gall goes mouldy, and the mites

are killed inside.

more economical in material and independent of atmospheric conditions, was

developed by Keifer as cited by Owen Evans, Sheals and Mac Farlane (Bib. 45;

pp. 67-70). Their method of soaking in a solution made of resorcinol (50 g.),

diglycolic acid (20 g.), glycerol (25 ml.), iodine water (about 10 c.c.), has been

tried without success as I have not been able to needle the mites out of this

solution which turned black and thick. That led me to investigate other methods

of soaking based on the destruction of cellulose in the plant cell: —

Method I: Soaking in a solution of enzymatic extracts of digestive tubes of

a snail; the snail is the only multiple-celled animal capable of digesting cellulose

directly.

Method II: Soaking in Schweitzer’s solution that dissolves cellulose and not

chitin.

INSECTS

I set out to culture gall-midges and wasps, and inquilines but, cecidiological

moths and beetles being rarer, did not provide sufficient material for applying the

methods described below to culture their larvae : —

(a) Method of D. v. L. R. (Bib. 22, m. 17). Galls are collected, opened up, and

placed with their pupated parasites in a closed tube with a piece of gauze. Many

wasps have been obtained by this method. The rearing of Diptera was much more

difficult: the pupae of midges were very weak, they dried out and became mouldy.

These unpredictable results led me to develop another technique described here.

(6) Placing cages on the gall-bearing plant. For descriptions and usage of various

types of cages see Leach*.

It is necessary to have an opening covered over by a piece of gauze, thereby

permitting aeration of the attacked part contained in the cage, and avoiding

excess condensation which may cause (1) galls to mould before the opening or (2)

the parasites to drown and to decompose rapidly after their exit. This method

seems perfectly adapted to the rearing of winged gall-producers. It permits the

observer to follow the development of the gall-makers and their cecidea in situ

under quasi-natural conditions. The disadvantages are subsequent re-location of

cages in the Nature Reserve and risk of destruction of those deposited in public

areas. Also, leaves may succumb under the heavy weight of the cage and insects

such as wasps and ants tend to cut out the gauze for making nests or to colonise

the cages.

Frequently, several different sorts of animals are obtained, e.g. Diptera and

Hymenoptera; it is at first sight difficult to discern as to which is the gall-maker,

the parasite or the inquiline.

*J. G. Leach: Insect transmission of plant diseases. McGraw-Hill Book Co., Inc. New

York and London. 615 pp. 238 figs. 1940.

Cecidia of Singapore 23

(Reena a

Fig. 2. Cages for rearing gall-makers; a, for foliar galls and acrocecidia; b, for pleurocecidia.

c; transparent plastic cylinder; cc: cylinder cut; cg: gauze glued with chloroform

to the cylinder; ga: gall bearing organ; s: perforated plastic stopper; sc: stopper

cut; sg: gauze held in position by the stopper.

PROBLEMS ON DISTRIBUTION OF GALLS AND

GALL-MAKERS

Range of geographical distribution

Of the hundred galls that are in my collection, only ten have been reported

by workers in other parts of Malesia. This applies proportionately also to gall-

producers: none of the animal species submitted to specialists for determination

has been hitherto described. Thus the list contains many unidentified species. A

new genus of Diptera was found on Calophyllum ferrugineum Ridl. (gall 68).

Another, new genus of Coccidae is Gallacoccus with species G. secundus (Gall 23

Ij) and G. anthonyae (Gall 23 V) both on Shorea pauciflora King described by

J. W. Beardsley*.

*J. W. Beardsley: A New Genus of Gall-Inhabiting Eriococcidae from Singapore

(Homoptera: Coccoidea). The Proceedings of the Hawaiian Entomological Society, (1971)

Vol. 21 (1) 31-39. N.B. See footnote on p. 19.

24 Gardens’ Bulletin, Singapore — X XVII (1974)

As many as seven different coccid-galls were collected; three occurred on

Shorea curtisii King (Dipterocarpaceae), one on Hypserpa cuspidata (Wall).

Miers (Menispermaceae), one on Calophyllum inophylloide King (Guttiferae), one

on Xylopia malayana Hk. f. and Th. (Annonaceae), and one on Lithocarpus sun-

daicus (Bl.) Rehd. (Fagacae). Now, all the works that I have consulted indicate

that coccids occupy a minor place among the gall-makers, except in Australia

where they attain their maximum development. (Bib. 28, Vol. I, p. 11, Vol. IL, p.

590. — Bib. 36, p. 27. — Bib. 22, p. 13). It seems so far that the galls on the flora in

Singapore are very special and original: this endemism is certainly linked with

the insular character of the region.

On Mount Kinabalu in Borneo, a coccid-gall on a young Dipterocarpus?

eracilis Bl. was collected by E. J. H. Corner in 1963 (Bib. 2, pp. 339-341). This

gall presents many analogies with Gallacoccus secundus i.e. gall 231 on Shorea

curtisii King. The discovery of gall-coccids in Singapore with related representa-

tions in Borneo suggests a wider distribution and cecidiologists will perhaps be

brought to reconsider the importance of coccids as gall-producers.

Ecological Problems on the distribution of galls

Plants of Actinodaphne sp. seemed to me equally spread out on the slope

ABC limited by Ginger Walk and Jalan Kutu (fig. 1, c). Actinodaphne sp. carries

several galls, two of which are caused by midges (gall 2411 & 99III, see p. 38).

One can find gall II everywhere in this location. In contrast the occurrence of gall III

seems strictly limited around point C. Why is the distribution of these two midge-

galls not homogeneous? Although a statistic study is necessary to confirm my

findings, I can nonetheless, beginning with these observations, show up the problems

on the distribution of the gall-makers and their ability to propagate. An ecological

study of the habit of these two kinds of Diptera in relation to the variations of the

microclimatic factors such as humidity, temparature, sun and others should be

also interesting.

Distribution of galls and gall-makers in the different plant families (table 1)

The table here follows the style of D.v.L.R. (Bib. 22, pp. 28-31). It does

not claim to embody exhaustive data but the following are noteworthy:

— The plant families of my list carrying the greatest number of different galls

are the Dipterocarpaceae, Guttiferae, Myrtaceae, Lauraceae and Euphorbiaceae;

the last three correspond to the collections of D.v.L.R. (ibid p. 28).

— Concerning the gall-makers, there are many obscure points. However midges

are preponderant among gall-producers and occur in nearly all the regions of the

world (Bib. 36, p. 30).

—In Bukit Timah 25 dipterocecidia and 7 acarocecidia were collected. The

ratio is comparable to that of 4:1 as given by D.v.L.R. (Bib. 22, p. 26) w.r.t. humid

forests in S. E. Asia. Future collections will perhaps modify my current results

though the present data confirm the assessments of those authors and correspond

with data related to forest and climatic types as given by them i.e. to the parti-

cularly high humidity of the undergrowth of Bukit Timah forest despite the relative

dryness of the November 1968 — January 1969 period.

TABLE 1

Gall-maker Mite Midge Wasp Moth | Thrips Insect | Fungus ? | Total

Plant-families Ae eee Ieee ean ior a. balsa. ib apabe las he by anal bilkca Wy

Anacardiaceae 2 | 3

Annonaceae Se Se ee ewe els oe 6

Apocynaceae eee eee on ee eS es hae aaa 1

Me clepiadiaceae re ee eae ee rhea Ira 1

Burseraceae ie race hee he. nee ret ea 4

Combretaceae ce eee eS ree) Sy ae ee ae Pri 1

Celastraceae ree Get sean aap re 1

Dipterocarpaceae ‘ogee ae a ae E35 Oe ie el ae ee a at Ey Pe ee rar ee 10

Euphorbiaceae IL ae Seg a a Vs Pen eee aes 8

Fagaceae ° eee et eo hh, Se ae an | he ae Rae vn Pheri y ea ea 5

Guttiferae . . ee aS Se ee ene ee eer aie nyc 9

Tauraceae . . | et eae ees iar ie rae ie ie Ere ale et 12

Leguminoseae Mma hao he ae eS ay 4

Loganiaceae Sieh ieee aan. Peat ein 22

Menispermaceae WT eR Sra ee ites aa a a 8 Se Se 1

Moraceae . . eas eee ea eS SG aa ea 3

Myristicaceae ae ee ee a eS ae aie re mie 3

Myrsinaceae ee ae aes | a ae Wetcnceet ce Se 1

Myrtaceae Sa a Sa ee es ae Bee cli eat as

Rhizophoraceae oer oe hen Pe cles aan a. = ie ae Recep tel 1

Rubiaceae . . ere es eae ee Sen ea rey ea aie ee oe. 3

re Sr eee le

Tiliaceae =e aa es a ee ee ee ee ie ee a ee a 1

Verbenaceae ee ee ee a ae aie x el Sa 6

ee Ten Tete oer see ia) Sa

a on re ae gs Pe Te 1

Total b / 75)

Totala+b 10 Poy | 7 5 9 12 1 8 103

b: galls from Bukit Timah Nature Reserve

a: galls from other localities

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Cecidia of Singapore 27

CONCLUSION

My results show that the gall flora of Singapore is special when compared

with those in Malesia hitherto published.

(i) About ten only of the cecidia of my list have been mentioned in previous

studies, the other galls appear, for the most part, on host-plants not mentioned in

the works of Houard or D.v.L.R.

(ii) Moreover, my coccid-galls appear in the present stage of knowledge as

pathological forms particular on the island of Singapore. Also my discoveries of

coccids, added to that of Corner in Borneo, commit me to question the dogma

generally accepted, i.e. that these Homoptera are hardly important as gall-producers

except in Australia.

The list shows that, despite their particularities, my cecidia present certain

typical characters of humid tropicai regions: abundance in some families such as

Lauraceae, Myrtaceae, Euphorbiaceae, and also abundance of gall-midges and

thrips.

Certain types of galls, provoked by the same animal genus on the same plant

genus are extremely widespread in the tropical world, for example the psyllid-galls

on the Eugenia genus “distributed from East Africa to the Gambier islands” (Bib.

28, p. 592) are numerous in the Malesian Region (Bib. 28, pp. 593-598; Bib. 22,

pp. 407-417), and in particular in Singapore (see list). This poses a phytogeo-

graphical problem of distribution and dispersion of the flora with the associated

fauna.

Finally, it is noted that, though the action of gall-makers belongs to the

same zoological group, galls of identical aspect and structure appear on taxono-

mically distant plant species. Examples are the ball-shaped, parenchymatous,

fleshy midge-galls of my list: gall 168I on Shorea sp. (Dipterocarpaceae), gall, 13II

on Grewia blattaefolia Corner (Tiliaceae), gall 10711 on Calophyllum pulcherrimum

Wall. (Guttiferae). In the formation of a gall two fundamental factors are distin-

guished: first, the role of the animal and second that of the host-plant. In the

cases mentioned above, it would seem that under the influence of certain gall-

makers, the same genetic factor of localisation would intervene in the gall morpho-

genesis.

ANNOTATED LIST OF GALLS

ANACARDIACEAE

Mangifera indica Linn. Gall. 140. Pepis Road (garden).

Midge-gall affecting the two faces of the lamina. Adaxially the cylindrical gall

is 1.5 mm to 2.5 mm in diam., and | to 1.5 mm in height, often around the gall

the lamina is elevated. The distal region of the cylinder has a small lid. Abaxially

the gall is cone-shaped: 1.5 to 2 mm high, the diam. of the base 1.5 to 2 mm.

Inside is a very small circular larval cavity. The galls on the specimens in Singapore

were mostly located along the midrib and side veins.

Distribution: Singapore-Java, Rhio Archipelago, Sebesy Island, Celebes, Malacca.

See Bib. 7, p. 107. No. 66, fig. 36-9, p. 187, No. 189.11, p. 92, No. 12.15, p.

36, No. 41.20, p. 303, No. 43.21, p. 137, No. 53, fig. 28-22, p. 325, No. 803,

ge. 519.=28, p. 467, No. 1719 & 2/1721.

28 Gardens’ Bulletin, Singapore — X XVII (1974)

Idem. Gall 82; Singapore, Changi Road, il4 m.s.

Midge-gall looking like the preceding one but smaller, Adaxially, the cylinder

is 1 mm high, and 0.5 mm in diam.; the apex of the cylinder is rounded. That is

where the dehiscence by a pore is located. The gall is not surrounded by an

elevation of the lamina as in the preceding case. Abaxially, cone of 1 mm high

on | mm in diam. of base. It seems to correspond to the description of Houard

(Bib. 28, p. 467, No. 1721).

Semecarpus sp. Gall 59 & 46. B.T.N.R.

Psyllid-gall. Disc-shaped thickening of the lamina 4 mm in diam. on 1 mm

thickness, visible on both faces of the leaf. The exit pore of the gall-maker is

abaxial and the tissue around it is slightly elevated.

ANNONACEAE

Artabotrys suaveolens (BI.) Bl. Gall 22 I, 26 & 127. B.T.N.R. (Pl. 1).*

Midge-gall, attached to one or the other face of the lamina and sometimes to

the twig by a large circular base 5 mm in diam. If the leaf is very infected it

becomes atrophic. The young gall is horn-shaped, more or less curved, up to

2.5 cm long. In the course of its development, its inferior third part becomes

hypertrophic and is 5 mm across; the superior part keeps its conical shape. In

the centre of the swollen part is the larval chamber, surrounded by a hard and

fibrous shell that digitates into the white parenchyma surrounding it (Pl. 1, e). The

gall is covered with a green and glabrous epidermis. In the gall-chamber is only

one midge-larva. On maturity the gall cavity is much widened; its wall becomes

blackish. The midge makes its way out by piercing a large canal through the wall

of the swoilen part of the gall (Pl. 1, d, f), and leaves its pupal integument partly

inserted in the gall (Pl. 1, b). Very often the gall contains a parasitic moth. There

is then deviation of the cecidiogenesis: the swollen part becomes preponderant,

reaching 1.4 mm in diam. The superior conical part is reduced to 2 mm in height.

The moth larva eats the internal part of the gall and, after metamorphosis leaves.

the gall through a large circular pore (Pl. 1, c). When the parasite or gall-maker

has left, the gall detaches itself easily from the leaf, therefore inducing the forma-

tion of an alveole on the lamina (PI. 1, a, d).

Distribution: Singapore-Java (Bib. 22, p. 185, fig. 285).

Idem. Gall 22 “Kk. B. TIN Ry (Ete):

Psyllid-gall, very wide-spread in the Nature Reserve. The lamina is marked

by conical elevations, 0.5 mm high, on a base 1 mm in diam., caused by the young

psyllids attached abaxially.

Cyathocalyx ramuliflorus (Hk. f. and Th.) Scheff. Gall 100. B.T.N.R. (fig. 15. a).

Mite-gall. Mamiliated pockets that pouch out from the adaxial leaf-suriace

between the side-veins, 3 to 5 mm high. The interior of the cavity is covered with

an erineum that is brown when the gall is adult.

Popovia tomentosa Maingay Gail 70 & 151. B.T.N.R., Jalan Kutu and Jungle

Walley. Gig. “13%: (Pl as ay.

*Figures 3 to 18 and all plates follow bibliography.

Cecidia of Singapore 29

Midge-gall often attacked by wasps. On the stem or leaf-stalk, the gall is

ovoid, 2 mm in height and 1 mm at the widest part, fixed by a large base. The

Opening is apical. On the leaf (PI. 11, d), the galls are attached laterally along the

midrib, usually packed closely in a more or less coalescent row. The isolated gall

presents itself adaxially as a protuberance 2 mm in height and 1 mm wide at the

base, and abaxially is a less distinct swelling, less than 1 mm in height. The

large gall-chamber, containing only one midge-larva, is located at the level of the

lamina; around it one finds the nutritive tissue (nt), a protective shell (s), and a

tanniferous parenchyma. Finally the gall is covered with a hairy epidermis (fig.

13, c). The opening is apical and adaxial. After the exit of the gall-maker, one

notices that the gall-chamber is plastered with a hard and blackish covering.

Xylopia malayana Hk. f. and Th. Gall 154. B.T.N.R., between Ginger Walk and

Jalan Kutu. (fig. 12, c).

Pleurocecidium caused by a coccid belonging to the family Lecaniodiaspididae,

Lecaniodiaspis sp. On the scale-mother periphery, a fusiform wad develops in the

direction of the axis of the stem, 1 cm long and up to 6 mm wide. The large

Opening of the gall cavity is covered by a waxy and resinous scale, 5 mm across,

secreted by the mother. The female deposits its eggs beneath this scale. Under the

growth of cortical tissues that actively participate in gall morphogenesis, the bark

is fissured.

Undetermined ANNONACEAE. Gall 164. B.T.N.R., Jungle Valley.

Psyllid-gall. Conical elevation of the lamina, less than | mm in height,

provoked by psyllids attached abaxially.

APOCYNACEAE.

Willughbeia coriacea Wall. Gall 28. B.T.N.R.

Cause not determined. Fusiform swelling | to 1.5 cm wide, affecting the stem

for several cm in length. Under the effect of the growth of internal tissues (xylem,

phloem, cortical parenchyma), the bark cracks and breaks up into fragments.

ASCLEPIADIACEAE.

Hoya diversifolia B\l. Gall 90. Dalvey Road.

Insect-gall. Leaf-gall affecting only the upper surface of the lamina. It

presents itself as a slight purple protuberance of 5 mm in maximum diam, yellow

in its centre.

BURSERACEAE.

Canarium pilosum A. W. Benn. Gall 69 I, 83 & 109. B.T.N.R. (PI. 5, c).

Gall caused by a mite, Eriophyes sp. close to E. reijnvannae Nal. fide Dr.

Keifer in litt. Hairy clavate finger-gall in 1 cm height by 5 mm width at the

maximum, projecting out from the adaxial side of the leaflet, has a small abaxial

Opening and is covered inside with straight hairs. This type of gall occurs frequently

on the genus Canarium. For example: Canarium hispidum (Bib. 28, P. 422, no

1550.—22, p. 262, no 627, fig. 436—16, p. 28, no 531, fig. 531); Canarium sp.

(Bib. 22, no 627 & 629, p. 263, fig. 438).

30 Gardens’ Bulletin, Singapore — XXVII (1974)

Idem. Gall 165 Il. B.T.N.R.

Thrips-gall. The leaflets become involute.

Idem. Gall 121 Ill. B.T.N.R., between Jalan Kutu and Main Road.

Midge-gall (Pl. 5, a, b). Ovoid cecidium, 2 mm long, less than | mm wide,

attached to the veins by a large base and opening by a round apical pore, when

mature. Inside is a large gall-chamber containing a midge-larva. The internal wall

of the gall is grey or blackish.

Idem. Gall 121 IV. B.T.N.R., between Jalan Kutu and Main Road.

Cause not determined (PI. 5, d). Under the influence of the parasite, there is

inhibition of the vein-growth, which creates the pleating of the lamina of leaflets

the edges of which, moreover, tend to become involute.

COMBRETACEAE.

Terminalia catappa L. Gall 145. Campus of the University of Singapore and

Angullia Park.

Mite-gall. On the adaxial face of the leaf this yellow, green or red pouch-

gall forms a rounded protuberance less than 1 mm high; the ostiole of the gall is

abaxially opened and leads to a small chamber.

Distribution: Singapore — Java, Krakatau Island, Sumatra, Salajar Group. (Bib.

15, p. 43, no 64, fig. 64.—16, p. 68; no 674.—17, p. 78, no 18, fig. 18—28, p. 582,

no 2104.—22, p. 404, no 1048, 20, fig. 752). This gall has a large distribution area,

following the pantropical character of the host-plant.

CELASTRACEAE.

Salacia korthalsiana Miq. Gall 169. B.T.N.R.

Insect-gall. Fusiform swelling of the midrib, abaxially visible. The main vein

can be affected locally and/or along its entire length. In the second case the lamina

is contorted towards its abaxial side. The side-veins can also become hypertrophic

at their junction with the midrib. The gall manifests itself adaxially by the unusual

width of the affected veins. The exit of the gall-makers is abaxial.

DIPTEROCARPACEAE.

Shorea curtisii Dyer ex King Gall 23 I. B.T.N.R.

Gallacoccus secundus Beardsley, new genus and species.

—— Description of an adult gall (Pl. 2, a, b, e): purplish-green pear-shaped gall,

3.5 cm long, 2 cm wide at its maximum. Its irregular surface is covered with bumps

each of which has an apical tuft of hairs. It is formed of foliaceous appendages

overlapping at their apex, with more or less thickened and fused bases. The two

median appendages, larger than the others, separate between them a large ovoid

gall-chamber containing many coccids.

—— Location on the host (figs. 3, 4; Pl. 2): it is found in the axil of the leaf or

on a leaf vein. Therefore this gall is hard to interprete without precise ontogenical

study. The study of the young stages shows that this gall occurs either on one of

the internal stipules (fig. 3, a, b; fig. 4. b), or on one of the young leaves of the

Cecidia of Singapore 31

bud (fig. 4, c, d; Pl. 2, f). Normally the stipules are deciduous but if one of them

has parasites, it participates in its totality to the cecidiogenesis without affecting

the growth of the rest of the bud (fig. 3, a, b). That is why at the adult stage this

gall gives the impression of being in the axil of a leaf: in fact it is in a lateral

position since it is a stipule transformed by the cecidiogenesis. An anatomical study

would be necessary to pinpoint to what degree the axillary bud of the leaf is

affected. At the beginning of the development of the gall, a wad is always formed

(fig. 3; 4), then there is the outgrowth of the foliaceous appendages. That leads

me to think that after the installation of the gall-maker, there occurs, dedifferentia-

tion, formation of a callus, then of an abnormal bud. One can note the resemblance

of the gall 23 I with those figuring in the bibliography: 2, pp. 339-341, fig. 1—-2.—22,

p39), fig. 721.

—— Deviation of the cecidiogenesis due to the presence of secondary hosts in the

coccid-gall 23 I on Shorea curtisii King (Pl. 2, c, d). For an analogous problem

see Bib. 39, Vol. 33, section 3, pp. 143-148, 2 Pl. The galls attacked by parasitic

wasps appear almost normal except that they never reach their maximum size.

They are more woody and dried out than normal galls. Moreover the parasitic

wasps do not leave through the apical canal, but through apertures ‘which they

make in the gall-wall. The gall-cavity is very reduced and irregular; coccids are

present but in very small numbers. On the other hand, the walls of the gall are

hypertrophic and chambered. The parasitic wasps develop themselves in these

oval gall-chambers which are 2 mm long and 1.5 mm wide. They protrude inward

in the gall-cavity which explains its reduced size and its irregularity.

Idem. Gall 23 Il. Wide-spread in B.T.N.R.

By means of cages, wasps and midges were obtained. Fide D.H. Murphy

(personal communication), wasps are the gall-makers, and midges are inquilines

(fig. 5, d, e; Pl. 4, e). Galls 23 II protrude onto the adaxial leaf-surface, mainly

alongside the midrib and sideveins where they form continuous, non-coalescent

alignments. At that level the lamina is deformed and has a tendancy to roll up,

thus partially covering them (PI. 4, e). A fully developed gall is column-shaped,

2 mm high and less than 1 mm wide, reddish brown, covered with simple hairs.

The gall can be divided into three superimposed parts: a pedicel, a median swollen

zone and an apical zone (fig. 5, d). The pedicel is roughly cylindrical and

perpendicular to the lamina. The gall swelling corresponds to the rounded larval

chamber which appears lateral in position relative to the median gall-axis. The

tapering apical zone is more or less curved. The position of the cavity and the

apex of the gall distinguish morphologically a longitudinal plane of symmetry,

perpendicular to the lamina. An anatomical study confirms this. The gall is

covered with a piliferous epidermis. A longisection of the gall, perpendicular to

the lamina shows that the epidermic gall-cells are longitudinally hypertrophic being

3.3 times longer than normal epidermic cells, and transversally only slightly higher.

Under the gall epidermis one finds a cortical tanniferous parenchyma which reaches

its maximum thickness in the apical gall part. Some or all cells of the outer cortical

layers are differentiated into sclerenchymatous cells forming more or less a con-

tinuous shell. The gall axis consists of a tanniferous medullar parenchyma.

Epidermis, cortical and medullar parenchyma are the only three tissues that

constitute the apical gall-part. The cells of those tissues are much elongated in

the direction of the longitudinal gall-axis. Level with the swollen part one finds

the larval cavity surrounded by a spherical mass of nutritive parenchyma the cells

of which are polygonal and characterised by a nuclear and nucleolar hypertrophy.

32 Gardens’ Bulletin, Singapore — XXVII (1974)

A section cut along the plane of symmetry shows that the nutritive tissue juts

out over the gall pedicel and that its protruding part is covered uniquely by epider-

mic and shell layers; on the internal side the medullar parenchyma runs along the

nutritive tissue. A vascular ring of tracheids and associated parenchyma runs the

length of the pedicel in between cortical and medullar parenchyma. That vascular

tissue in relation with the leaf-veins at the base reaches the nutritive tissue: the

vascular bundles that come right up to the nutritive tissue spread and surround it

on both sides of the gall plane of symmetry. Those bundles which are diverted

from the nutritive tissue joins the latter by running obliquely round the medullar

parenchyma. On maturity there is necrosis of the medullar parenchyma and forma-

tion of a dehiscing canal in the centre of the gall-pedicel. This canal opens through

an abaxial pore.

Idem. Gall 23 TT & 77, B,TN.R. (fig..d, 2..D.¢5 FL 4 2, 9),

By means of cages, Diptera and Hymenoptera have been obtained. Diptera:

this species belongs to the subfamily Cecidomyiinae, supertribe Asphondyliidi. The

genus seems to be underscribed fide Dr. Nijveldt in litt. Hymenoptera: Chalcidae

of the family Eulophidae, genus near Pediobius fide Kerrich in litt. The midges

are the gall-makers, the wasps are parasites. At the height of its development, the

gall reaches 2 mm long by 1.5 mm wide. It is mushroom-shaped: pedicel topped

by a cap. It stands more often on the abaxial surface of the leaf. The pedicel

inserted on a small leaf-vein between the side-veins stands out adaxially as a smail

bright red protuberance; it is red at its base as well, becoming lighter and turning

yellow towards the top. The cap is green-yellow. The gall turns brown and dries

out when mature. The young gall is green-yellow, the red pigmentation does not

appear until later. The gall-chamber containing one midge-larva is situated under

the cap at the apex of the pedicel. This chamber remains open by a small

ostiolar canal surrounded by hairs and located in the centre of the cap (fig. 5, c).

The gall-cavity is surrounded by a sclerenchymatous shell made of an upper part

and a lower part. The latter is covered by a thin nutritive epidermis. The epidermis

of the gall is glabrous, level with the pedicel and hairy, level with the cap: on the

external cap-side (e) these hairs are tufted and relatively short; on the internal

side (i) they are very long and form a sleeve surrounding the apex of the pedicel.

Under the epidermis lies a hypertrophic and tanniferous cortical parenchyma. The

centre of the gall is occupied by a medullar parenchyma the cells of which are

elongated vertically in the pedicel and radiate round the gall cavity, level with the

cap. In the internal part of the parenchyma one finds about 24 vascular bundles

disposed in a circle which run up the pedicel parallel to its axis. These bundles

enter the cap and reach the ostiolar canal as shown on fig. 5, c. Ontogenically, the

hairy cap forms first, the pedicel develops subsequently. The gall cavity, very

small at the beginning, becomes enlarged due to the development of nutritive pillars

that disjoint the lower and upper parts of the shell. After pupating the midge

makes its way through the wall of the cap, leaving its pupal integument partly

inserted in the gall. When the gall is attacked by parasitic wasps, its apical part is

deformed by a lateral swelling, and the cap of the cecidium is rejected on the side

(fig. 5, a).

Idem. Gall 23;:1Va...B.T.N.Ri (PL. 3.. a). Bsc):

Cause: Coccidae, genus Beesonia, undescribed species, fide Beardsley in litt.

These coccids are very unusual according to this specialist. They provoke a fusiform

hypertrophy affecting the stem on a variable length, sometimes for more than

Cecidia of Singapore 33

10 cm. The placement of the coccids is marked by a small depression, in the

centre of which with the binocular magnifying glass one notices the brown cuticle

of the posterior part of the scale mother. When mature the latter has a distended

body in the shape of a flat bag, reaching 1 cm long by 8 mm wide, lying between

the central cylinder and the cortical parenchyma. Dr. Beardsley comments as

follows: “‘the posterior end of the bag forms a constricted neck which extends

through the bark of the twig, and the young coccids probably leave the mother’s

body and the gall by way of an opening at the tip of this constricted portion’.

Under the influence of the parasitic coccids, the plant tissues necrotize, become

brown, and the whole twig on which the gall occurs dries out (PI. 3, a). The gall

is often attacked by parasitic wasps that make their ovoid larval cavities in the

hypertrophic cortical gall-tissue. They leave the gall by piercing the gall-tissues

(Pra: D:-c).

Idem. Gall 23 [Vb, 77, 75, 106. B.T.N.R. (Pl. 3, d, e).

Wasp-gall. The attacked part of the stem becomes hypertrophic, fusiform and

covered with hemispheric swellings of approximatively 3 mm in diam. Each of

these growths contains several wasp-larvae, lodged in a rounded gall-cavity. The

exit of the gall-makers is central and at the distal extremity of the vertical axis.

Idem. Gall 23 V. B.T.N.R., between Jalan Kutu and Main Road. (PI. 4, a, b, c, d).

Gallacoccus anthonyae Beardsley. Same genus as the gall forming coccid 23 [.

(for reference see foot-note on p. 23). The gall, abaxially attached on the basal

half of the midrib or the petiole, has the form of a bud, constitutes overlapping

foliaceous appendages, the base of which are on two generating lines (distichous

disposition). It can reach 6 cm long and 2 cm at the widest part. Light-green and

whitish when young, yellowing at maturity, it turns brown and dries out like the

leaf that carries it. The young coccids are found in a translucid bag. They are

surrounded by their waxy whitish secretion and hold themselves in the axil of the

seventh leaf counting from the base.

Idem. Gall 23 VI. B.T.N.R.. Ginger Walk.

Insect-gall. Ball-shaped, 1 to 2 mm in diam., yellow, becoming red-brown at

maturity. The cecidia located on the veins are visible only on that face of the

lamina on which they find themselves. They detach themselves easily.

Idem. Gall 23 VI. B.T.N.R., Ginger Walk.

Wasp-gall. Disc-shaped gall, less than 1 mm in thickness, 4 mm in diam.,

affecting symmetrically both faces of the lamina. The exit of the gall-makers is

abaxial.

Idem. Gall 23 VIII. B.T.N.R., Ginger Walk.

Insect-gall. Adaxially, it is a conical protuberance 1 mm in height, on a base

1 mm in diam., to which corresponds abaxially a similar protuberance. The galls

are located along the length of the side-veins, and on the side of the petiole. When

they are close together, they fuse at the base, forming alignments that look like

cockscombs.

Shorea sp. Gall 168 I. B.T.N.R., Jungle Valley.

Midge-gall. Bright-yellow spheroid gall 3 or 4 mm in diam., attached by a

short pedicel to the under-side of the leaf. The point of attachment is marked on

34 Gardens’ Bulletin, Singapore -— XXVII (1974)

the upper-surface of the leaf by a little protuberance surrounded by a depigmented

area. This gall resembles the gall 21437 no. 1007 on Shorea belangeran Burck.

(Bib. 22, p. 391).

EUPHORBIACEAE.

Acalypha wilkesiana Muell. Arg. Gall 161. Coronation Road (garden) Singapore,

and Botanic Gardens of Tjibodas, Java.

Cause not determined. It appears as a globulous mass of witches’ broom about

6 cm in diam., sprouting from the twigs. This gall is made up of a tangle of filaments

radicular in nature, and of twigs carrying reduced leaves.

Aporosa benthamiana Hook. f. and Thom. Gall 98 I. B.T.N.R., Jalan Kutu. (fig.

6..Pie kL), db).

Thrips-gall. The leaf becomes involute; the leaf-tip is not involved in the

cecidiogenous morphogenesis. Abaxially the lamina is punctuated between the

side-veins by little pustules 1 mm in diam., with a black point in their centre.

Idem. Gall 98 III. B.T.N.R. Jalan Kutu. (Pl. 11, b, c).

Insect-gall. Petiole or midrib develops roundish or fusiform swellings, 4 to 10

mm long and about 4 mm wide, visible on both faces of the lamina. The ovoid

gall-cavity is located in the hypertrophic medullar pith. Exit of the gall-maker is

by a large pore.

Idem. Gall 98 IV. B.T.N.R., Jalan Kutu. (Pl. 11, c).

Wasp-gall. Yellowish fleshy mamillated swelling affecting laterally the stem

along a few cm. A longisection of the gall shows that the cortical parenchyma is

the only stem-tissue to be hyper-developed under the influence of the numerous

wasp-larvae that it contains; pith and vascular bundles are not modified by the

gall-makers. That explains why the stem-growth is not affected by the gall. On

maturity the wasps leave the gall through a large pore.

Baccaurea ? griffithii. Gall 130. B.T.N.R., Ginger Walk.

Cause undetermined. Globular or more or less elongated deformation of the

stem or of the petiole for about 1 cm.

Coelodepas glanduligerum Pax. Hoffm. Gall 71. B.T.N.R., Jungle Valley track.

PPL 11 oa).

Midge-gall. Abaxial disc-shaped gall, 5 mm in diam., by 1 mm in height,

depressed in its centre. The rim formed in this way is irregular and has a

brownish-red tint. The central depression is cylindrical, 1 mm in diam., 0.5 mm

in height. In it a fine epidermic pellicle forms the roof of the gall-cavity where one

midge-larva is found. At maturity the parasite leaves the gall through a semi-

circular orifice formed by piercing the membrane. On the adaxial side, the gall

presents itself as a yellowish rounded spot which has a very slight central brownish

point.

Macaranga triloba Muell. Arg. Gall 124. B.T.N.R., North View Path.

Midge-gall. Ovoid, attached to the abaxial face of the lamina, 4 mm long by

! mm wide; its surface is green and covered with long white hairs. Its attachment

is marked on the adaxial face of the lamina by a slight protuberance covered

with white hairs. The exit of the gall-maker is adaxial.

Cecidia of Singapore 35

Distribution: Singapore — Java, Sumatra, Malacca (Bib. 10, p. 80, no. 232.—12,

p. 32, no. 304, fig. 131-14, no. 31-22, p. 313, no. 771.28, p. 455, no. 1675 &

1676, fig. 983).

Mallotus penangensis Muell. Arg. Gall 111. B.T.N.R., North View Path.

Psyllid-gall? All the galls found were already open and devoid of gall-makers.

Adaxially this leaf-gall is in shape of a hemispheric hollow dome 6 mm in diam.,

covered with long hairs. Abaxially the walls of the gall are divided into several

reflexed lobes.

FAGACEAE

Castanopsis wallichii King Gall 25. B.T.N.R., on the corner of Main Road and

Jalan Kutu. (PI. 6, d, e).

Midge-gall. Gali of the stem and the leaves affecting the axillary and terminal

buds, the petiole and the veins. The parasitic midges provoke hypertrophy and

inhibit growth of the attacked organ.

— Petiole: fusiform swelling, 3 mm wide.

— Veins: the gall, especially visible abaxially, presents itself as a fusiform

swelling 3 mm in width on the midrib, and 1 mm in width on the side-veins.

Moreover, the lamina is deformed and curved around the attacked zone, caused

by inhibition in growth.

— Terminal bud: hypertrophy of its axis transforms the bud into an irregular

sphere 8 or 9 mm in diam. From this sphere contorted leaf-shoots project. Inside

are many midge-larvae.

Lithocarpus conocarpus (Oudem.) Rehd. Gall 120 I. B.T.N.R., between Main Road

and Jalan Kutu. (fig. 7, c, d; Pl. 6, c).

Wasp-gall (probably a cynipid). Brown ovoid cylindrical gall protruding more

or less equally on both faces of the lamina, reaching 2 mm in total height and 1

mm in width. The apex of the gall on the adaxial face is equiped with a flat lid

that has a small central erect prickle. The gall is sessile, detaches itself before

maturity from the orange wasp-larva contained in its large cavity and leaves an

alveole in the lamina (PI. 6, c). This gall has a rapid development from January

to March (see p. 20).

Idem. Gall 135 IJ. B.T.N.R., between Ginger Walk and Jalan Kutu. (fig. 7, b).

Moth-gall. Fusiform swelling of the midrib, 1.5 cm long, 0.6 cm wide at its

centre, visible on both faces of the lamina, and is of woody consistence. The

interior is a long gall-chamber.

fore, Gall 135011. B.T:N-R. (fig. 7, a; Pl. 6, a, 0).

Moth-gall developed on stem and petiole.

— Stem: fusiform swelling of the stem. One moth-larva makes a central long

gall-chamber. It induces the fragmentation of the cambium into four parts; after

departure of the gall-maker, this divided cambium keeps on functioning and

leads to the formation of four stem units (pl. 6, b). Due to the proliferation of

these internal stem-tissues, the gall-cavity disappears and the bark is fragmented

into four longitudinal strips. At the end of its development this gall reaches 2.5

cm in length and 8 mm in width (PI. 6, a: fig. 7, a).

36 Gardens’ Bulletin, Singapore — X XVII (1974)

— Petiole: level with the insertion of the petiole on the stem, adaxial side, the

internal hypertrophic tissues become a small irregular growth about 3 mm diam.

(fig. 7, a).

Lithocarpus sundaicus (Bl.) Rehd. Gall 153 1V. B.T.N.R., very close to the summit.

(fig. 8; 9). |

Gall caused by a coccid that belongs to the family Diaspididae fide Beardsley

in litt. The young Homoptera are localised at the axil of young leaves of lateral

buds (fig. 8, a). They create:

— Shortening of the internodes; the leaves are tufted in a rosette (fig. 8, d). Their

bases become hypertrophic (fig. 8, c, d).

— Cessation of growth of the principal axis.

— Formation of secondary and tertiary shoots by axillary buds (fig. 8, a, b). The

young leaves are modified in rhabdodes.

One notices the development of vigorous shoots under the gall-zone which

appear to be the strongest shoots in the plant (fig. 9, d); this poses the problem

of causal determination. According to Dr. Cusset in litt. it might only be due to

an inherent phenomenon in the considered species if the normal growth of the

plant is either monopodial with flush (fig. 9, a) or sympodial and more or less

acrotonic (fig. 9, b). But if the normal growth of the plant is monopodial and

without rhythm (fig. 9, c), it might be a phenomenon linked to the remote action

of the cecidiogenous agent. One could then take in consideration the possible

interaction between the distal zone with axillary buds abundantly developed (BA),

and the subjacent zone with axillary buds strongly developed (BS), fig. 8, d.

GUTTIFERAE

Calophyllum ferrugineum Ridl. Gall 68 I, 101, 136, 141. Widespread in B.T.N.R.

ig. 13.d5. 61.5: 6)

Midge-gall. This species belongs to the subfamily Cecidomyiinae, supertribe

Cecidomyiidi, but the genus seems to be undescribed fide Nijveldt in litt. Parasites

or inquilines: thrips and wasps. The latter are chalcids of the family Eulophidae,

belonging to the genus Tetrastichus fide Kerrich in litt. The gall is formed by the

edges of the lamina which fold downwards thus resulting in a cylinder 1.5 mm. in

diam. One can, parallel to the midrib, distinguish three parts in each half lamina,

the behaviour of which is different in the morphogenetic movement: proximal,

median and distal zones, with respect to the midrib. The median zone only of

each half lamina participates in the formation of the wall of the cylinder. This

tegion of the lamina is considerably thickened and its surface is irregular. The

proximal and distal zones are clung together, lower epidermis against lower

epidermis (fig. 13, d). In the long cavity thus delimited, one finds many midge-

larvae which at maturity protrude from the channel of the cylinder by sliding

between the distal and proximal zones, where they leave their empty pupal

integument. However the parastic wasps work their way out by piercing the wall

of the cylinder.

Idem. Gall 68 II, 101, 136, 141. Rather wide-spread in B.T.N.R. (Pl. 5 f).

Midge-gall, affecting the stem, petiole and midrib. On the midrib the isolated

gall looks like a hemispheroid swelling of 5 mm in maximum diam., more often

abaxial, visible only one one side of the leaf. Frequently the swellings are fused

=* i -

Cecidia of Singapore 37

and the whole vein looks hypertrophic. The attacked petiole is considerably larger

(5S mm versus 2 mm). There are hemispheroid swellings laterally disposed on the

stem.

Idem. Gall 141 III. Very rare in B.T.N.R., localised between Main Road and

Jalan Kutu. (fig. 11, a).

Moth-gall. (See also description of gall 107 IV on Calophyllum pulcherrimum

Wall. on p. 38). Swelling of both faces of the lamina, elongating parallel to

the midrib, 25 mm in length, 8 mm in maximum width, and a few mm in thickness.

The larvae eat the interior of the leaf except the epidermis. However it is a gall

and not a mine: there is formation of a very definite growth due no doubt to a

parenchymatous nutritive proliferation. The gall i.e. 141 IIT looks like that described

on Calophyllum inophyllum L. (Bib. 22, p. 386, no. 989, Ist, fig. 705).

Calophyllum inophylloide King var. singaporense Henderson & Wyatt-Smith. Gall

43 I. MacRitchie Reservoir. (fig. 12, b).

Gall caused by a coccid: Amorphococcus sp. (Lecanodiaspididae), fide Dr.

Beardsley in litt. The gall develops abaxially on the midrib. Each gall is made up

in the same way as bivalve molluscs, of two shells contacting one another by the

edge, limiting between them a large rounded gall-cavity containing the young

coccids that form an orange mass surrounded by the product of their whitish waxy.

secretion. When the coccids settle on a young leaf they inhibit the development of

the lamina and the internodes are shortened: the apex of the stem is crowned by

a globular bunch of more or less spherical galls each 1 cm in diameter.

Idem. Gall 43 Il. MacRitchie Reservoir. (fig. 12, a).

Psyllid-gal!l. The leaf becomes revolute. Each leaf-half can be affected

separately, forming a cylinder, 1 mm in diam. Mr. D. H. Murphy notes that leaf

roll galls of this type on C. inophyllum are well known and are caused by

Leptynoptera sulphurea.

Calophyllum pulcherrimum Wall. Gall 107 I. Wide-spread in B.T.N.R. (fig. 10, a).

Psyllid-gall. The leaf is conduplicate and tends to become revolute. Its edges

are undulated and crisped. The midrip is curved in a hook towards the abaxial

leaf-side. The petiole is sometimes twisted, bringing the abaxial side of the leaf

in adaxial position.

Idem. Gall 107 I. B.T.N.R. (fig. 10, c,d).

Midge-gall. The midge belongs to the genus Bruggmanniella but the species

seems to be undescribed fide Nijveldt in litt. Fleshy spheroid gall 3 mm in diameter.

pale-green or light-yellow, more often abaxially attached to the midrib by a little

pedicel. In aging, it becomes yellow, orange, then brown and dries out. Inside the

gall-cavity, one midge-larva is found. The gall is often attacked by parasitic wasps:

its consistence is then firmer, and its color changes to dark green, marbled with

brown. This gall i.e. 107 II looks like that described on Calophyllum javanicum

Mig. (Bib. 22, p. 386, no 991, Ist, fig. 760).

Idem. Gall 107 III. Quite rare in B.T.N.R. (fig. 10, b).

Midge-gall. Swelling of the stem, petiole and midrib. The stem-gall is spheroid,

about 1 cm across; it is due to a considerable hypertrophy of the central cylinder.

38 Gardens’ Bulletin, Singapore -— XXVII (1974)

and contains several cortical gall-chambers. When the stem is strongly attacked,

the galls are more or less fused, linked by hypertrophic stem-tissue, i.e. without

larval cavity.

Idem, Gall 167 IV and 107 IV. Rare in B.T.N.R. (fig. 11, b).

Moth-gall. See also description of gall 141 III on C. ferrugineum Ridl. on

p. 37. Swelling visible on each face of the lamina; it is 4 cm long, 0.4 cm

wide, 0.2 cm in height abaxially and about 1 mm in height adaxially. It is elongated

parallel to midrib, in form of a “S’, of which the basal extremity (petiole side),

pushes away from the vein, while the distal extremity converges towards it. The

adaxial dehiscence is caused by the wall of the gall collapsing inward.

LAURACEAE.

Actinodaphne sp. Gall 24 I, 67, 73, 93, 99, 138, 143. Wide-spread in B.T.N.R.

CPL. 75.8).

Gall caused by a mite “Eriophyes sp., very similar to Nalepa’s gyrographus

(on Litsea) and also close to Corti’s Linderae, but the host species is different and

Corti’s mite makes conical galls.” fide Dr. Keifer in litt. Pockets 1 cm high which

appear like blisters on the adaxial leaf-surface. They are abaxially covered with

a whitish erineum becoming brown on maturity.

Idem. Gall 24 II, 67, 73, 93, 99, 138, 143. Widespread in B.T.N.R..

alongside Jalan Kutu; the trees are sometimes entirely covered with galls. (fig. 13.

be P75):

The gall-maker is a midge belonging to the subfamily Cecidomyiinae, super-

tribe Cecidomyiidi, fide Nijveldt in litt. Gall of the leaf, stem and petiole, On the

stem it is a lateral spheroid outgrowth, attached by a large base. On the leaf (case

of an isolated gall), it is a sphere 5 mm in diam., having the lamina as diametrical

plane. The glabrous green galls are located principally on the midrib; they are

very often fused, and the whole leaf can be affected. The dehiscence is abaxial

(fig. 13, b). In the centre of the abaxial hemisphere one finds a small depression

that eventually becomes the exit pore for the gall-maker. There is a cellular

degeneration along a central axis, more or less perpendicular to the lamina. Often

the galls are attacked by parastic wasps belonging to the superfamily Proctotrupoi-

dea, family Scelionidae, Platygaster sp. fide Nixon in litt. Then the galls are

dark green, marbled with brown.

Idem. Gall 99 III. B.T.N.R., at the extremity of Jalan Kutu, around

point 'C on ‘fig 1)¢."@ 1. 7,’ e%d).

Midge-gall. Spheroid acrocecidium reaching 3 cm in diam. with a greenish

surface, marbled with brown spots more or less suberized. It contains a number

of gall-cavities distributed in a median horizontal plane. The young galls carry

modified leaves in the shape of spines, with lamina absent or very reduced. These

appendages are adaxially plane, and abaxially rounded: there is therefore bilateri-

sation. The exact nature of these foliar appendages poses a morphological problem.

One can admit that, under the influence of gall-makers, there is an inhibition of

the foliarisation phenomenon depending on the stem-apex, and formation of

thabdodes. For experiments of microsurgery of the apex see Wardlaw, Bib. 51.

Cecidia of Singapore 39

Idem. Gall 143 IV. B.T.N.R., at the corner of Main Road and Jalan

Kutu, near a thinning of the forest. This gall is not widespread. (fig. 13, a).

Midge-gall. Spherical cecidium of 5 mm in diameter, more often attached

abaxially to the midrib by a large base, and visible only on the side on which it

finds itself. It is fleshy, chlorophyllous, covered with a brown suberized layer. The

interior is a large rounded gall-chamber containing one midge-larva. The animal

pierces the wall at maturity.

Cryptocarya sp. Gall 128. B.T.N.R.

Cause undetermined, but the presence of numerous ants at the interior of the

tolled leaf suggests that it might be caused by Homoptera. Edges of lamina

revolute. The leaf curves downwards forming a hook.

Endiandra sp. Gall 156 I. B.T.N.R., Main Road. (fig. 14).

Thrips-gall. Edges of lamina involute and crisp. The leaf is contorted, swollen,

depigmented, yellow, marbled with brown.

Idem. Gall 156 Ill. B.T.N.R., Main Road near the summit. (fig. 14, b).

Cause undetermined. Cylindrical conical adaxial expansion on the midrib, 3

mm in height and less than 1 mm in diam. at the base. This gall is not very

widespread.

Litsea elliptica (Bl.) Boerl. Gall 86. MacRitchie Reservoir. (Pl. 8, f).

Gall caused by a mite: Eriophyes gyrographus Nal. fide Keifer in litt.

Pockets varying in size and height from a few mm to a few cm pouch out from

the adaxial leaf-surface. They are covered abaxially with a bright erineum which

browns on maturity. It is formed by unicellular clavate hairs.

Litsea grandis Hook. f. Gall 144. B.T.N.R.

Homoptera-gall. Spots, visible on both sides of the lamina; roundish, | cm

in diam. approximately, often confluent and forming large depigmented patches.

The gall-makers, fixed abaxially to the centre of the depigmented area, induce a

very slight elevation of the lamina. They are surrounded by their whitish waxy

secretion and are covered with an hemispheric shield (skin of the first larval instar)

which lifts at maturity like a lid.

Nothaphoebe kingiana var. malvescens Gamble. Gall 92 I, 79, 94, 133. Wide-spread

mm B-T.N.R. (Pl. 8, d).

Psyllid-gall. The young psyllids fixed abaxially create adaxial laminal eleva-

tions 2 mm in diam. at base, and 1 mm in height.

Idem. Gall 94 II, 79, 92, 133. B.T.N.R. (fig. 15, d, e).

Midge-gall, visible on both faces of the lamina, forming a circular swelling

3 mm in diam. approximately, slightly thicker than the lamina, affecting especially

the adaxial face. The midge-larva fills completely the small central gall-cavity. On

the adaxial face the gall is dark green, almost black and lighter in its centre. At

dehiscence the centre becomes white-grey adaxially and red-brown abaxially. The

gall opens adaxially by a central ostiole.

40 Gardens’ Bulletin, Singapore —- X XVII (1974)

Idem. Gall 133 Il, 79, 94. Wide-spread in B.T.N.R. (fig. 15, b, c,; Pl. 8, e).

Mite-gall. Pouch-gall affecting both faces of the lamina, generally isolated,

sometimes two or three are fused. Abaxially, the isolated gall is in the shape of a

mushroom, 1.5 mm high, the cap is 2.5 mm in diameter. To the insertion point of

the gall corresponds adaxially a depression 0.5 mm in depth, and 2 mm in diameter.

The gall is formed abaxially by development of lips enclosing a space in which

the parasites are lodged. This gall-cavity is crowed with fleshy parietal projections

directed inward. The gall stays open by an ostiole situated in the centre of the

slight apical depression of the cap: this orifice is surrounded by hairs.

LEGUMINOSEAE. ae

Dalbergia rostrata Grah. Gall 134, 162. Wide-spread in B.T.N.R. (Pl. 9, d, e).

Midge-gall. Leaflets conduplicate. The two leaflet-halves thicken and are more

or less fused. The place of each gall-cavity is marked by peg-like pustules on the

abaxial surface. The leaflets can be affected partially or entirely. In the latter case,

they take the appearance of a pod.

Dialium maingayi Baker Gall 147. B.T.N.R., between Jalan Kutu and Main Road.

Thrips-gall. Leaflets revolute and retrorse. The distal extremity of the leaflet

is not involved in the cecidiogenous morphogenesis. The gall surface is yellow-

green, spotted with red.

Pithecellobium clypearia Benth. Gall 115 I. Quite wide-spread in B.T.N.R. (fig.

1G) be: BISONS bc).

Gall caused by a midge belonging to the supertribe Asphondyliidi fide

Nijveldt in litt. Gall occurs on the main leaf-stalk, on the midrib and veins of the

leaflets. On the lamina, the galls on the veins are isolated, ovoid, 2.5 mm in length

and 1.5 mm in maximum width. They are attached by a large base to the main

vein, abaxially and sometimes adaxially. At the interior one finds a very large

larval chamber. The circular emergence-hole is made on the gall-top. On the

main leaf-stalk, the galls are less definite since they are often fused. The main

leaf-stalk can be infected, starting from its distal extremity along a varied length:

it is then hypertrophic and contorted.

Idem Gall 115 IJ. Very rare in B.T.N.R., located between Jalan Kutu and Main

Road. (fig. 16, a). |

Homoptera-gall? The leaflets tend to be conduplicate. Their lamina is entirely

undulate. The distal end of the leaflets becomes retrorse.

LOGANIACEAE.

Fagraea fragrans Roxb. Gall 45, 87. MacRitchie Rerservoir. (Pl. 10, e).

Thrips-gall. The affected leaf becomes revolute. The surface of the gall is more

or less granulate. .

Strychnos sp. Gall 27, 74. B.T.N.R. (Pl. 11, e).

Gall caused by a midge the species of which belongs to the genus Bruggman-

niella and seems to be undescribed fide Nijveldt in litt. Brownish or ‘greyish

fusiform swelling, 5 to 10 mm long, laterally affecting the stem, petiole or midrib.

The galls are sometimes fused. At the interior one finds a larval cavity elongated

in the direction of the axis of the organ.

Cecidia of Singapore: 4]

MENISPERMACEAE.

Hypserpa cuspidata (Wall.) Miers Gall 123, 108. Wide-spread in B.T.N.R. (PI.

13, a).

Coccid-gall; the gall-maker “appears to belong to the family Eriococcidae, ...

It probably represents an undescribed genus and species, . . . It is not closely

related to the new Eriococcids from Shorea pauciflora King.’* fide Beardsley

in litt. Gall of the stem and leaf (lamina, petiole, veins). Under the effect of

the gall-makers, there is an inhibition of growth and hypertrophy occurs. The

young coccids are contained in large numbers in fine-walled bags (0.1 mm in

diam. These bags fill completely the gall-cavity that opens out by a circular ostiole.

They fit in the cavity and the short exit canal so that they have the form of a

bottle with a large body and a very short neck. On the adaxial face of the lamina,

the placement of the coccids is made visible by the local depigmentation which

they cause: the leaf is punctuated by little yellow spots 0.5 mm across, with

central ostioles, where a tuft of white filaments (waxy secretion. of the. coccids)

comes. out. During the development of the lamina-gall, one notices a slight

augmentation in thickness of the lamina, located at the placement of the gall-

makers, then necrosis and darkening of. the gall-tissues. The coccids installed in

the veins inhibit their growth and create a distortion of the leaves. The parasites

localised in the stem create a general hypertrophy of this part which when thus

affected can reach 8 mm across, and cause some inhibitions of growth: shortening

of the internodes, contortion of the stem, inhibition of leaf-development.

MORACEAE.

Ficus pumila Linn. Gall 48, 150. B.T.N.R., Jalan Kutu and Main Road. (fig. 18).

Gall caused by a wasp belonging to the superfamily Chalcidoidea. Gall of

terminal or axillary bud. This cecidium is fleshy, globulous, | cm in diam.

approximately, often slightly elongated in the direction of the axis. It is light

yellow when juvenile, becoming brown and drying out on maturity. At the interior

of a young gall one finds three or four larval cavities, each containing a wasp-

larva. Those chambers become united into one large irregular cavity as the larvae

feed on the nutritive gall-parenchyma. The gall is situated in the leaf axil or at

the apex of the stem: it is a bud gall. In fact the gall is largely constituted of the

petiole of the first leaf fused with the axis of the bud, both enlarged. The lamina

of the first leaf does not participate in the formation of the gall: it emerges from

the globulous mass and is sessile or subsessile. The fusion between the lower part

of the bud-axis and the petiole of the first leaf is proved by the presence of the

stipules of the latter on the gall, and by the presence of the apex of the bud at

the apex of the gall. There is a resemblance between the gall ic. 48 and 150 and

the wasp-gall on Ficus recurva Bl. of other authors (Bib. 22, p. 136, no 20145, fig.

172-28, pp. 186-187, no. 679-680, fig. 385).

Ficus retusa L. Gall 33. Mangrove, West Coast Road.

Thrips-gall. Leaf conduplicate, with a brown surface, marked with red. It is

one of the first thrips-galls that has been described and studied. Distribution: this

gall is spread throughout tropical Asia (Singapore — Siam, Cochinchina, Hongkong,

Malacca, Java, Sumatra, Celebes), and also in Africa. (Bib. 1, p. 250.—S, p. 154.-15,

p. 30.-17, p. 54, no 18-19, p. 272-21, p. 128, no 31.—22, p. 140, no 269, 8th, fig.

14, Pi. I1V.—25, pp. 56-58, no 108, fig. 103—108.—26, p. 95.27, pp. 248—250.-28, p.

188, no 685, fig. 372—377.-30, pp. 131, 150, 152, 160-31, pp. 302, 309, 314-37,

pp. 251-253.-46 pp. 423-424.-52, p. 117, Pl. XVI, fig. 16-18.).

*See footnote p FP:

42 Gardens’ Bulletin, Singapore — XXVIII (1974)

Ficus sinuata Thunb. ssp. sinuata Gall 146. Tyersal Road and 44 Nassim Road.

(Pl. 8, a, b, ¢).

Gall caused by a mite, “Eriophyes sp. which does not closely resemble any

of Nalepa’s Ficus spp. either as to mite description or gall . . .” fide Keifer in

litt. Irregular adaxial outgrowth, nodulous or more or less conical, 5 mm in

maximum height. To this corresponds a conical protuberance abaxially, 6 mm

in maximum height, the apical extremity is perforated by an orifice leading to the

gall-cavity. This cavity is crowded by parietal out-growths.

MYRISTICACEAE.

Knema communis J. Sinclair Gall 129 I. B.T.N.R., between Jalan Kutu and Main

Road. (Pl. 13, d).

Insect-gall. Symmetrical gall in relation to the leaf surface. On the adaxial

and abaxial faces, the gall presents itself as a conical growth 1 mm in height and

2 mm in diam. at the base. At dehiscence the outer wall of the gall divides into

brown woody lobes which become reflexed, leaving a central erect column. This

column forms a stopper that detaches itself at maturity, allowing therefore the exit

of the gall-makers.

Idem. Gall 129 II. B.T.N.R., between Jalan Kutu and Main Road.

Thrips-gall? The edges of the leaf tend to become involute. Between the side-

veins, the lamina is more or less crisped.

Gymnacranthera eugeniifolia (A. DC.) Sinclair var. eugeniifolia Gall 118. B.T.N.R.,

Jalan Kutu.

Cause non determined. Local inhibition of growth of the stem and the midrib,

provoking contortion. Moreover the edges of the affected leaves tend to become

revolute. The lamina is contorted and crisped.

MYRTACEAE.

General comments concerning the galls on genus Eugenia: the galls found in

Singapore on the genus are of classic types, already described by many authors

on different species of Eugenia. One can distinguish four types of galls:

(1) Stem-galls: spherical or fusiform swellings formed by hypertrophy and hyper-

plasy, caused by insects.

(2) Leaf-galls:

(a) Edges of the leaf involute or revolute, caused by thrips.

(b) Adaxially hemispheric, abaxially conical equipped with an ostiole, caused by

psyllids.

Eugenia cumingiana Vidal Gall 155 I. B.T.N.R.

Wasp-gall. Fusiform swelling of the stem, containing a number of gall-cavities.

Idem. Gall 139 If. B.T.N.R.

Psyllid-gall. Conical depression of the lamina less than 1 mm in depth from

base less than 1 mm in diameter.

Cecidia of Singapore 43

Idem. Gall 157 III. B.T.N.R., Jungle Valley.

Thrips-gall? The edges of the leaf tend to become involute.

Eugenia malaccensis Linn. Gall 81. Singapore Botanic Gardens.

Psyllid-gall caused without doubt by Megatrioza vitiensis, the biology of which

was studied by Zehnter but not consulted. Adaxially, it is a red hemisphere 5 to

8 mm in diam. containing a large gall-cavity. Abaxially, the young gall has the

form of a conical hollow swelling 5 mm in height, the wall of which splits and

curls at maturity. This gall is very wide-spread: Singapore-Java, Ceylon, Singa-

pore, Malacca, Amboina, Sumatra, Salajar Island and Fiji Island (Bib. 3, p. 266.4,

p. 195.-5, p. 172, fig. 175-12, p. 6, no 37-13, p. 6, no 8-14, p. 6, no 10-15, p.

29, no 19-21, p. 126, no 26-22, p. 411, Ist, no 1074-28, p. 594, No 2142, fig.

1235 & 1236.33, p. 103, fig. 3, 4,549, p. 150.—52, p. 3, fig. 1 to 5.).

Eugenia rugosa (Korth.) Merr. Gall 158 I. B.T.N.R., Jungle Valley track. (fig. 17,

mek:

Psyllid-gall. Adaxially, hemispherical or conical protuberance, reaching 3 mm

in height on a base 6 mm in diam. Abaxially, conical outgrowth 4 mm high,

Opening at the apex by a large circular ostiole. Galls are often fused. of a lighter

green than the leaves, turning yellow or orange on maturity.

Idem. Gall 158 Il. B.T.N.R., Jungle Valley track. (fig. 17, a. b).

Insect-gall. Spherical swelling of the stem, 1.5 cm in diam. All the tissues

seem to participate in the cecidiogenous morphogenesis, in particular the vascular

tissue, xylem and phloem, and the cortical parenchyma. The gall-cavity is irre-

gularly surrounded by a red-brown parenchymatous nutritive tissue; at maturity

the gall-makers pierce a cylindrical canal through the gall-tissues leading to a

circular orifice.

Idem. Gall 158 I. B.T.N.R., Jungle Valley track.

Psyllid-gall. Conical depression of the lamina approximately 1 mm deep,

caused by the psyllids fixed adaxially.

Idem. Gall 143 1V. B.T.N.R., Ginger Walk. (fig. 17, d).

Wasp-gall? Irregular fusiform swelling of the twig, containing a number of

gall-cavities. The bark pushed by the internal tissues breaks into flakes.

Eugenia subdecusscata Duthie Gall 148 I. B.T.N.R.

Psyllid-gall. Conical elevation of the lamina of | mm in height on a base 2

mm in diam.

Idem. Gall 166 I. B.T.N.R. (Pl. 10, d).

Thrips-gall? The leaf is involute. The surface of the gall is more or less

marked with red spots.

Rhodamnia cinerea Jack (— RA. trinervia Bl.). Gall 88 I. MacRitchie Reservoir.

(Pl. 10, a, b).

Moth-gall. Woody fusiform swelling of the stem, reaching 6 cm in length and

1 cm in width. At the interior one finds a large gall-cavity containing only one

larva.

Distribution: Singapore-Java. Sumatra (Bib. 21, p. 143, fig. 35-22, p. 418 no

1098, Ist).

44 Gardens’ Bulletin, Singapore-— X XVII (1974)

Idem. Gall 88 II. Mac Ritchie Reservoir.

Psyllid-gall. Adaxially it is a depression of the lamina 0.5 mm in depth on a

base 0.5 mm in diam., caused by the psyllids installed adaxially.

Idem. Gall 72 IV. B.T.N.R. and Mac. Ritchie. Reservoir. (Pl. 10, c).

Homoptera-gall? Local inhibitions of growth of the veins, provoking the

distortion of the leaf. :

RHIZOPHORACEAE.

Anisophyllea disticha Baill. Gall 114. Everywhere in B.T.N.R. (Pl. 12, b).

Midge-gall. Gall pyriform with a long neck, 3 mm long and 1 mm wide,

covered with long hairs, attached at the base of the leaf to the adaxial edge of the

lamina, containing a large longitudinal gall-cavity, in which one finds a midge-

larva. Galls are sometimes fused in twos or threes.

Distribution: Singapore-Bangka, Rhio—Archipellago (Bib. 22, p. 402, no 1040,

fig. 746). |

RUBIACEAE.

Lasianthus maingayi Hk. f. Gall 76. B.T.N.R.

Cause undetermined. Inhibition of growth of the veins, hence the contorted

aspect of the leaf.

Psychotria ovoidea (Hook f.)} Wall. Gall 110. B.T.N.R. (Pl. 12, d).

Midge-gall affecting the leaf. Abaxial roughly hemispherical outgrowth of the

lamina, 5 mm in diam, corresponding adaxially to an umbonate outgrowth 2 mm

in height. The gall develops by hypertrophy: spectacular elongating of the cells

perpendicularly to the surface of the leaf, then division of the cells of the spongy

mesophyll. In the centre of the gall one finds a large rounded gall-cavity. Exit

of parasites abaxial.

Urophyllum hirsutum Hook. Gall 159. B.T.N.R., Jungle Valley track. (pl. 12, c).

Midge-gall. Fusiform swelling 6 mm in maximum width, affecting the stem,

petiole and midrib. In the last case, the gall is mostly visible abaxially. The organ

can be hypertrophic along a variable length. The gall-cavity is elongated in the

direction of the spindle-axis.

SAPOTACEAE.

Palaquium obovatum (Griff.) Engl. var. obovatum. Gall 125. B.T.N.R.

Aphid-gall? Under the influence of the Homoptera fixed on them, the various

organs of the host-plant (stem, veins), are submitted to inhibitions of growth that

cause them to be twisted. The leaves are contorted, their edges become involute.

Palaquium semaram H. J. Lam. Gall 78, 116. B.T.N.R. (Pl. 12, a).

Midge-gall. Disc-shaped gall affecting both faces of the lamina, gall less than

1 mm thick, and diam. approximately 4 mm. Inside one finds a small larval cavity

containing one midge larva. That larva leaves the gall through an adaxial

emergence hole before pupating. The gall is yellowish, browning and drying out

on maturity.

Cecidia of Singapore 45

TILIACEAE.

Grewia blattaefolia Corner Gall 131. B.T.N.R., Jalan Kutu. (Pl. 12, e, f,).

Midge-gall. This light yellow spherical gall, less than 5 mm across is attached

on the adaxial or abaxial side of the leaf, laterally to the veins. The pyriform

gall-chamber stretches out perpendicular to the vein.

VERBENACEAE.

General comments concerning the galls occuring on the genus Clerodendron:

Aphids are parasitic on a number of Clerodendron and provoke the formation of

characteristic acrocecidia :

— Inhibition of the growth of the veins, hence the contorted aspect of the leaves.

— Inhibition of the growth of the internodes.

These galls were considered as pseudogalls by previous authors, on account of

their inhibitionary effect. See an illustration of this type of gall Pl. 13, b. This type

of gall has been found on:

C. deflexum Wall. Gall 95. B.T.N.R.

C. laevifolium Bl. (= C. disparifolium Bl.). Gall 119 I; 122. B.T.N.R.

Clerodendron laevifolium Bl. ( = C. disparifolium Bl.) Gall 119 II, 219. B.T.N.R.

(fig. 5, f).

Midge-gall. The edges of the lamina are involute. The distal end of the leaf

does not participate to the gall-morphogenesis. The external surface of the gall is

covered with white spots. Distribution: Singapore—Java (Bib. 22, p. 488, no 1300.

fig. 934).

Clerodendron sp. Gall 137, 151. Coronation Road (garden).

Mycocecidium. Rounded elevation of the lamina, a few mm in height, and 5

to 8 mm in diam., abaxially covered by small whitish vesicles (sori), disposed

concentrically. The yellowish gall turns brown on maturity, therefore causing the

whole leaf to dry out.

Vitex pubescens Vahl. Gall 64, 160. B.T.N.R. and Cherengi Road. (Pl. 13, f).

Mite-gall caused by Erviophyes cryptotrichus Nal. fide Keifer in litt.

Globulous and wart-iike pockets that project from the adaxial leaf-surface, 0.5 to

6 mm in diam., covered adaxially by short hairs, and abaxially by a dense

erineum. Distribution: this gall is very widespread. Singapore-Java, Sumatra,

Sebesy Island, Malacca, Siam (Bib. 12, p. 49, no 342, fig. 153.-14, p. 19.no 49-18.

p. 61, no 34.20, p. 311, no 77.-21, p. 148, no 82, fig. 44.-22, p. 493, no 1317, fig.

943-28, p. 766, no2756.—42, p. 59).

Vitex vestita Wall. Gall 126. B.T.N.R.

Mite-gall. Whitish erineum browning at maturity, covering the lamina in

rounded or reticular blotches abaxially but also adaxially.

VITACEAE.

Vitis gracilis Wall. Gall 80, 163, B.T.N.R.(Pl. 13 c).

Psyllid-gall. The psyllids fixed adaxially create a depression of the lamina,

less than 0.5 mm in depth on a base 0.5 mm in diam.

Gardens’ Bulletin, Singapore — X XVII (1974)

PTERIDOPHYTES. OLEANDROIDEAE.

Nephrolepis biserrata (Sw.) Schott Gall 240 I. Catchment Area. (PI. 13, e).

Gall caused by a mite, Nothopoda pauropus Nal. fide Keifer in litt. Small

finger-galls that project out from the adaxial and more frequently the abaxial

surface of the fronds. It is conical, 3 mm high, 1 mm wide at the base, having an

apical ostiole surrounded by hairs. The internal wall of the gall-cavity is glabrous

and longitudinaly fluted. When galls develop at margin of pinnae they are sub-

globular, about 2 mm high and 4 mm wide.

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Cecidia of Singapore 47

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Cecidia of ‘Singapore... | , 49

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1900. De Gallen der Djamboe-bladeren. De Indische Natuur. Algemeen

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50 Gardens’ Bulletin, Singapore — X XVII (1974)

ARR

Kv oT

2 hat) ‘

\ OM

‘i

Fig. 3. a & b, two different views of two young coccid-galls 23 I developed on a stipule

of Shorea curtisii King (on b tufts of hairs not drawn).

B: bud: G: gall; Is: foliar scar; st: stipule; sts: stipular scar; w: wad.

‘PEM 2 AK ‘Sapndis JOJ ¢ 8 ‘Z IS ‘| IS ApAEPWIS STapIO dAI}DOdSaI 9Y} JO Jeo] v Yow ‘¢/ *Z/ “T] ‘[[es :H ‘julod Buimoi3s :y

*d 9989

UBY} YOR}]v JO 93v}S JoNIVD UL O} spuOodsarIOS 1] ‘OIYdoIye Aja}o{]dWIOD sWUIOd9q sey }eY} Jeo] B UO podolsAop [[ed BunoA

‘p ‘yeWIOU SI JVoyT dy} JO JSd1 DY) ING ‘padnpor AJOA SI BUILT] 94) [[es OY) 0} 1XOU ‘qlIprur sy) Jo dy 94) 1e podojoaop

wy jes Bunod ‘9 S1ap1O puoses oy} JO sayndys oy} JO suo UO podo]sAap [[eVd ve JO odeRIS A[IvO *q SpoAOWDI Us0q sey IIed

yord jo aindns sug ‘png [eunou fe fsury rsiyund vasoysg UO T ¢Z S{[es-pi9909 BunoA oy} Jo syedse swOg ‘p BI

~~,

1S)

iy)

~~)

52 Gardens’ Bulletin, Singapore --.X XVII (1974)

geo,

Coe

. Mego

i 2,

~ "= 0tefegee

Fig. 5. a-e, galls on Shorea curtisii King; a, midge-gall 23 III attacked by a parasitic

chalcid; b, two developing stages of the midge-gall 23 III (left: early stage; right:

older stage.); ec, longisection of the midge-gall 23 Ill; d, wasp-gali 23 IH;

e, longisection of the same; f, thrips gall 219 on Clerodendron laevifolium BI.

ab: abaxial leaf-surface; ad: adaxial leaf-surface; cl: larval cavity; cp: cortical

parenchyma; e: epidermis; ex: external cap-side; i: internal cap-side: 1: larva;

mp: medullar parenchyma; o: ostiole; s: shell; tn: nutritive tissue; Wb-L/G:

vascular bundles of the leaf/gall.

Cecidia of Singapore

Fig. 6. Thrips-gall 98 I on Aporosa benthamiana Hk. f. & Th.

54 Gardens’ Bulletin, Singapore — X XVII (1974)

Fig. 7. Galls of Lithocarpus conocarpus (Oudem.) Rehd.; a, moth-gall 135 III on stem and

petiole; b, moth-gall 135 II on midrib; ¢, sessile wasp-gall 120 I; d, longisection

of the gall 120 I. On top, the detached lid of the gall.

ab: abaxial leaf-surface; ad: adaxial leaf-surface;

al: alveole; cl: larval cavity; li: gall-lid.

Cecidia of Singapore 55

AIX

Fig. 8. Witches’ broom 153 IV caused by coccids on Lithocarpus sundaicus (Bl.) Rehd.

a, pathological development of an axillary bud: formation of secondary and

tertiary shoots and reduction of leaves; b, sketch of the line drawing a,

c, hypertrophic base of a leaf; d, sketch of the witches‘broom.

A1X: cessation of growth of the principal axis; An, An+1, An+2, each a shoot

of the respective order: H: Homoptera; R: rosette of leaves due to the shortening

of internodes; BA: axillary buds abundantly developed: BS: axillary buds

strongly developed.

56 Gardens’ Bulletin, Singapore — X XVII (1974)

a PE erase e2es4 1 equivalnt plaat- parts

h equivalat sheots

bee ae owe om ow co om mm

few ewe ee -

yb i ae oe

normal growth \,

fom me me om { ih on Sl

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Fig. 9. Hypothesis on the growth vigor under the gall-zone, fide Dr. Cusset in litt. (see

p. 36 in text); a, monopodial growth with flush; b, sympodial and more or less

acrotonic growth; ¢c, monopodial growth without rhythm; d, observed gall-growth.

BA: axillary buds abundantly developed; BS: axillary buds strongly developed.

Cecidia of, Singapore We i 57

p. eh.

Fig. 10. Galls on Calophyllum pulcherrimum Wall.; a, psyllid-gall 107 I; note the twisted

petioles; b, midge-gall 107 III on stem longitudinally cut on the left; ec, midge-galls

107 il; d, longisection ‘of gall 107 II.

ab: abaxial leaf-surface; ad: adaxial leaf-surface cl: larval cavity: h: emergence

hole; N: midrib; p. chl; parenchyma with chlorophyll; TC: vascular bundle.

Gardens’ Bulletin, Singapore — XXVIT (1974)

FOO NTN ~

FoR GGG en

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a, Calophyllum ferrugineum Ridl. gall 141 II;

b, Calophyllum pulcherrimum Wall. gall 167 IV.

Leaf-galls caused by moths;

Fig. 11.

Cecidia of Singapore 59

Fig. 12. a, psyllid-gall 43 If on Calophyllum inophylloide King; b, coccid-galls 43 I on

same: ¢, coccid-gall 154 on stem of Xylopia malayana Hk. f. & Th.

Gardens’ Bulletin, Singapore — XXVIII (1974)

Fig. 13. a, midge-gall 143 IV on vein of Actinodaphne sp; b, longisection of the midge-gall

138 II on leaf of same; ¢, longisection of the midge-gall 151 on midrib of Popowia

tomentosa Maingay; d, transection of the midge-gall 68 I on leaf of Calophyllum

ferrugineum Ridl.

ab: abaxial leaf-surface; ad: adaxial leaf-surface;

cl: larval cavity; mr: midrib; nt: nutritive tissue;

o: ostiole; s: shell.

Cecidia of Singapore 61

Fig. 14. Endiandra sp.: a, gall 156 I caused by a thrips; b, gall 156 III, gall of the midrib

developed on the upper-surface of the leaf and gall 156 I caused by a thrips, on

the right part of the leaf.

Gardens’ Bulletin, Singapore — XXVII (1974)

Fig. 15. a, mite-gall 100 on leaf of Cyathocalyx ramuliflorus (Hk. f. & Th.) Scheff.: b-e, galls

on Nothaphoebe kingiana var. malvescens Gamble; b, abaxial view of the mite-

gall 133 III; ¢, section of the same; d, adaxial view of the midge-gall 94 II;

e, section of the same.

ab: abaxial leaf-surface; ad: adaxial ieaf-surface;

cl: larval cavity 1: larva: o: ostiole.

Cecidia of Singapore

Fig. 16. Pithecellobium clypearia

b, gall 115 I, median

chamber (CL); ¢, midge-gall 115 I abaxially attached to the midrib (n).

Benth.; a, gall 115 II, probably caused by Homoptera:

longisection of the midge-gall, showing the large larval

Gardens’ Bulletin, Singapore — X XVII. (1974)

Fig. 17. Galls on Eugenia rugosa (Korth.) Merr.; a, stem-gall 158 II, caused by an unidenti-

fied insect; b, longisection of the same gall; ¢c, longisection of the psyllid-gall

158 I; d, wasp-gall 142 IV, on stem;

ab: abaxial leaf-surface; ad: adaxial leaf-surface; c: emergence canal; cl: larval

cavity; h: emergence hole; nt: nutritive tissue; 0: ostiole; pc: cortical parenchyma;

ph: phloem; xy: xylem.

Cecidia of Singapore

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midge-galls (D) and galls containing a parasitic moth (L): ec, longisection of a gall

after emergence of parasitic moth with cocoon left behind in the gall cavity;

d, longisection of a gall after emergence of the midge; e, longisection of a young

midge-gali; f, a dehiscent midge-gall; g, psyllid-gall 22 11 on abaxial (ab) and

adaxial (ad) sides.

al: alveole; cl: larval cavity; co: moth-cocoon; dp: pupal integument of midge:

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Plate 2. Coccid-gall 23 I on Shorea curtisii King; a, a fully developed gall; b, longisection

of the same; ec, 2 coccid-gall attacked by parasitic wasps; d, longisection of the;

same; e, fully developed galls on the petiole (left) and on the leaf-margin (right);

f, young gall developed on the petiole (left) and on the leaf-margin (right).

co: coccids; cocl: larval cavity of coccids; o: ostiolar emergence canal; wel:

larval cavity of wasp; wh: emergence hole of wasp.

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Annotated list of seed plants of Singapore (II)

HSUAN KENG

Department of Botany

University of Singapore

II. Angiosperms — Dicotyledons

8. MAGNOLIACEAE

Key to the genera

A. Flowers terminal.

B. Aggregate fruit of a mass of fleshy, indehiscent and fused carpels; perianth-segments

18 or more, narrow (less than Icm wide) Aromadendron

B. Aggregate fruit of dry, dehiscent and + free carpels; perianth-segments 15 or less,

relatively broader.

C. Mature carpels dehiscing along the dorsa! (or outer) side Magnolia

C. Mature carpels dehiscing along the ventral (or inner) side, and separating from the

floral axis Talauma

A. Flowers axillary; mature carpels free, dehiscent along the dorsal side Michelia

Aromadendron elegans Bl.

Large tree with grey bark; flowers white, sweet-scented; fruit an ovoid mass;

found on Bukit Timah (Ngadiman 243), very rare. Vern. Champaka hutan.

Magnolia maingayi King

Shrub or smal! tree; flowers scented, white; formerly also found in Gardens’

jungle, now only on Bukit Timah (Ngadiman 36547), very rare.

Magnolia coco (Lour.) DC.

Shrub; flowers white, nodding, scented, the petals falling off shortly after

blooming; native of China. KBAR o

Michelia alba DC.

Tree; flowers white, strong scented; native of S. China, propagation by

marcots. Vern. Champaka puteh, A=E RR °

Michelia champaca Linn.

Tree; flowers orange, strongly scented; native of India, Thailand and northern

Malaya, producing viable seed. Vern. Champaka kuning, Kf °

Michelia figo (Lour.) Spr.

Small shrub; flowers ivory-coloured, scented but barely opened: native of S.

China. @27£

68 Gardens’ Bulletin, Singapore — X XVII (1974)

Talauma singaporensis Rid1.

Small tree; leaves very large; flowers white, large; carpels glabrous; formerly

found at Chua Chu Kang (Ridley s.n. date 1894) and elsewhere, now surviving

in Water Catchment Area, rare.

Talauma villosa Miq.

Recorded as T. lanigera Hook. f. in Ridley’s Flora; small tree; flowers white,

large; carpels woolly; formerly near Chua Chu Kang, Ang Mo Kio, Kranji

and elsewhere, now limited to Water Catchment Area (Nee Soon, Keng &

Jumali 6354), rare.

9. SCHISANDRACEAE

Kadsura scandens BI.

Big woody climber with smooth or rough bark; flowers rosy, on stems; Gardens’

jungle, Bukit Timah (Ridley 6354), rare. (A small leaved form of this species

was formerly called K. cauliflora Bl.) Vern. Akar dama-dama, AUAKF- °

10. ANNONACEAE

Key to the common genera

A. Carpels united into a fleshy mass, especially in fruit (cultivated fruit trees; connective

of anther truncate or oblong, concealing the anther locules.) Annona

A. Carpels free or slightly united but always quite free in fruit.

B. Petals 2-seriate, one or both series imbricate in bud (inner petals with a broad base;

anther connective broad).

C. Petals usually large (1-4 cm long), imbricate Uvaria

C. Petals smaller (below 8 mm long), valvate at base, imbricate at tips Cyathostemma

B. Petals 2-seriate or sometimes 1-seriate, all valvate.

D. All petals, or the inner ones cohering (or connivent) by their edges.

E. Petals 3, entirely connate into a + conical hood (flowers solitary, long-

pedicelled, pendulous, 6-8 cm long; ripe carpels necklace-shaped;

erect shrubs) Desmos dasymaschala

E. Petals 6 in 2 series, inner ones often shorter than the outer; cohering

into a long hood.

F. Erect trees or shrubs; flowers axillary, inner petals clawed (ripe

carpels bright red) Goniothalamus

F. Climbers; flowers extra-axillary; inner petals not clawed (ovary

densely pubescent) Friesodielsia

D. All petals free or separating very readily or shortly connate at the base only.

G. Connectives narrow, pointed, not concealing anther locules; trees or

errect shrubs.

H. Outer petals not or only slightly longer than calyx; the inner

much larger with saccate base; ovaries numerous (ovules 1-2)

Miliusa

H. Outer petals much longer than calyx; the inner not or only

slightly longer than the outer, sometimes smaller.

I. Flowers 4 to many in cymes, only 1-2 in each cyme open at

a time; petals with saccate base; all stamens perfect, ripe

carpels ellipsoid, fleshy, 3-12 seeded Alphonsea

I. Flowers solitary; petals not with saccate base, sometimes

obtuse; ripe carpels club-shaped, 2 valved, 1-2 seeded

Anaxagorea

Annotated list of seed plants of Singapore (11) 69

G. Connective widened and truncate, concealing anther locules.

J. Flowers borne on hooked, rigid, laterally compressed short

shoots; base of petals cohering into a cap over stamens and

carpels; climbers

Artabotrys

J. Flowers not borne on such hooked, short shoots.

K. Outer petals slightly longer than calyx (the inner

petals erect, concave inside; ripe carpels 1-2 seeded:

trees or shrubs) Popowia

K. Outer petals much longer than calyx.

L. Anthers transversely septate (trees, flowers axillary,

petals usually linear lanceolate: ripe carpels several

seeded)

Xylopia

L. Anthers normal, not septate.

M. Apex of connective acute; flowers very

fragrant, in pendulous, umbellate inflore-

scence; petals lanceolate or linear-lanceolate:

Ovaries numerous; ripe carpels ellipsoid, 2-

many seeded; trees Cananga

M. Apex of connective obtuse or truncate.

Alphonsea maingayi Hook. f. & Thoms.

Flowers solitary, extra-axillary or oppo-

site the leaves, pendulous, yellow; the in-

ner petals clawed at base; ovaries many;

Tipe carpels necklace-shaped with 1-8

segments; shrubs often with drooping

twigs Desmos

O. Not as above

P. Climbers, densely brown pubescent:

flowers in terminal cymes or panicles;

the outer petals erect or spreading,

the inner slightly shorter; ovaries

many, pubescent; ripe carpels stalked,

subglobose, 1-8 seeded _—_ Fissistigma

P. Trees or erect shrubs, rarely slightly

climbing.

Q. Flowers axillary, solitary or 2-3

together; petals subequal or the

inner ones shorter; all stamens

fertile, subequal; ovaries 2-5 with

several ovules in 2 rows; ripe

carpels thick-walled, fleshy, sessile

Meiogyne

Q. Not as above.

R. Trees with a crown of

branches near top; ficwers

usually opposite the leaves:

petals linear or oblong,

clawed; inner petals conceal-

ing the stamens and carpels

by the broadened base; ripe

carpels many-seeded

Cyathocalyx

R. Trees or shrubs; flowers

axillary or on branches or

stem; petals usually spread-

ing; Tipe carpels 1-5 seeded

Polyalthia

Tall tree; flowers in cymose clusters, opposite the leaves; outer petals with

teflexed tip and saccate base; ripe carpels oblong: seeds many: Chua Chu

Kang (Ridley 6758), Gardens’ jungle.

70 Gardens’ Bulletin, Singapore — X XVII (1974)

Anaxagorea javanica BI.

Shrub; petals spreading, the edges very thick; ripe carpels of 4-8 clavate

follicles; seeds 2, black, shining. Called A. scortechinii in Ridley’s flora. In

Singapore only found in Gardens’ jungle (Keng & Jumali 3027). 32i%f- «

Annona muricata Linn.

Shrub or small fruit tree; this and the two following species are native to

trop. America. Vern. Soursop, Durian belanda, i) GFA -

An. reticulata Linn.

Vern. Custard apple, Bullock’s Heart, 4-i(>% o

An. squamosa Linn.

Vern. Sweet-sop, sugar apple, ZA > eR

Artabotrys costatus King

Climber; petals creamy white, tomentose; Mandai (Ridley 10921).

Art. crassifolius Hook. f. & Thoms.

Stout climber; petals rose-pink; Bukit Timah (Ridley 10853).

Art. maingayi Hook. f. & Thoms.

Climber; common in MacRitchie Reservoir jungle (Sinclair 4876); petals green

to creamy yellow.

Art. suaveolens (Bl.) BI.

Climber; young branches glabrous or nearly so; peduncles hooked; flowers

creamy white, fragrant; the commonest species of the genus in Singapore;

Peirce Reservoir (Wee 1/84).

Aft.

unicinatus (Lam.) Merr.

Scandent shrub, native of Ceylon and S. India, sometimes cultivated, known

as Art. odoratissimus R. Br. ‘®)\ZE °

Art. wrayi King

Climber; petals pale yellow; Bukit Mandai (Ridley 3630).

Cananga odorata (Lamk.) Hook. f. & Thoms.

Tree, wild in N. Malaya and elsewhere, cultivated in gardens; Flowers very

fragrant; called Canangium odoratum King in Ridley’s Flora; a dwarf culti-

vated variety (var. fruticosa J. Sinclair) about 2 m. high, often has supernu-

merary petals but never sets fruit. Vern. Kenanga.

Cyathocalyx remuliflorus (Maingay ex. Hook. f. & Thoms.) Scheff.

Tall monopodial tree; branches spreading at top; Jurong (Corner 26162); called

Drepananthus ramuliflorus Maingay in Ridley’s Flora.

——————

Annotated list of seed plants of Singapore (11) 71

Cyath. ridleyi (King) Sinclair

Small tree, with pole-like trunk and crown branches at top; petals filiform,

pale yellow, pink at base, the claws orbicular; Bukit Timah (Sinclair 38449)

and elsewhere; called Xylopia ridleyi King in Ridley’s Flora.

Cyath. sumatranus Scheff.

Tree, with a crown of spreading branches at top; petals green, tomentose with

pink claws: Gardens’ jungle (Corner s.n. in 1943); called Xylopia curtisii King

in Ridley’s Flora.

Cyathostemma hookeri King

Climber, twigs glabrous; petals waxy-yellow; Gardens’ jungle (Ridley 4790).

Cyath. viridiflorum Griff.

Climber, often in swampy forest; flowers on stem, small petals, greenish

yellow; fruit orange, large; Seletar Reservoir (Sinclair SFN 39246).

Desmos chinensis Lour.

Bushy shrub, wild in Malaya and elsewhere, occasionally cultivated.

Desmos dasymaschala (BI.) Safford

Bushy shrub; petals cream coloured, 3-4 in one row (inner row absent);

MacRitchie Reservoir (Wee 49).

Desmos dumosus (Roxb.) Safford

Straggling shrub; petals greenish yellow, 6 in 2 rows; Cluny Road (Ridley

6305).

Ellipeia cuneifolia Hook. f. & Thoms.

Climber, young twigs rusty; petals fleshy yellow; Gardens’ jungle (Ridley

4919).

Fissistigma fulgens (Hook. f. & Thoms.) Merr.

Large climber; flowers few, in cymes opposite to a leaf; petals thick, ovate,

orange; ripe carpels globose to oblong, 3-4 cm long; seeds several in 2 rows;

called Melodorum fulgens Hook. f. & Thoms. in Ridley’s Flora. MacRitchie

Reservoir (Wee 135).

Fis. lanuginosum (Hook. f. & Thoms.) Merr.

Climber, young twigs rusty-tomentose; petals oblong-lanceolate, MacRitchie

(Wee 46).

Fis. latifolium (Hook. f. & Thoms.) Merr. var. ovoideum Sinclair

Large climber, twigs rusty-tomentose; flowers like F. fulgens but larger;

Seletar Reservoir (Wee 301).

72 Gardens’ Bulletin, Singapore — X XVII (1974)

Fis. manubriatum (Hook. f. & Thoms.) Merr.

Stout climber; flowers terminal, opposite a leaf; MacRitchie (Wee 173).

Friesodielsia biglandulosa (Bl.) Steenis

Climber; flowers extra-axillary; mature carpels ovoid or oblong, thin-walled,

l-seeded, usually stalked; Seletar Reservoir (Sinclair 38870); called Oxymitra

biglandulosa Scheff. in Ridley’s Flora.

Fri. glauca (Hook. f. & Thoms.) Steenis

Climber; Jurong (Corner SFN 26151).

Fri. latifolia (Hook. f. & Thoms.) Steenis

Climber; young branches rusty-tomentose; flowers solitary, extra-axillary;

petals creamy white; ripe carpels ovoid, clustered; Gardens’ jungle (Ridley

10809).

Goniothalamus malayanus Hook. f. & Thoms.

Shrub or small tree; flowers solitary, axillary or from the axils of fallen

leaves; petals valvate, greenish yellow, outer ones oblong-ovate; ripe carpels

oblong, 2—5 seeded, clustered; Seletar (Ridley s.n. in 1894).

Gon. ridleyi King

Tree; flowers in fascicles from warty tubers at base of trunk; ripe carpels

numerous, obovoid, glabrous, 1|-seeded, short-stalked; Bukit Timah (Ridley

4450).

Gon. tapis Miq.

Shrub or tree; flowers solitary or in pairs; leaves broader than Gon. malayanus;

ripe carpels red, 1-seeded; Bukit Timah (Ridley 8405).

Mezzettia leptopoda (Hook. f. & Thoms.) Oliv.

Tall monopodial! tree; petals linear, pale yellow, 1-!.2 cm long; fruit globose

with a resinous odour; Mandai (Corner SFN 33147).

Meiogyne virgata (Bl.) Mig.

Tree; petals lanceolate, 1.5-2.5 cm long, greenish yellow or pink, reddish at

base; ripe carpels oblong, brown-tomentose; Bukit Timah (Ridley 4457);

called Cyathocalyx virgatus King in Ridley’s Flora.

Miliusa longipes King

Shrub or small tree; flowers solitary, axillary, sepals and outer petals alike

(resembling Phaeanthus); inner petals greenish yellow outside, dark red inside;

ripe carpels globose, 1-2 seeded; Bukit Timah (Ridley 8450).

Mitrella kentii (Bl.) Mig.

Climber; leaves very variable; flowers 1-3, axillary; Bukit Mandai (Ridley

2117); called Melodorum elegans Hook. f. & Thoms. in Ridley’s Flora.

Annotated list of seed plants of Singapore (II) 73

Monocarpia marginalis (Scheff.) J. Sinclair

Tree; ripe carpels 1-3, subglobose, 4-6.5 cm across; called Cyathocalyx

maingayi Hook. f. & Thoms. in Ridley’s Flora.

Phaeanthus ophthalmicus (Roxb. ex Don) J. Sinclair

Small shrub, common in shady forests; flowers 1—2, extra-axillary; outer petals

and sepals alike; inner petals large, green, flat, ovate-lanceolate; ripe carpels

numerous, ovoid to oblong, l-seeded; MacRitchie (Keng & Jumali s.n., April,

1961); called Ph. nutans in Ridley’s Flora.

Polyalthia angustissima Ridley

Small tree; petals pink, linear; Bukit Timah (Ridley 8050, type).

Poly. cauliflora Hook. f. & Thoms.

Shrub or small tree; flowers arising mostly from woody tubercles on the

trunk and branches; petals pinkish brown, narrow ianceolate; Bukit Timah

(Ridley 8117), with 2 varieties, var. beccarii (King) Sinclair (with narrowed

leaves) and var. desmantha (Hk. f. & Thoms) Sinclair (with denser brown

tomentum esp. on midrib).

Poly. glauca (Hassk.) Boerl.

Tree; flowers fragrant; petals linear, oblong; Mandai (Holttum 37706).

Poly. hookeriana King

Tree; petals pale yellow; Fern Valley in B.T.N.R.* (Sinclair 39149).

Poly. hypoleuca Hook. f. & Thoms.

Tree: petals yellow, linear oblong; Jurong (Corner SFN 26163).

Poly. lateriflora King

Monopodial tree; leaves oblong, 20-40 cm long; petals pale yellow, with

tinges of red; Mandai (Corner SFN 33141); another related species, Poly.

sclerophylla Hook. f. & Thoms. with smaller leaves (15-20 cm long), also

found in Mandai (Kiah s.n. in 1940).

Poly. macropoda King

Tree; petals green, yellowish to white; Seletar (Ridley 429, type); a closely

related species, Poly. clavigera King with much longer stalked fruiting stalks

(2-3.5 cm vs. 0.5 cm), was found at Fern Valley in B.T.N.R. (Ridley 5857).

Poly. rumphii Merr.

Tree; Gardens’ jungle (Ridley 3863); called Poly. scortechinii King in Ridley’s

Flora.

Poly. sumatrana (Miq.) Kurz

Tree; petals greenish, linear, spreading: Seletar (Wee 338).

* Bukit Timah Nature Reserve.

74 Gardens’ Bulletin, Singapore — XXVII (1974)

Popowia fusca King

Shrub or tree; leaves thinner than Pop. pisocarpa; ripe carpels 1-4 globose-

falcate; Bukit Timah (Ngadiman SFN 36364).

Pop. pisocarpa Endl.

Shrub or small tree; flowers extra-axillary and leaf-opposed in small fasicies;

ripe carpels globose, 1-4 seeded: common; MacRitchie (Wee 1/53). Called

Pop. ramosissima Hk. f. & Th. in Ridley’s Flora.

Pop. tomentosa Maing. ex Hook. f. & Thoms.

Tree; Gardens’ jungle (Ridley 4916).

Pyramidanthe prismatica (Hk. f. & Th.) Sinclair

Climber; flowers axillary or terminal, solitary or in pairs; ripe carpels oblong,

tuberculate, 3-5 cm long; seeds in 2 rows; Kranji (Ridley 6344). Called

Melodorum prisomaticum Hook. f. & Th. in Ridley’s Flora.

Uvaria curtisii King

Large climber; petals white or greenish-yellow; Bukit Timah (Sinclair SFN

39652).

Uvar. cordata (Dunal) Alston

Climber; petals dark-red; common; Seletar (Wee s.n, Dec. 1961). Called

Uvar. macrophylla Roxb. in Ridley’s Flora.

Uvar. grandiflora Roxb.

Climber: petals purplish red; common; Bukit Panjang (Wee 90). Called Uvar.

purpurea Bl. in Ridley’s Flora.

Uvar. hirsuta Jack

Woody climber; petals dark red; Bedok (Ridley s.n. in 1877).

Uvar. leptopoda (Ding) R. E. Fries

Climber, petals dark crimson; Bukit Mandai (Ridley 4708, type): called

Uvariella leptopoda Ridl. in Ridley’s Flora; often confused with Uvar. rufa BI.

Uvar. lobbiana Hook. f. & Thoms.

Climber; collected once at Gardens’ jungle (Ridley 9211).

Uvar. pauci-ovulata Hook. f. & Thoms.

Stout climber with rusty-brown, stellate-hairy twig; petals yellow-brown; Bukit

Timah (Ngadiman SFN 36432).

Xylopia caudata Hook. f. & Thoms.

Tree; flowers very small; petals pale yellow; Jurong (Corner SFN 26030).

Annotated list of seed plants of Singapore (11) 75

Xyl. ferruginea Hook. f. & Thoms.

Tree, with stilt roots; leaves ferrugineous pubescence beneath; ripe carpels

numerous, elongate, constricted between the seeds; Gardens’ jungle (Abu

Kassim 2687).

Xyl. magna Maing. ex Hook. f. & Thoms.

Tree; petals linear, 8-10 cm long; Bukit Timah (Sinclair SEN 39690).

Xyl. malayana Hook. f. & Thoms.

Tree: flowers fragrant; petals pale yellow; ripe carpels 3 to many in a

bundle, irregularly dehiscent; seeds 2-7 in 2 rows; common; Chua Chu Kang

(Ridley 6759).

N.B. For the full range of species distribution of Annonaceae in Singapore, see

J. Sinclair, Gard. Bull. Sing. (1955) 14: 149-477.

11. MYRISTICACEAE

Key to the genera

A. Twigs not striate, often angled; leaves whitish beneath, reticulations not visible above

(aril laciniate to the base or nearly so; androecium columnar; anthers free at apex)

Gymnacranthera

A. Twigs striate (namely marked with fine grooves or ridges) or at least in the older parts.

B. Leaves often witish beneath.

C. Reticulations forming a dense, close network raised above in dried material,

(aril laciniate at the apex only; androecium a stalked disc with free anthers)

Knema

C. Reticulations few above, not forming a dense network, mostly sunk (aril laciniate

to the base; anthers completely fused to the stalked column) Myristica

B. Leaves not whitish beneath (aril entire, covering the seed; androecium various, always

sessile; anthers completely fused) Horsfieldia

Gymnacranthera bancana (Miq.) Sinclair

Tree, young twigs rusty-tomentose; flowers unisexual, in branched inflores-

cences; fruit rusty-tomentose, ovoid, 2.8 by 1.5 cm; Seletar (Ridley 1835),

Mandai. Called G. murtonii Warb. in Ridley’s Fiora.

Gymn. eugenifolia (A. DC.) Sinclair

Bukit Timah, Woodlands (Ridley 11646).

Gymn. forbesii Warb.

Seletar (Ridley 6270, type), Mandai

Horsfieldia irya Warb.

Tree, often slightly buttressed, by streams in swampy places; fruit round, 2-3

cm across, in bundles of 2-4; ochre yellow to pinkish orange; aril orange-red:

Changi (Ridley 4814), Gardens’ jungle.

Horsf. macrocoma Warb. var. canarioides (King) Sinclair

Bukit Mandai (Ridley 6324), Jurong.

76 Gardens’ Bulletin, Singapore — X XVII (1974)

Horsf. subglobosa Warb.

Bukit Timah (Ridley 6451), Mandai

Horsf. sucosa Warb.

Gardens’ jungle (Ridley 6559), Changi.

Horsf. superba Warb.

Bukit Timah (Negadiman 36141), Seletar. Vern. Pendarah.

Horsf. tomentosa Warb.

Collected only once from Bukit Timah (Cantley 30); recorded in Thailand,

Malaya and Sumatra. Vern. Manchong.

Horsf. wallichii Warb.

MacRitchie Reservoir (Corner 33556).

Knema conferta Warb.

Bukit Timah (Ridley 442, type), Mandai Vern. Panara Batu.

Knema curtisii Warb.

Bukit Mandai (Goodenough 3376), Jurong.

Knema furfuracea Warb.

Botanic Gardens (Sinclair 39488).

Knema glaucescens Jack

Jurong (Ridley 3873).

Knema globularia Warb.

Changi (Ridley 13342), Pulau Ubin.

Knema hookeriana Warb.

Small tree, leaves very large, densely covered with brown wool when young;

fruit also covered with brown wool; Water Catchment Area, Bukit Timah

(Ridley 3701). Vern. Singgah putih.

Knema intermedia Warb.

Bukit Timah (Ridley 2107), Changi, Jurong.

Knema latericia Elm.

Common in Water Catchment Area, Bukit Timah (Cantley 2/).

Knema laurina Warb.

Water Catchment Area, Bukit Timah (Cantley 3083).

Knema malayana Warb.

Bukit Timah (Cantley 20), Seletar.

Annotated list of seed plants of Singapore (11) 77

Myristica cinnamomea King.

Tree: fruit ellipsoid, reddish yellow, 7-8 cm long; aril thin, red, deeply

laciniate; Sembawang, Bukit Mandai (Ridley 3581la). Nee Soon, Sungei Buloh.

According to Ridley (Flora 3:66), this is the only Malayan species in which

the seed and aril have any spiciness. The seeds in all other species seem to

be deficient in aromatic properties, and are not used by the villagers at all.

Vern. Maiang pahong.

Myrist. crassa King.

MacRitchie Reservoir (Sinclair 39490).

Myrist. elliptica Hook f. & Th.

Bukit Timah (Ridley 6920), Seletar. Vern. Tabah, Pala hutan.

Myrist. fragrans Houtt.

The nutmeg of commerce is the seed. and mace is the red aril of the seed

of this species. A native of the Moluccas, once plantation crop in Singapore.

Vern. Pokok pala. AG °

Myrist. maingayi Hook. f.

MacRitchie Reservoir (Corner 33555).

Myrist. maxima Warb.

Bukit Timah (Ridley 3363) and Nee Soon.

N.B. For the full range of species distribution of Myristicaceae in Singapore, see

J. Sinclair, Gard. Bull. Sing. (1958) 16: 205-472.

12. MONIMIACEAE

Matthaea sancta BI.

Large shrub: leaves opposite, light green; flowers yellow: drupe blue black.

In forests and thickets, rare: Gardens’ jungle, Chua Chu Kang, Ang Mo Kio:

Bukit Timah (Ridley 9165).

13. LAURACEAE

Key to the genera

A. Hemi-parasitic herbaceous twiner: leaves totally reduced Cassytha

A. Trees or rarely shrubs: leaves normal. alternate, opposite or whorled.

B. Inflorescence umbellate (in Actinodaphne, a very short panicle is seen on full

expansion of the infl.). surrounded by an involucre of bracts.

C. Leaves alternate: bracts persistent: fruit seated on a cup formed by the per-

sistent perianth-tube.

D. Anthers 2-loculate Lindera

D. Anthers 4-loculate Litsea

C. Leaves in false whorls: bracts deciduous; fruiting pedicels strongly thickened.

fleshy Actinodaphne

78 Gardens’ Bulletin, Singapore — X XVII (1974)

B. Inflorescence paniculate.

E. Fruit on naked pedicel, or if perianth persistent, not cup-like.

F. Perianth in fruit hardened, clasping the base of fruit Phoebe

F. Perianth in fruit mostly deciduous, if persistent, neither hardened nor

clasping the base of fruit.

G. Fruiting pedicel strongly thickened, fleshy.

H. Anthers 2-loculate Dehaasia

H. Anthers 4-loculate Alseodaphne

G. Fruiting pedicel hardly or not thickened.

I. Anthers 2-loculate Beilschmiedia

I. Anthers 4-loculate.

J. Anthers sessile Nothaphoebe

J. Anthers with long filaments Persea

E. Fruit seated either on a shallow or deep cup, or completely included in the

persistent perianth-tube.

K. Fruit seated on a shallow cup; anthers 4-loculate Cinnamomum

K. Fruit completely included in and adnate to the persistent and enlarged

perianth-tube; anthers 2-loculate Cry ptocarya

Actinodaphne glomerata Nees

Tree; Bukit Timah, Bukit Mandai (Ridley 6747), Bukit Panjang.

Actin. hullettii Gamble

Botanic Gardens (Cantley 38); ‘“‘A single tree on the bandstand, which died

about 1911” (Ridley).

Actin. maingayi Hook. f.

Tree, in dense forests; Bukit Timah, Bukit Mandai, Pulau Ubin (Ridley 9489).

Actin. malaccensis Hook. f.

Tree, in forests; Gardens’ jungle, Bukit Timah (Ridley s.n. in 1897). Vern.

Medang kachigawei.

Actin. pruinosa Nees

Tree; Botanic Gardens, Changi (Ridley 2768).

Alseodaphne bancana Miq.

Shrub; Gardens’ jungle, Bukit Timah, Mandai Road (Sinclair 10900).

Beilschmiedia kunstleri Gamble

Big tree; Bukit Timah (Bayliss 5885).

Beils. malaccensis Hook. f.

Tree; Bukit Timah, Gardens’ jungle (no specimens available).

Cassytha filiformis L.

Leafless, hemi-parasitic twiner, yellow, common in open bushes near the

sea; Changi (Ridley 3380). Vern. Chemar hantu, 8G #8 i ©

Annotated list of seed plants of Singapore (11) 79

Cinnamomum camphora Sieb.

The oriental camphor is obtained by distillation from the wood of this

tree, native of S. China and Japan: occasionally planted. ifs} o

Cinn. iners Reinw. ex BI.

Small or medium-sized tree, very common in open places all over the Island:

young leaves reddish; fruit blue-black; Tyersall Road (Ridley 3370). Vern.

Kayu manis.

Cinn. javanicum Bl.

Tree, in dense woods: Bukit Timah (Ridley 15621), Gardens’ jungle.

Cinn. ridleyi Gamble

‘“‘A single tree on the Changi Road near Changi’ (Ridley), (no specimens).

Cinn. verum J.S. Presl

Native of S.W. India and Ceylon, formerly called Cinn. zeylanicum Garc. ex

BI.; of which the bark is cinnamon of commerce; a smaller flowered variety

(or called Cinn. deschampsii Gamble} was found in the ground of Botanic

Gardens. Vern. Kayu manis, #4) RAKE

Cryptocarya caesia BI.

Tall tree; Bukit Mandai, Chua Chu Kang (no specimens), Vern. Kayu Grisek.

Crypt. ferrea Bi.

Big tree; Chua Chu Kang (Ridley 6158), Bukit Mandai. Vern. Medang merah.

Crypt. griffithiana Wight

Tree; Tempinis, Changi, Seletar, Tebau (Goodenough 3382). Vern. Medang

buaya.

Crypt. impressa Mig.

Tall tree, rare; Gardens’ jungle, Bukit Timah (Ngadiman 36147). Vern.

Munyjuat.

Crypt. kurzii Hook. f.

Tree; recorded from Chua Chu Kang (no specimens).

Dehaasia incrassata (Jack) Kosterm.

Tree, formerly known as D. microcarpa Bl.; Sungei Jurong, Bukit Mandai

(no specimens).

Dehaas. nigrescens Gamble

Tree, once collected from Gardens’ jungle (no specimens).

Lindera lucida (Bl.) Boerl.

Medium sized tree, formerly called L. malaccensis Hook. f. Small tree;

Gardens’ jungle, Tanglin, Bukit Timah, Seletar (Ridley 3373).

80 Gardens’ Bulletin, Singapore — X XVII (1974)

Litsea accedens (Bl.) Boerl.

Tree, formerly known as Lits. singaporensis Gamble; Gardens’ jungle, Bukit

Timah, Kranji (Ridley 6740), Tuas.

Lits. cordata Hook. f.

Tree; Kranji (Mat. 4743), Bukit Timah.

Lits. costalis (Bl.) Kosterm.

Large tree, formerly called /its. megacarpa Gamble; Tuas, Reservoir Jungle

(Corner s.n. in 1930).

Lits. erectinervia Kosterm.

Tree, formerly called Lits. griffithii Gamble; Gardens’ jungle, Reservoir woods

(Ridley 4823).

Lits. ferruginea Bl.

Tree; Gardens’ jungle, Seletar (Ridley 6151).

Lits. firma Hook. f.

Tree, fruit pink; Bukit Timah (Ngadiman 35509), Chua Chu Kang.

Lits. garciae Vidal

Tree, formerly called Lits. sebifera Bl.; native of Java, occasionally cultivated.

Vern. Malek.

Lits. grandis Hook. f.

Tree, common in open places; Tanglin, Bukit Timah, Changi (Ridley 4700).

Vern. Medang busuk, Medang Daun Lebar.

Lits. lanceifolia Hook. f.

Small tree; Gardens’ jungle, Bukit Timah (Ridley 12563), Seletar.

Lits. lanceolata (Bl.) Kosterm.

Small tree, once collected from Reservoir woods (Ridley 4817); formerly

called Lits. sarawacensis Gamble.

Lits. machilifolia Gamble

Tree; Gardens’ jungle, Bukit Mandai (Md. Shah & Kadim 413).

Lits. myristicaefolia Hook. f.

Tall tree; Geylang, Changi, Tanglin (no specimens).

Lits. perakensis Gamble

Large tree; recorded once in Bukit Timah (no specimens).

Lits. petiolata Hook. f.

Large tree: highly valued in native medicine; Gardens’ jungle, Cluny Road,

Geylang (no specimens).

= . ls

Annotated list of seed plants of Singapore (11) 81

Lits. ridleyi Gamble

Small tree: Changi, Gardens’ jungle, Bukit Mandai (Ridley 5064).

Lits. robusta BI.

Tree; collected once from Singapore by Wallich (no specimens).

Micropora curtisii Hook. f.

Tree, recorded once from Jurong swamp forest (no specimens available). The

monotypic genus Micropora Hook. f. is endemic to the Malay Peninsula

(Penang) and Singapore.

Nothaphoebe coriacea Kosterm.

Tree; Tuas (Goodenough s.n. in 1890).

Notha. umbelliflora (Bl.) BI.

Tree; Chua Chu Kang; Bukit Timah (no specimens). Vern. Medang Losa.

Persea americana Mill.

Native of C. America, cultivated in gardens for the large edible fruit known

as “‘avacado” or “‘alligator pear’. #24! » RE4i o

Phoebe cuneata BI.

Tree; once collected at Changi (no specimens).

Phoebe declinata Nees

Tree; Changi (no specimens).

Phoebe grandis Merr.

Tree; MacRitchie Reservoir (Corner 37269).

Phoebe opaca Bl.

Large tree, Gardens’ jungle, Changi, Pulau Ubin (Ridley 5915).

14. HERNANDIACEAE

Hernandia nymphaeifolia (Presl) Kubitzki

Tree with large glossy peltate leaves; collected from Singapore coast by

Wallich in 1822, no specimens available in Singapore; extinct long ago but

survived in S. Johore and elsewhere; formerly called H. peltata Meisn. or

H. ovigera L. {= fil ©

Hern. cordigera Vieill.

A bush, native of New Caledonia, rarely cultivated and under the erroneous

name of Aralia triloba Hort.; flowering frequently but never sets fruit.

Illigera trifoliata (Griff.) Dunn

Woody climber; leaves alternate, 3-foliolate; recorded from Bukit Timah

forest (no specimens available); called I/ligera appendiculata BI. in Ridley’s

Flora.

82 Gardens’ Bulletin, Singapore — X XVII (1974)

15. RANUNCULACEAE

Clematis dioscoreifolia Levl. & Van. var. robusta Rehder

A slender twiner of Japanese origin (possibly also one or two other horti-

cultural species), is sometimes cultivated in gardens for its white flowers

and plumose achenes. {i fRiE ©

16. MENISPERMACEAE

Key to the genera

A. Leaves distinctly peltate (namely, petiole attached to the under surface of blade);

inflorescence a large panicle.

B. Leaves round, with dense white tomentose hairs beneath Coscinium

B. Leaves ovate or heart-shaped, not with white hairs beneath Cyclea

A. Leaves not peltate (namely, petiole attached to or near the base of blade).

C. Inflorescence a few-flowered, axillary cyme; leaves 3-nerved from base.

D. Slender woody climber; leaves glabrous Cocculus, Hypserpa

D. Large woody climber; leaves tomentose Limacia

C. Infl. raceme-like.

E. Racemes mostly solitary.

F. Leaves cordate, glabrous, 5-nerved from base Tinospora

F. Leaves ovate, tomentose, 3-nerved from base Limxia

E. Racemes few to many in fascicles from old twig; leaves ovate, glabrous, 3-

nerved from base Timomiscium

C. Infl. paniculate; leaves ovate, glabrous, 3-nerved from base Fibraurea

Cocculus trilobus (Thunb.) DC.

Slender climber, twining on bushes near sandy shore; formerly called C. ovali-

folium DC.; Changi (Sinclair 40021). AKG

Coscinium wallichianum Miers

Woody climber with yellow sap; leaves large rounded, white tomentose be-

neath; flowers in dense globose heads, on racemose or paniculate infi.; Chua

Chu Kang, Changi (Ridley 3967). Vern. Akar Kunyit.

Cyclea laxiflora Miers

Slender hairy climber; leaves ovate or heart-shaped; in hedges and forests;

local medicinal plant; Changi, Bukit Timah (Ridley s.n. in 1898), Tanglin.

Vern. Akar gasing Bukit.

Fibraurea chloroleuca Miers

Woody climber, with yellow sap formerly used in dyeing; fruit orange; com-

mon in forest; Bukit Mandai (Ridley 3834), etc. Vern. Akar Kuning, Akar

Kenching kerbau.

Hypserpa cuspidata Miers

Woody climber, in hedges and thickets; Chua Chu Kang (Ridley 6189). Seletar,

Tanglin. Vern. Akar minyak.

Annotated list of seed plants of Singapore (11) 83

Limacia velutina Miers

Woody climber; branches and leaves covered with rusty hair; common in

open thickets; Chua Chu Kang, Bukit Mandai (Ridley s.n. in 1893).

Tinomiscium petiolare Miers ex Hook. f. & Th.

Woody climber, similar to Fibraurea chloroleuca in general appearance but

with profuse white sap and green fruit: common on forest edge; Bukit Mandai

(Goodenough s.n. in 1892).

Tinospora crispa Miers ex Hook. f. & Th.

Woody climber with numerous warts on stem; native of India, cultivated as

a medicinal plant.

17. NYMPHAEACEAE

Key to the genera.

A. Petiole stout; leaf-blade, high above the water; flower showy; carpels free, immersed

on ob-conical receptacle Nelumbo

A. Petiole slender; !eaf-blade generally floating on the water surface: carpels united into

a many-chambered ovary.

B. Flower hardly expanded; outer perianth-lobes (sepals) 5 Barclaya

B. Flower showy; outer perianth-lobes (sepals) 4 Nymphaea

Barclaya motleyi Hook.

Aquatic, rhizomatous herb, in shallow muddy streams in thick jungle; leaves

round, dark-green; Bukit Timah (Ridley s.n. in 1892), Ang Mo Kio, Nee Soon.

Vern. Daun kelapu.

Nelumbo nucifera Gaertn.

“The Indian Lotus” is sometimes cultivated as an ornament; the rhizome

imported from China as a vegetable. Native of India and China. {aj7§ > si ©

Nymphaea capensis Thunb.

“The cape-blue water lily” originated from S. Africa. Leaves glabrous beneath:

sepals without dark markings; petals blue-purple or purple.

Nymphaea nouchali Burm. f.

‘Blue lotus of Egypt’, often called Nymphaea stellata Willd., is a native of

tropical Africa. Leaves glabrous beneath; sepals with dark lines or marks;

petals light blue or purple. + )X fHETHE ©

Nymphaea pubescens Willd.

Sometimes erroneously called Nymphaea lotus (non L.), native of Africa:

leaves densely short hairy beneath; flowers mostly white, occasionally pink or

red.

18. CERATOPHYLLACEAE

Ceratophyllum demersum L.

Hornwort, a submerged water plant with whorls of simple, but finely dissected

leaves; occasionally found growing in aquaria. 4 43s ©

yyy ~

————---

Morphology and ecology of some introduced herbaceous legumes

K. H. CHOW

Department of Botany, University of Singapore

Abstract

Eight tropical herbaceaus legumes were introduced into Singapore and evaluated for

their possible roles in soil fertility improvement and soil erosion control on the island,

and also in the other wet tropical areas. Desmodium canum is too woody and grew into a

small bush in two years; Desmodium intortum is seriously attacked by insects; Desmodium

sandwicense has upright growth habit and poor nodulation; Glycine wightii is attacked by

insects and has not flowered in Singapore; Phaseolus atropurpureus and Phaseolus lathroides

are two climbers and can be used as ornamental plants on fence; Stylosanthes humilis is an

annual plant and not well adapted to the wet tropics. Desmodium uncinatum is the only

legume showing potentials in fertility improvement and erosion control of soils in Singapore.

It has moderate vegetative growth, spreading growth habit and strong stoloniferous develop-

ment. It nodulates well, regrows fast after being cut and is well adapted to the wet tropics.

Introduction

In addition to the direct use as food and feed, the herbaceous legumes play

a significant and fundamental role in the soil fertility improvement and soil erosion

control. They are characterized by two important morphological features: 1) Their

roots bear nodules containing Rhizobium species which play a unique and vital

role in nitrogen fixation; and 2) they tend to root at the stem nodes and produce

stolons which are essential in soil erosion control.

The purpose of this study is to examine morphologically and ecologically

some introduced herbaceous legumes grown under the natural conditions in Singa-

pore so as to evaluate their usefulness on soil fertility improvement and soil

erosion control.

Materials and methods

Eight herbaceous legumes had been introduced from 4 tropical and subtropical

areas and grown for morphological and ecological investigations. A list of the

legumes and seed sources is presented in Table 1.

Table 1. Legume species introduced to Singapore for morphological and ecological studies

Species Identification Seed Source

Desmodium canum (Gmel.) Schin. & Thell. PI 317045 Hawaii

Desmodium intortum (Mill.) Urb. Greenleaf Australia

Desmodium sandwicense E. Mey. PI 322470 Brazil

Desmodium uncinatum (Jacq.) DC Silverleaf Australia

Glycine wightii (R. Grah. ex Wight & Arn.) Verdc. Perenn. soybean Brazil

Phaseolus atropurpureus DC. Siratro Australia

Phaseolus lathyroides L. Phasemy bean Puerto Rico

Stylosanthes humilis H.B.K. Townsville style Australia

io ¢)

86 Gardens’ Bulletin, Singapore — X XVII (1974)

One hundred seeds of each species were counted and weighed to the nearest

mg and scarified by rubbing between two fine carborundum papers before germina-

tion. Each lot of hundred seeds were placed on moistened blotting paper in a

petri dish. The time interval leading to germination was recorded. When the radicle

reached length of about 1 cm, all the young seedlings from each lot were trans-

planted to 8 cm-diameter plastic pots containing local soil on which Desmodium

triflorum and Desmodium heterophyllum have previously been grown. The two

legume species nodulated well so that the soil contained sufficient nitrogen-fixing

bacteria. The pots were placed under shade and gently watered twice daily. When

the seedlings reached the Sth true-leaf stage, they were transplanted to the field in

the Department of Botany at the University of Singapore and grown under natural

conditions. At the time of transplanting, the field is fully covered with grass-legume

mixture, with about 20-30% of legumes, mainly Desmodium heterophyllum and

Desmodium triflorum.

Morphological descriptions, measurements, and other observations of seed,

seedling and adult plant are recorded under individual species. Leaflet size was

measured by automatic area meter (Model AAM-5 by Hayashi Denko & Co. Ltd.)

to the nearest mm’. Pubescence on stems and leaves were classified into 4 classes,

namely glabrous, slightly pubescent, pubescent, and densely pubescent.

Ecological observations are aslo listed under each species. A minimum of 5

plants from each species were examined for nodulation. The nodulation value for

each species were recorded according to the following scale:

Rate 0 no nodulation.

Rate 1 poor nodulation: Sparse, small nodules.

Rate 2 fair nodulation.

Rate 3 good nodulation.

Rate 4 excellent nodulation: Abundant, large nodules.

Results and Discussions

Desmodium canum (Gmel.) Schinz & Thellung, Kaimi clover. — Plate 1A and 1B

This species originated from West Indies (Bryan, 1966). Younge et al (1964)

reported that it has been naturalized in the Hawaiian Islands and is known to the

farmers as Kaimi clover. This legume has been tested in Queensland, where it

persisted in spite of severe competition with the aggressive grasses.

Morphology

Seeds: Yellow to light brown, kidney-shaped, 1.2 x 2.0 x 3.5 mm, 1000-seed

weight 4.5-5.0g (180,000—220,000 seeds per kg); hilum oval, dark brown, at the

centre of straight edge; germination epigeal; time to germination at 27°C, 52 hours.

Seedlings: Cotyledons oblong, 4-5 x 6-8 mm; eophylls 2, opposite, unifolioliate,

cordate, 8-10 x 10-12 mm, margin entire, apex obtuse, base cordate: hypocotyl

6-8 mm; time to first true leaf, 25 days.

Stems: Decumbent to erect, cylindrical, green to brown, pubescent, fairly woody,

internodes 3-4 cm long, rooting at nodes.

Leaves: Alternate, pinnately trifolioliate, dark green, slightly pubescent on both

surfaces; petioles 3-5 cm long, pubescent; terminal leaflet oval, 8.8 cm’, 2-3 x 4-5,

margin entire; lateral leaflet 7.3 cm; stipules deltoid, 3-4 x 6-8 mm, apex pointed,

stipels minute, lanceolate.

Morphology and ecology of some introduced herbaceous legumes 87

Inflorescence: A terminal raceme, 10-20 cm long, with 20-30 flowers; flowers small

5-6 mm long, paired at node; pedicels 3-4 mm long, calyx campanulate, 5-cleft,

joined at base, teeth lanceolate; corolla pink to red, 4-5 mm long.

Pods: Straight, sickle-shaped toward end, 2-4 cm long, 5 mm wide, 4—8-seeded,

deeply indented at lower suture, covered with hooked hairs, indehiscent, readily

breakable into single-seeded segments.

Ecology

Kaimi clover has fairly big seed, about 4-5 grams per 1000 seeds, as com-

pared with other Desmodium species (Rotar and Urata, 1966). It germinated slowly

and the percentage of plant establishment was low, less than 20% under intensive

care. lt is a slow-growing plant and took 4 months to reach the flowering stage.

This legume was reported as a long-day plant (Chow, 1972). It flowered

throughout the year under the natural day length in Singapore. The percentage

of pod formation was low, less than 15%. Because of its low seedling vigour and

poor pod formation, it has difficulties in establishment and regeneration. However,

once it is established, it is very persistent and drought-tolerant, and competes very

well with the aggressive grasses. It was hardly attacked by insects and also roots

freely at nodes.

The plant did not nodulate well, with an average nodulation value of 1.1.

Moreover, the plant grew into a small bush in two years because of its very

woody stems. Thus it is concluded that this legume is fair in soil fertility improve-

ment but good in soil erosion control.

Desmodium intortum (Mill) Urb., Greenleaf Desmodium or Kuru vine. — Plate

IC and 1D

_D. intortum originated in Central and South America. It is found at altitudes

up to 2,500 m at the equator and at about 800 m at its geographical limit of

latitude 25°N and S (Bryan, 1969). This legume is used primarily for grazing.

It recovers rapidly and can be grazed periodically throughout the growing season

(Younge et al., 1964). This legume has grown in popularity in Queensland, Australia

and commercial seeds are available under the name “Greenleaf desmodium”’.

Morphology

Seeds: Light brown to brown, kidney-shaped, 0.6-1.0 x 0.8-1.2 x 1.0-1.4 mm,

1000-seed weight 1.5 g (400,000 seeds per kg); non-shattering; hilum small, nearly

round, covered with dark brown material, at the centre of the nearly straight edge;

germination epigeal, time to germination at 27°C, 26 hours.

Seedlings: Cotyledons oblong, 2-3 x 4-5 mm; eophylls 2, opposite, unifoliolate,

ovate, 7-10 x 8-12 mm, margin entire, apex obtuse, base obtuse; hypocotyl 4—6

mm; time to first true leaf, 22 days.

Stems: Decumbent to sub-erect, cylindrical to triangle, green to reddish brown,

pubescent, internode 4-8 cm long, 2-4 mm in diameter; rooting readily at nodes.

Leaves: Alternate, trifoliolate, green to dark green, pubescent on both surfaces:

margin entire; petioles 3-4 cm long, pubescent; terminal leaflet ovate, size 24.9

cm’, 5-6 x 6-8 cm, pubescent, with brown flecking; apex acute, base obtuse:

lateral leaflets smaller, 16.8 cm?, 4-5 x 5-7 cm; stipules deltoid, 3-4 x 4-5 mm;

Stipels small, linear.

Inflorescence: This plant has not flowered under the natural day length in Singa-

pore.

88 Gardens’ Bulletin, Singapore — X XVII (1974)

Ecology

Greenleaf desmodium has rather small seeds, about 1.5 g per 1000 seeds,

but the percentages of seed germination and seedling establishment were high.

The plant started slow at seedling stage, and then grew fast and vigorously after

it was established. The plant is not very tolerant to drought; vegetative growth

of this plant is limited to a certain extent in the drier season. This legume seems

to be best adapted to humid and subhumid climates in the tropics and subtropics.

The plant rooted freely at the stem nodes and nodulated very well. The

average nodulation value was 3.2, the highest among the legumes in this study.

The leaves of this plant were seriously attacked by insects (Chrysomelidae spp. and

Cicadellidae spp.). For many old leaves, only mid-ribs were left. D. intortum is

a shortday plant and flowered during December to March in Hawaii or under 10

to 11 hours of light (Rotar and Chow, 1971). Under the natural photoperiod in

Singapore, it has not flowered.

Although greenleaf desmodium is one of the most important legumes for

grazing animals in the tropics and subtropics, it can not be grown in Singapore

for soil conservation due to two reasons: (1) The plant is seriously attacked by

insects and (2) it does not flower with the intermediate day length at the equator,

thus will have difficulties in regeneration.

Note: Taxonomists have not clearly distinguished between D. intortum and

D. aparines (Link). DC. The type specimen of D. intortum is from Jamaica with

articles of the loment almost symmetrical. Some materials from Mexico, Central

and South America resemble the type but the loment articles are smaller and

asymmetrical. D. intortum ranges from southern Mexico, through Central America,

Jamaica and Haiti, into north Colombia and western Venezuela. D. aparines

reportedly occurs from Lake Titicaca northward, through the highlands of Vene-

zuela and Central America, to Vera Cruz in Mexico.*

This species designation of D. intortum has been used in this study.

Desmodium sandwicense E. Mey., Spanish clover. -— Plate 2A and 2B

This legume originated in South America and has been introduced into

tropical and subtropical Asia, Africa and Australia. It crosses readily with Des-

modium intortum and Desmodium uncinatum, the two most important pasture

species in the genus Desmodium (Rotar and Chow, 1971), and thus it was sug-

gested that this legume could be used as bridging species to combine genes from

these two species.

Morphology

Seeds: Light brown to brown, kidney-shaped, 1.2 x 2.0 x 3.0 mm, 1000-seed

weight 3-4 g (250,000-330,000 seeds per kg): non-shattering; hilum oval, dark

brown, at the centre of straight edge; germination epigeal, time to germination at

21, C. 2: Rots:

Seedlings: Cotyledons oblong, 4-6 x 8-10 mm; eophylls ovate, 6-10 x 5-8 mm,

margin entire, apex obtuse, base obtuse; hypocotyl 10-14 mm; time to first true

leaf, 18 days.

Stems: Erect to sub-erect, cylindrical, 1.5-3.0 mm in diameter, green, brown or

dark red, woody, glabrous; internode 3—5 cm long.

* Correspondence from B. G. Schubert, Arnold Herbarium, Harvard University.

‘yueld -q “Bullpass -D ‘wnjsojul wnipouseg “juejd :g ‘Buipoos :p ‘wnuns wnipowsaqg | 31eId

MLN INL IN UtI pamninad eta Reens poxmlgs Manin imnttiinipineg sh pipeoee

STRIATE

i dipesmsenutienae

Plate 2. Desmodium sandwicense, A: seedling, B: plant.

Desmodium uncinatum, C: seedling, D: plant.

Morphology and ecology of some introduced herbaceous legumes 89

Leaves: Dark green, glabrous above, slightly pubescent beneath, margin entire:

petioles 4-8 cm long, glabrous; terminal leaflet lanceolate, 9.8 sq. cm, 3-4 « 4-6

cm; apex acute, base acute; lateral leaflet smaller, 8.4 sq. cm, 2-3 x 4-5 cm;

stipules small, deltoid, 2-4 mm long, 1 mm wide, stipels minute, linear.

Inflorescence: A terminal raceme, 15—25 cm long, with 30-40 flowers; flowers 1 cm

long, paired at node; pedicels 3-5 mm long, calyx campanulate, 5-cleft, joined at

base, teeth lanceolate; corolla 8 mm long, pink.

Pods: Brown, straight, sickle-shaped toward apex, 3-5 cm long 0.5 cm wide, 5—11-

seeded, covered with hooked hairs, deeply indented at lower suture, indehiscent,

readily breakable into several single-seeded segments.

Ecology

Desmodium sandwicense showed high percentages of seed germination and

seedling establishment. The plant grew well and fast at seedling stage, but vegeta-

tive growth slowed down when the plant started to flower which took place within

two months from seeding. The plant flowered profusely but did not set seed well;

percentage of pod formation was less than 10% under the natural environment in

Singapore. This was probably owing to the high temperatures.

The plant had rather woody stems, and thus had upright to intermediate

growth habit. Nodulation was poor, with an average nodulation value 1.4. The

leaves were attacked by insects. It is concluded that this legume is not well

adapted to the rainforest conditions and is not recommended to grow in Singapore.

Desmodium uncinatum (Jacq.) DC., Silverleaf desmodium.— Plate 2C and 2D

This legume is native to tropical America, most likely Brazil (Bryan, 1969).

Its distribution is restricted to humid or subhumid tropical or subtropical climates

with a rainfall of 1000 mm or more. This species is one of the most promising

tropical pasture legumes. However, it is susceptible to ‘legume little leaf’ and is

not as palatable as D. intortum selections (Hutton, 1960).

Morphology

Seeds: Light brown, kidney-shaped, 1.5 x 2.3 x 3.3 mm, 1000-seed weight 5 g

(200,000 seeds per kg); non-shattering; hilum oval, dark brown, at the centre of

straight edge; germination epigeal, time to germination at 27°C, 25 hours.

Seedlings: Cotyledons oblong, 4-6 x 7-11 mm, apex obtuse, base obtuse; eophylls

2, opposite, unifoliolate, ovate, 8-12 x 10-15 mm, margin entire, apex obtuse,

base obtuse; hypocotyl 8-12 mm; time to first true leaf 20 days.

Stems: Decumbent, cylindrical to angled, green to light brown, 3.5 mm in diameter,

soft, densely covered with hooked hairs, very sticky; internodes 6-10 cm long,

rooting at the nodes on moistened soils.

Leaves: Alternate, pinnately trifoliolate, dark green with very distinctive silver

marking on the mid-rib, densely covered with hooked hairs on both surfaces;

petioles 6-8 cm long, densely pubescent: terminal leaflet elliptic and ovate, 15.5

cm’, 3-5 x 5-8 cm, margin entire, apex acute, base obtuse, lateral leaflets smaller,

12.5 cm?, 3-4 x 4-5 cm; stipules deltoid, 2-3 x 5-8 mm, apex pointed: stipels

minute, 3-4 mm long, lanceolate, pointed.

Inflorescence: A terminal raceme, 30-50 cm long, with 25-45 flowers; flowers

1.3-1.5 cm long, paired at node, pedicels 1-1.5 cm long, densely pubescent; calyx

campanulate, 5-cleft, joined at base, teeth acuminate, densely pubescent: corolla

pink, 1.1—1.4 cm long.

90) Gardens’. Bulletin, Singapore — X XVII (1974)

Pods: Sickle-shaped, 5—9-seeded, 4-6 cm long, 0.6 cm wide, upper suture straight,

lower suture deeply indented, covered with harsh and hooked hairs, very sticky,

indehiscent but readily breakable into several single-seeded segments.

Ecology

Silverleaf desmodium has fairly large seeds, 5 g for 1000 seeds, and grew well

at seedling stage. It is a short-day plant and flowered under 11 hours of photo-

period (Chow, 1972). In the natural environment of Singapore, it took 6 months

to reach flowering stage. The plant flowered well throughout the year but pod

formation was poor, less than 10%. Most of the flowers dropped within two days

after opening. This is probably due to the high temperatures in Singapore. Rotar

and Chow (1971) reported that pod formation of D. intortum and D. uncinatum

was negatively correlated with temperatures. Pods harvested in Singapore con-

tained an average of 2.8 seeds which is significantly less than the average of 4.5

seeds per pod reported by Rotar and Chow (1971) under the natural conditions in

Hawaii in winter.

After flowering, vegetative growth slowed down considerably. The plant rooted

freely at stem nodes and also nodulated well, with an average nodulation value 2.6.

Owing to its heavy pubescence, the plant is hardly attacked by insects. It is also

tolerant to shade, grew well in dry season, and regrew fast after being cut. It is

concluded that this legume shows potentials in soil fertility improvement and soil

erosion control in the wet tropics because of its high nodulation value, stoloni-

ferous habit, moderate vegetative growth, and tolerance to insects, shade and

drought. It has a spreading growth habit and builds up a dense ground cover in

half a year.

Glycine wightii (R. Grah. ex Wight and Arn.) Verdc., Glycine, perennial soybean

or Rhodesian Kudzu— Plate 3A and 3B

This legume is a native of tropical Africa, most likely Kenya. It is now

widely distributed in India, Ceylon, Malaya, and Java (Hutton, 1968). This plant

has received much attention in Australia, Africa and South America as a pasture

legume for semi-arid and for moderately humid tropics. In Kenya, Bogdan (1966)

reported that glycine mixed well with a number of cultivated grasses and is suitable

for improved permanent pastures.

Morphology

Seeds: Dark brown to black, oblong, 1.5 x 2.0 x 2.5 mm, 1000-seed weight 5.8 g

(172,000 seeds per kg); hilum very small, circular, white, at the centre of straight

edge: germination epigeal, time to germination at 27°C, 28 hours.

Seedlings: Cotyledons oblong, 5-7 x 8-12 mm: eophylls 2, ovate, 1.4-2.0 x

1.2-1.8, margin entire, apex obtuse, base obtuse; hypocotyl 10-14 mm; time to

first true leaf, 20 days.

Stems: Twining, cylindrical, soft, green, pubescent; internodes 8—15 cm long, rooting

at nodes.

Leaves: Green, pubescent on both surface; petioles slender, 3-5 cm long, pubescent;

terminal leaflet ovate, 14.1 sq. cm, 3-4 x 4-6 cm, margin entire, apex obtuse,

base obtuse; lateral leaflet smaller, 10.6 sq. cm: stiples small, deltoid, 3-4 x 6-8

mm, apex pointed; stipels minute, linear.

Inflorescence: This plant has not flowered under the natural day length in Singa-

pore.

Morphology and ecology of some introduced herbaceous legumes 91

Ecology

Glycine wightii grew slowly at seedling stage. Wilson (1972) stated that the

slow growth of this species at seedling stage was due to the relative slow initial

nodule development in this species. The plant grew and nodulated fairly well after

being established, with an average nodulation value of 2.2. This legume is a

short-day plant and has not flowered in Singapore. The leaves of this legume have

been attacked by insects, but not very seriously. Furthermore, this legume is a

climber and does not root well at stem nodes, and thus is not recommended for use

in soil conservation in Singapore.

Phaseolus atropurpureus DC., Siratro— Plate 3C and 3D

This species is native to Mexico, Central America and parts of South

America. It is usually found in drier areas at sea level or in the drier transition

zones at around 300 m elevation between the wet coastal lowlands and the higher

sub-coastal plateau (Hutton, 1962).

Morphology

Seeds: Dark brown, oblong-ellipsoid, 2.0 x 2.5 x 3.5 mm, 1000-seed weight 12.54 g

(80,000 seeds per kg); hilum oval, small, central, white; germination epigeal, time to

germination at 27°C, 31 hours.

Seedlings: Cotyiedons oblong, thick, small, 3 x 5S mm; seedling leaves 2, opposite,

cordate, 1.5-2.0 x 1.5-2.5 mm, apex, obtuse, base obtuse; hypocotyl 8-12 mm;

time to first true leaf, 15 days.

Stems: Trailing or climbing, cylindrical, green, soft, slightly pubescent; internodes

elongating.

Leaves: Alternate, trifoliolate, dark green, margin entire, pubescent beneath;

petioles 4-8 cm long, slightly pubescent; terminal leaflet ovate, size 11.8 cm’,

3-4 x 4-6 cm, apex obtuse, base obtuse; lateral leaflet smaller, 8.4 cm*; stipules

small, deltoid, 2-3 mm long, | mm wide, apex pointed; stipels minute, 1-2 mm

long, deltoid.

Inflorescence: An axillary raceme, with 6-10 flowers crowded at the apex of the

raceme; flower 1.5—2.0 cm long; calyx campanulate S-cleft, joined at base, teeth

short, pubescent; corolla dark purple, much exserted, keel prolonged, spirally

twisted, style filiform, wings conspicuous, 15-17 mm across.

Pods: Linear straight, 7-10 cm long, 0.4 cm wide, containing 11-13 seeds, dehi-

scent when ripe, splitting along both edges.

Ecology

The percentage of seedling establishment for this legume was low, and the

seedlings grew slowly in the environment of Singapore. The plant, however, grew

well after being established and flowered profusely throughout the year. Despite the

hot climate, it produced a large number of seeds. The pods of this legume are

dehiscent upon maturity. The plant did not volunteer well due to its poor seedling

establishment in wet and hot environment.

This legume is very resistant to insects and nodulated fairly well, with an

average nodulation value of 1.7 when the plant was 6 months old. The plant rooted

poorly at stem nodes. It is thus not recommended for soil conservation use in the

wet tropics. Being a climber, this legume can be used as an ornamental plant on

fence.

92 Gardens’ Bulletin, Singapore-- XXWVII (1974)

Phaseolus lathroides L., Phasemy bean — Plate 4A and 4B

This legume originated in India and is now widely grown in many parts of the

world. The plants are palatable and highly nutritive to animals. However, it is

susceptible to bean virus 2 and is badly attacked on occasion by bean flies

(Agromyza phaseoli Coq.) in Queensland, Australia, particularly at the seedling ©

stage (Hutton, 1962). In Australia, this legume is being replaced by ‘Siratro’

(Phaseolus atropurpureus L.) in many areas.

Morphology

Seeds: Gray, mottled with brown spots, oblong with rounded ends, solid, 1.6 x 2.0

x 2.2 mm, 1000-seed weight 6.5 g (150,000 seeds per kg); hilum oval, central;

germination epigeal, time to germination at 27°C, 19 hours.

Seedlings: Cotyledons oblong, small, thick, 3-4 x 4-6 mm; eophylls 2, opposite,

unifoliolate, cordate, 1.2-1.5 x 1.5-2.0 cm, margin entire, apex obtuse, base

cordate; hypocotyl 10-15 mm; time to first true leaf, 14 days.

Stems: Sub-erect, climbing, cylindrical, purplish green, slightly pubescent, soft;

internodes elongating.

Leaves: Trifoliolate, alternate, green, glabrous on upper surface, slightly pubescent

on lower surface; margin entire; petioles 4-6 on long, slender; terminal leaflet

ovate, size 12.5 cm’, 3-4 x 5-6 cm, apex obtuse, base obtuse; lateral leaflet size

9.2 cm’, deltoid, 1-2 x 4-6 mm, apex acuminate, stipules minute, 1-2 mm long,

lanceolate.

Inflorescence: An axillary raceme, 20-30 cm long, with 5—10 flowers crowded at

the top of raceme; flowers 1.5—2.0 cm long, purplish red, pedicels 0.5 cm long;

calyx campanulate, 5-cleft, joined at base, teeth short, pubescent; corolla purplish

red; keel prolonged, spirally twisted; wings conspicuous 16-18 mm across; style

filiform.

Pods: Linear, 10-14 cm long, 3 mm wide, covered with brown hairs, nearly

straight, containing 15-25 seeds, dehiscent when ripe, splitting along both edges.

Ecology

The seeds of phasemy bean germinated well, the seedlings grew fast and the

plants were very vigorous after being established. It flowered profusely throughout

the year in Singapore. The percentage of pod formation was about 65% and

each pod contained approximately 20 seeds, thus it produced a great number of

seeds. The pods dehisced when ripe and the plant volunteered well.

The young plants of two months old nodulated well, with an average nodula-

tion value of 3.1. For 6-month-old plants, the nodulation value decreased to 1.5.

The fast growth at the seedling stage of this legume is probably due to the fast

initial nodule development. Phasemy bean is probably a biennual plant. Its growth

slowed down considerably in the second year and gradually died off. The plant is

very resistant to insects.

The above results indicate that this legume is very well adapted to the natural

environment in Singapore. It is good on soil fertility improvement but fair on

soil erosion control because it is a climber and produces stolons poorly. It can —

be used as an ornamental plant on fence. However, precaution has to be taken

to prevent the plant from growing beyond control.

SS 8 CW TOC S28 ve ttt. &

Plate 3.

Glycine wightii, A: seedling, B: plant.

Phaseolus atropurpureus, C: seedling, D: plant.

Plate 4. Phaseolus lathroides; A: seedling, B: plant.

Stylosanthes humilis; C: seedlings, D: plants.

Morphology and ecology of some introduced herbaceous legumes 93

Stylosanthes humilis H.B.K., Townsville style — Plate 4C and 4D

This legume is a native of South America. It was introduced into Queensland,

Australia through the port of Townsville in the early years of this century (Shaw

et al., 1961). By 1945 this plant had become naturalized over a wide area in the

northern part of Queensland and is the only annual pasture legume in the sub-

tropical areas of the world. Hutton (1968) stated that the success of this legume

in Australia comes from its adaptability, good seed production—as high as

1,000 Ib per acre—ease of re-establishment, and tolerance to grazing due to

moderate palatability.

Morphology

Seeds: Brown or light brown, glossy, somewhat rhombic, with one end beaked,

1.2 x 1.5 x 2.0 mm, 1000-seed weight 2.13 g (470,000 seeds per kg); hilum small,

circular, brown, near the beaked end; germination epigeal, time to germination at

27° C,.20. hours.

Seedlings: Cotyledons oblong, 2-4 x 4-7 mm; eophyll 1, trifoliolate, lanceolate,

2-4 x 5-9 mm, margin entire, apex obtuse, base obtuse, hypocotyl 6-10 mm;

days to first true leaf, 20.

Stems: Erect or sub-erect, attaining 0.5 m high or more, green, soft, cylindrical

when young and slightly angled when old, pubescent; internodes 3-6 cm long.

Leaves: Alternate, palmately trifoliolate, green, slightly pubescent on both surfaces;

petiole slender, 1-2 cm long, pubescent; terminal leaflet lanceolate, size 0.78 cm’,

0.5 x 2-3 cm, apex acute, base acuminate; lateral leaflets smaller, 0.41 cm?, 0.4

x 1.5-2.5 cm; stipules conspicuous, 5-8 mm long, adnate to petioles, forming a

sheath of 1 cm long at the base; stipels absent.

Inflorescence: terminal or axillary head with 5—10 flowers; flowers 0.8 cm long,

yellow or light orange; calyx tubular, 5-cleft, joined at base, teeth acuminate, short,

covered with fine hairs.

Pods: Single-seeded, dark brown, 2-3 mm long, 2 mm wide, with a fine and

hooked stalk extending from one end, indehiscent.

Ecology

Townsville style has small, rhombic seeds which germinate well in petri dish.

The plant grew slowly at seedling stage and vegetative growth was somewhat

limited under the natural conditions in Singapore. It flowered profusely after 4

months from seeding and set a large number of shrivelled seeds. This is probably

due to the high temperatures. This species is annual and started to die off after

10 months from seeding in Singapore. The plant is very resistant to insects. It

did not produce any stolons and nodulated poorly. Thus this species is not recom-

mended to be grown in Singapore for soil conservation purpose. This species has

been reported to be best adapted to the drier areas in the tropics (Fisher, 1969).

Literature Cited

Bogden, A. V. 1966. Glycine javanica under experimental cultivation in Kenya.

Trop. Agr. 43: 99-105.

Bryan, W. W. 1966. The pasture value of species of Desmodium. Inter. Grassld.

Coner. Proc. 10: 311-315.

. 1969. Desmodium intortum and Desmodium uncinatum. Herbage Abs.

39 (no. 3): 183-191.

94 Gardens’ Bulletin, Singapore — X XVII (1974)

Chow, K. H. and L. V. Crowder. 1972. Hybridization of Desmodium canum (Gmel.)

Schin. & Thell. and Desmodium uncinatum (Jacq.) DC. Crop Sci. 12: 784-785.

Chuang, C. C. and C. Huang. 1965. The leguminosae of Taiwan for pasture and

soil improvement. JCRR, Taipei, Taiwan. 268 pp.

Fisher, M. J. 1969. The growth and development of Townsville lucerne (Stylosan-

thes humilis) in ungrazed swards at Katherine, N.T., Aust. J. Exp. Agr. &

Anim. Husb. 9: 199-208.

Hutton, E. M. 1960. Flowering and pollination in /ndigofera spicata, Phaseolus

lathroides, Desmodium uncinatum and some other tropical pasture legumes.

Emp Jour. Exp. Agr. 28: 235-243.

. 1962. Siratro —a tropical pasture legume bred from Phaseolus atropur-

pureus. Aust. J. Exp. Agr. & Anim. Husb. 2: 117-125.

—_———. 1968. Australia’s pasture legume. J. Aust. Inst. Agr. Sci. 34: 203-218.

Rotar, P. P. and K. H. Chow. 1971. Morphological variation and interspecific

hybridization among Desmodium intortum, Desmodium sandwicense and

Desmodium uncinatum. Hawaii Agr. Exp. Stat. Tech. Bull. No. 82, 28 pp.

Shaw, N. H., C. T. Gates and J. R. Wilson. 1966. Growth and chemical composition

of Townsville lucerne (Stylosanthes humilis). 1. matter yield and nitrogen

content in response to superphosphate. Aust. J. Exp. Agr. & Anim. Husb.

6;,, 159. |

Wilson, J. R. 1972. Comparative nodulation, nitrogen fixation and growth of

Glycine wightii cv. Cooper and Phaseolus atropurpureus cv. Siratro seedlings.

Aust. J. Agr. Res. 23: 1-8.

Younge, O. R., D. L. Plucknett and P. P. Rotar. 1964. Culture and yield per-

formance of Desmodium intortum and Desmodium canum in Hawaii. Hawaii

Agr. Exp. Stat. Tech. Bull. No. 59. 28 pp.

Anatomical Features of the Dipterocarp Timbers of Sarawak

F. CHU FEI-TAN

Forest Department, Sarawak*.

Summary

Economically, the family Dipterocarpaceae is one of the most important in Malaysia,

producing timber and other commodities.

The timber of some 200 species of dipterocarps indigenous to Sarawak has been

anatomically examined and found to conform to about a dozen homogeneous groupings

corresponding, in the main, to generic categories. The characteristics of these categories are

described in detail and summarised in tabular form. Anatomical relationships between the

groupings are discussed and attention is directed to similarities and differences between them.

Also, indicative specific gravities for the 200 species examined have been listed.

A key to the timber groupings, which have commercial application, has been prepared.

Introduction

About two-thirds of the total area of Sarawak is occupied by forests. Browne

(1955) has remarked upon the fact that, although these forests may contain as

many as 3,000 different species, they are dominated by the Dipterocarpaceae. ‘This

family of commercially valuable trees is, with the exception of the sub-family

Monotoideae which occurs only in Africa, confined to the Indo-Malaysian region

where the timber is used locally for many purposes and is an important com-

ponent of the nation’s overseas trade. In addition, these trees furnish dammar

(amber) used in the manufacture of varnish and an oil which is used in foodstuffs,

medicines and cosmetics. This oil is obtained from illipe nuts — the fruits of the

Engkabang (Shorea spp.) trees.

Over 200 species of Dipterocarpaceae have been recognised from Sarawak

but, for commercial purposes, these fall into about a dozen categories correspond-

ing, in the main, to generic groupings. The appearance and properties of many

of the timbers within the family are very similar and the differences between the

several groupings are not always easy to recognise.

Desch (1957) has described the gross structure of the dipterocarp timbers of

West Malaysia. Metcalfe and Chalk (1950) have summarised the anatomical features

of all the genera within the family but make little reference to specific differences

within each genus. This paper presents the results of an intensive study of specific

variation in the 9 genera of dipterocarp indigenous to Sarawak. The results of

this study have been summarised in the form of two tables, one dealing with

the anatomical features characteristic of each genus, the other setting out the

densities of the 200 species examined. In addition, a key for the separation of

the various timber groupings recognised by the timber trade has been prepared.

* Current affiliation: Timber Office, Department of Trade, Singapore.

96 Gardens’ Bulletin, Singapore —— X XVII (1974)

In the text, genera have been described in alphabetical order, regardless of

natural affinities. Field characteristics have been summarised from descriptions

published by Browne (1955) or Ashton (1964). The technical description of the

wood structure conforms to the usage of the International Association of Wood

Anatomists (1957). The density classification proposed by Kauman and Kloot

(1968) has been accepted as conforming sufficiently closely to commercial dif-

ferences in the dipterocarp timbers.

A projection microscope and superimposed scale (designed by F.P.R.L.,

Princes Risborough, U.K.) were used to measure the sizes of pores/vessels and

rays, and the sizes quoted conform to the system of the British Forest Products

Research Laboratory (1960). Hardness was determined subjectively, by shaving the

end-grain with a pocket knife, the rating being: —

soft, moderately hard, hard and very hard.

Acknowledgements

I am indebted to many members of the Forest Department, Sarawak, for the

collection of the timber specimens examined in this study. The botanical identity

of each specimen studied was established either by the Department’s former

Forest Botanist, Dr. P. S. Ashton, or, more latterly, by Dr. J. A. R. Anderson,

the Department’s Forest Research Officer. This contribution is gratefully acknow-

ledged. My thanks are also due to the staff of the Wood Technology Section,

Timber Research Centre, and particularly to Enche Adenan bin Dillah, who

prepared the microscopic sections for examination and who made the multitude

of measurements necessary to establish the quantitative data presented. Finally,

1 am most grateful for the constructive criticism and helpful advice of Mr. J.

Beesley, from Australia and to Mr. L. S. V. Murthy, Conservator of Forests, for

permission to publish this paper and for allowing me to have access to his

laboratory records.

KEY TO THE DIPTEROCARP TIMBERS OF SARAWAK

1 (a) Porés ‘exclusively solitary 22,02.) eset ate Zz

(b) Pores not exclusively ‘SOlitaty --2fo on ences fone se ee 7

2 (a) Wood distinctly yellow .... MERSAWA (Anisoptera spp.)

(b) Wood ‘not yellow 5.0342 ccc ucts eee co0 es epee et 3

3 (a) Axial intercellular canals in long tangential series ........................0.

KAPUR (Dryobalanops spp.)

00 6 6 6 6 e's 46 wie 6 6 OVS BO 6 86 wd: 5 6 0 6s wiel.o Sb G6 CKe Cen s

(b) Axial intercellular canals not in long tangential series ..................... 2

4 (a) Axial intercellular canals diffuse; wood texture fine ..................... a

(b) Axial intercellular canals in short tangential series; wood texture

WUAESE “1. cect ete eee ae KERUING (Dipterocarpus spp.)

5, (a). Pores, numerous to Very (NUMErOUS} .nipncagt. cgnteers- teal eee 6

(b) Pores moderately few ............ PENYAU (Upuna borneensis)

6 (a) Silica ‘present = Ag. 73s. RESAK (Cotylelobium spp.)

(b) -Sifica: absent 5... sete RESAK (Vatica spp.)

Anatomical Features of the Dipterocarp Timbers of Sarawak

7 (a)

(b)

13 (a)

(b)

Crystals in parenchyma, chambered, usually in long strands

IPAS A hur. Us. we. WHITE SERAYA (Parashorea spp.)

Crystals in parenchyma, not chambered, usually not in long strands

a OM a oe cc tec Sass « Be pn Si Sete Met aca eden hE ms ddan oe © « 8

aoe weed. Belew 00 ‘la. iperveu: ft. ali-dty ;... 20d d pS, kL... 9

enum ter TAP) 10e PSF CU IT, AIE-OTY <5... isc achencuepge see sce oversees 14

Pores in characteristic groups of 3-5; silica present ..................4.....

Piston tg iis. Ss aansacares «29035 WHITE MERANTI (Shorea spp.)

Poe Mi VeroupS OL 3-5: Silica absent ....670 00. Oe 10

fa) Wood texture. fine «......-.: SELANGAN (Hopea spp.)

en ER GS Arse A COTS i pi ce rn edd D.Talai 11

Wood yellow; radial intercellular canals always present; rays bright

yellow (transverse section) .... YELLOW MERANTI (Shorea spp.)

Wood not yellow: radial intercellular canals present only in a few

species of Red Meranti; rays not bright yellow ........................... 12

(a) Wood parenchyma prominent, tending to short lines between rays;

axial intercellular canals prominent on all surfaces

(b) Wood parenchyma not prominent; axial intercellular canals distinct

on all surfaces but not prominent

LIGHT RED MERANTI (Shorea spp.)

nici eo (ee 6 = ia alles a ©. 0. 6 a 0.0, 0 @ £2 @ 6 aie C48 C16 6.6 @ CUP at wa wis S62 4 e's we

RED SELANGAN (Shorea spp.)

DARK RED MERANTI (Shorea spp.)

(a) Diameter of axial intercellular canals considerably smaller than that

of vessels; wood texture coarse

bth bocive !.. forvlialy. des SELANGAN BATU (Shorea spp.)

(b) Diameter of axial intercellular canals often as large or larger than

that of vessels: wood texture fine

GIAM (Hopea spp.)

Pees esr aeees essere eeeeeeeereeneeeeeennee®

SUMMARY OF QUANTITATIVE VALUES

DENSITY CLASSES (see Kauman & Kloot, 1968)

Light < 30 lb. per cu. ft.

Medium 30 — 45 Ib. per cu. ft.

Heavy 45 — 60 Ib. per cu. ft.

Very heavy > 60 lb. per cu. ft.

98 Gardens’ Bulletin, Singapore— X XVII (1974)

PORE NUMBER (see F.P.R.L., Princes Risborough, 1960)

Very few < 12 per 10 mm?

Few 12— 30 per 10 mm?

Moderately few 30— 65 per 10 mm?

Moderately numerous 65 — 125 per 10 mm?

Numerous 125 — 250° per 10.mm?

Very numerous > 250 per 10 mm?

PORE and RAY SIZE (see F.P.R.L., Princes Risborough, 1960)

Vessel Diameter | Ray Width

_ — Tangential (Transverse section)

Microns Microns

2 | < 50 | Very small

a 50 — 100 | Moderately smail Ue Fai

+ 100 — 200 | Medium 50 — 100 | Medium

5 | 200 — 300 | Moderately large 100 — 200 Moderately broad

6 > 300 | Very large 200 — 400 | Very broad

Anisoptera

FIELD CHARACTERISTICS

Of the five species of Anisoptera (Mersawa) known to occur in Sarawak, two

are rare and of no commercial importance. Of the other three species A. grossi-

venia is the most common but not enough to rate the genus as important a timber

source as it is in the forests of northern West Malaysia, Thailand and Indo-China.

Commercial stands of this species and of A. laevis are found in the lowland forests

whereas A. marginata is restricted to peat swamp forests. The genus usually occurs

on yellow sandy soils, calcareous shales or sandstone hills and on peat. All species

develop into medium to large-sized trees, maximum height between 150-180 feet,

with a girth of about 15-18 feet, with straight, cylindrical boles. Buttresses are

prominent, thick and rounded. Bark surface is dull brown. dotted with warty

lenticels, rather irregularly fissured. Dammar (amber) exuding from surface fis-

sures appears as smears on the bole.

WOOD ANATOMY

Pores/Vessels. Almost exclusively solitary, with a few radial, tangential and

oblique pairs, uniformly distributed, moderately numerous, generally oval, medium-

sized to moderately large (size 4-5); intervascular pitting alternate, moderately

coarse, pits vestured; perforation plates simple; tyloses abundant.

Anatomical Features of the Dipterocarp Timbers of Sarawak 99

Parenchyma. Paratracheal, as incomplete sheaths to the vessels, occasionally

aliform; apotracheal, diffuse and in short uniseriate bands between rays, the

uniseriate bands being most distinct and numerous in A. Jaevis; around the axial

intercellular canals, sometimes forming more or less continuous tangential layers.

Rays. Mostly 2-7 cells wide (medium-sized), uniseriates (fine) short and rare,

less than 2 mm. in height; heterogeneous (Kribs’s types II] and II) with 1-3

marginal rows of square or upright cells.

Vessel-ray Pitting. Simple, round, with large aperture.

Tracheids and Fibres. Fibres with moderately conspicuous bordered pits, walls

thick. Cells intermediate between vasicentric tracheids and fibre-tracheids occur

in the immediate proximity of the vessels.

Intercellular Canals. Axial canals scattered singly and in pairs throughout the

fibres, sometimes in more or less continuous tangential rows; diameter usually

less than that of the vessels; canals commonly blocked with chalky deposits.

Crystals. Wanting.

Silica. Abundant in all species in ray cells, occasionally in parenchyma.

Growth Rings. Absent.

PHYSICAL PROPERTIES

Sapwood fairly distinct, pale yellow; about 2 inches wide.

Heartwood yellow, later darkening to yellow-brown.

The timber is moderately hard and medium to heavy, with an average

density of 47 Ib. per cu. ft. (range: 42-56 Ib. per cu. ft.) air-dry.

Grain shallowly to rather deeply interlocked; texture moderately coarse, even.

DISTINGUISHING FEATURES

(i) The presence of axial intercellular canals distinguishes this timber from

non-dipterocarps.

(ii) The diffuse axial canals preclude the possibility of confusing this timber

with other dipterocarp timbers except Keruing (Dipterocarpus spp.), Resak

(Cotylelobium spp. and Vatica spp.) and Penyau (Upuna borneensis).

(iii) The timber can be separated from Keruing, Resak and Penyau by its

distinctive yellow colour. Further, the timber of Anisoptera is much coarser in

texture, and less hard, than Resak and Penyau.

Cotylelobium

FIELD CHARACTERISTICS

Three species of this genus are known to occur in Sarawak on a variety of

soils on sandstone ridges and limestone hills. In Sarawak, all species of Cotyle-

lobium and Vatica are collectively known as Resak, although as trees, the two

genera are quite distinct in many of their characters. All species of Cotylelobium

grow to medium-sized trees, with a maximum height of 120-140 feet and a girth

of 8-12 feet. Frequently, the trees have twisted boles, with low and rounded but-

tresses, similar to Vatica. Bark surface is at first smooth, greyish and hoop-marked,

becoming irregularly, shaggily flaked with age. Creamy or yellow smears of

dammar present on the bole.

100 Gardens’ Bulletin, Singapore -- XXVII (1974)

WOOD ANATOMY

Pores/Vessels. Exclusively solitary, generally round, numerous, moderately

small to medium-sized (size 3-4); intervascular pitting rare, alternate, moderately

coarse, pits vestured; perforation plates simple; tyloses present.

Parenchyma. Paratracheal, usually as incomplete borders to the vessels, abax-

ially aliform, occasionally confluent; diffuse and diffuse-in-aggregates, sometimes

tending to short, uniseriate bands between rays; also around the axial intercellular

canals.

Rays. Uniseriate (fine) and 2-8 cells wide (medium-sized); low, less than 2

mm. in height; heterogeneous (Kribs’s types II and III, type I occasionally present

in C. burckii) with 1-3 marginal rows of square or upright cells; sheath cells

occasionally present.

Vessel-ray Pitting. Simple, round to elongated, with large aperture.

Tracheids and Fibres. Fibres with moderately conspicuous bordered pits, walls

very thick.

Intercellular Canals. Axial canals scattered singly and sometimes in pairs

throughout the fibres, diameter less than that of the larger vessels but as large

as some of the smaller; canals commonly blocked with chalky deposits.

Crystals, Wanting.

Silica. Present in ray cells of all species, occasionally in parenchyma but rarely

in fibres.

Growth Rings. Absent.

PHYSICAL PROPERTIES

Sapwood yellow-brown and distinct from heartwood; about 4-1 inch wide.

Heartwood red-brown when freshly sawn, later darkening to dark brown.

The timber is very hard and very heavy, with an average density of 64 lb.

per cu. ft. (range: 58-67 lb. per cu. ft.) air-dry.

Grain straight or shallowly interlocked; texture fine and even.

DISTINGUISHING FEATURES

(i) The timber of Cotylelobium and Vatica are so similar in appearance and

properties that the common name Resak is used for both. They may be recognised

as dipterocarps by the presence of axial intercellular canals and separated by the

presence of silica which is abundant in the ray cells of Cotylelobium but absent

from Vatica.

(ii) By the arrangement of its diffuse axial canals, this timber is readily dis-

tinguished from Kapur (Dryobalanops spp.), Selangan Batu and all Merants

(Shorea spp.), Selangan and Giam (Hopea spp.) and White Seraya (Parashorea

spp.). In those four genera, the axial canals are arranged in long tangential series.

(iii) Cotylelobium can be readily distinguished from Keruing (Dipterocarpus

spp.) and Mersawa (Anisoptera spp.) by its finer texture and higher density.

(iv) The numerous pores and presence of silica in Cotylelobium readily

distinguish the timber from Penyau (Upuna borneensis).

Anatomical Features of the Dipterocarp Timbers of Sarawak LO]

Dipterocarpus

FIELD CHARACTERISTICS

All thirty-two species of Dipterocarpus, which occur in Sarawak are known

as Keruing. It is one of the most characteristic trees of the lowland dipterocarp

forests, extending into the hills to an altitude of about 3,000 feet, although one

or two species are to be found in the peat swamps. Dipterocarpus usually occurs on

yellow sandy soils, narrow shale ridges, clay hillsides, alluvium or peat. Some

species are widespread but others tend to be localised. Dipterocarpus trees are

large, with a maximum height exceeding 200 feet and a girth of over 20 feet;

but more commonly they occur as trees of 150-180 feet in height with a girth

range of 12-16 feet, according to species. The trees have tall, straight, cylindrical

boles, with thick, rounded, buttresses. Bark surface is pale or dark grey to orange-

brown, sometimes pink-brown, flaked or shaggy, with prominent warty lenticels.

Dammar does not occur on unbroken bark surfaces but appears rapidly as clear

sticky drops on freshly cut surfaces.

WOOD ANATOMY

Pores/Vessels. Almost exclusively solitary. rarely with radial pairs, oblique

in arrangement, generally oval in outline, usually moderately few, medium-sized

to moderately large (size 4-5); intervascular pitting alternate, moderately coarse,

pits vestured; perforation plates simple; tyloses present.

Parenchyma. Paratracheal, as incomplete sheaths to the vessels; diffuse; form-

ing short tangential layers around the axial intercellular canals.

Rays. Uniseriate (fine) and 2-8 cells wide (medium-sized); low, less than 2

mm. in height; heterogeneous (Kribs’s types II and III, type I occasionally present)

with 1-3 marginal rows of square or upright cells; sheath cells sometimes present.

Vessel-ray Pitting. Simple, round to elongated, with large aperture.

Tracheids and Fibres. Vasicentric tracheids observed in most species.

Fibres with moderately conspicuous bordered pits, walls thick.

Intercellular Canals. Axial canals arranged singly and in short tangential

series of 2-7, diameter considerably less than that of the vessels; canals blocked

with whitish resinous contents.

Crystals. Wanting.

Silica. Abundant in ray cells of all species except D. borneensis in which it

is sparse.

Growth Rings. Absent.

PHYSICAL PROPERTIES

Sapwood grey-brown or yellow-brown and distinct from heartwood; 14-3

inches wide.

Heartwood pink-brown, purple-brown or red-brown, with a purple tinge,

darkening on exposure.

The timber of Dipterocarpus is hard but varies in density, ranging between

38-62 Ib. per cu. ft. air-dry, according to species. An average air-dry density of

45 lb. per cu. ft. may serve as an useful criterion to separate the timber into

two classes, the medium weight and the heavy weight.

102 Gardens’ Bulletin, Singapore — X XVII (1974)

Grain straight or shallowly interlocked: texture moderately coarse to coarse

and even.

DISTINGUISHING FEATURES

(i) The presence of axial intercellular canals, arranged singly and in short

tangential series of 2-7, can be used to distinguish the timber of Dipterocarpus

from that of non-dipterocarps and from that of most other dipterocarps.

(ii) Two features can be used to distinguish the timber of Dipterocarpus from

that of Anisoptera (Mersawa): (a) the timber of Dipterocarpus has a pink-brown

cast, that of Anisoptera a yellowish colour; (6) in Dipterocarpus the pores are

arranged in an oblique pattern but, in Anisoptera, are evenly distributed with no

apparent pattern. ,

(iii) Confusion with Resak (Cotylelobium spp. and Vatica spp.) or Penyau

(Upuna borneensis) is unlikely because of the much coarser texture of Diptero-

carpus.

Dryobalanops

FIELD CHARACTERISTICS

Six species of Dryobalanops (Kapur) are known to occur in Sarawak. They are

common trees of the lowland and hill forests but one species, D. rappa is to be

found in the peat swamps. They usually grow on shallow yellow sandy soils

derived from ferrugineous sandstone, white sand terraces, clay soils derived from

basalt, sandy loam soils or peat. All species develop into large or very large trees,

often growing to a height of over 200 feet and a maximum girth of 20 feet. The

trees have tall, straight and cylindrical boles, with long, concave, rather thick,

plank buttresses. Bark surface is of variable colour according to the species, evenly

or persistently flaked, with distinct lenticels on fresh surfaces. Dammar is not

exuded from cut surfaces, nor does it occur on the bole. Some species (D. aro-

matica, D. lanceolata) produce crystalline camphor in fissures within the heartwood.

WOOD ANATOMY

Pores/Vessels. Almost exclusively solitary, with a few radial multiples of 2-3,

oblique in arrangement, generally round to oval in shape, moderately numerous,

medium-sized to moderately large (size 4-5); intervascular pitting alternate,

moderately coarse, pits vestured; perforation plates simple; tyloses abundant.

Parenchyma. Paratracheal, as incomplete borders to the vessels, aliform, occa-

sionally confluent; diffuse and diffuse-in-aggregates; forming continuous tangential

layers around the axial intercellular canals.

Rays. Uniseriate (fine) and 2-6 cells wide (medium-sized); low, less than 2 mm.

in height; heterogeneous (Kribs’s types II and III, type I occasionally present in

D. beccarii) with 1-3 marginal rows of square or upright cells; sheath cells

occasionally present. Rays tend to be storied or arranged in echelon (except D.

rappa).

Vessel-ray Pitting. Simple, round to elongated, with large aperture.

Tracheids and Fibres. Fibres with moderately conspicuous bordered pits, walls

thick, Cells intermediate between vasicentric tracheids and fibre-tracheids often

occur in the immediate proximity of the vessels. |

Anatomical Features of the Dipterocarp Timbers of Sarawak 103

Intercellular Canals. Axial canals in continuous tangential rows, diameter

considerably less than that of the vessels; canals filled with whitish resinous con-

tents.

Crystals. Only in D. fusca and D. rappa in idioblasts and parenchyma, rhom-

boidal in shape.

Silica. Abundant in ray cells of all species except D. rappa in which silica

is sparse.

Growth Rings. Absent.

PHYSICAL PROPERTIES

Sapwood yellow-brown and clearly differentiated from heartwood; 1—24 inches

wide.

Heartwood red-brown or pink-brown, darkening on exposure.

The timber is hard and heavy; air-dry density of the hill species (Kapur Bukit)

averaging about 50 Ib. per cu. ft. (range: 45-55 lb. per cu. ft.) and the swamp species

(Kapur paya) being a little lighter, averaging only 45 lb. per cu. ft. (range: 42-48

Ib. per cu. ft.).

Grain straight or shallowly interlocked; texture moderately coarse and even;

freshly cut wood has a pronounced camphor-like odour but the smell tends to dis-

appear on seasoning.

DISTINGUISHING FEATURES

(i) The presence of axial intercellular canals arranged in long tangential series

provides a reliable diagnostic feature for distinguishing this timber from non-dip-

terocarp timbers and from Keruing (Dipterocarpus spp.), Mersawa (Anisoptera spp.),

Penyau (Upuna borneensis) and Resak (Cotylelobium spp. and Vatica spp.). In

those four dipterocarps, the axial canals are diffuse or in short tangential series.

(ii) The timber can be distinguished from all Merantis and Selangan Batu

(Shorea spp.), Selangan and Giam (Hopea spp.) and White Seraya (Parashorea

spp.) by its almost exclusively solitary pores and camphor-like odour when freshly

cut.

Hopea

FIELD CHARACTERISTICS

In this genus a considerable amount of taxonomic work has still to be done

but, already, about forty species of Hopea are recognised in Sarawak where they

form a very variable group in both the lowland and the hill forests. Within the

genus two groupings of species are commonly recognised: (i) Giam — a very hard,

heavy, durable timber closely related to the Selangan Batu group (Shorea spp.);

common species H. nutans, H. pentanervia, and H. semicuneata (ii) Selangan (Mer-

awan in West Malaysia) — timber less hard and heavy than Giam; common species

Ai. beccariana, H. nervosa and H. sangal. Some Hopea species are locally abundant,

usually growing on alluviums, calcareous shales, clay soils or yellow sandy soils

on ridges and hillsides, but a few are to be found on damp hillsides, near rivers,

shallow peat or swampy soils overlying white sand. The trees are usually small,

occasionally medium-sized, maximum height 90-120 feet, with a girth of 6-10 feet.

Boles are usually tapering, frequently branching low. Buttresses are usually thin,

104 Gardens’ Bulletin, Singapore — X XVII (1974)

sometimes thick; stilt roots sometimes present. Bark surface is at first smooth,

chocolate- and grey-mottled, hoop-marked, becoming cracked and flaked or fis-

sured. The Giam trees usually have a pale yellowish, the others a clear white

dammar, sometimes in stalactitic form.

WOOD ANATOMY

Pores / Vessels. Solitary and sometimes in radial multiples of 2—4, occasionally

in small clusters, round in shape, moderately numerous to numerous, evenly distri-

buted, moderately small to medium-sized (size 3-4); intervascular pitting alternate,

moderately fine, pits vestured; perforation plates simple; tyloses abundant.

Parenchyma. Paratracheal, usually as incomplete borders to the vessels, aliform,

occasionally confluent; apotracheal, in irregularly spaced, narrow, terminal bands

or in short, uniseriate lines between rays; forming continuous tangential layers en-

closing the axial intercellular canals.

Rays. Uniseriate (fine) and 2-5 cells wide (medium-sized), occasionally up to

7 cells wide; low, less than 2 mm. in height; heterogeneous (Kribs’s types If and

JI, sometimes type I) with 1-3 marginal rows of square or upright cells; sheath

cells occasionally present.

Vessel-ray Pitting. Simple, round to elongated, with large aperture.

Tracheids and Fibres. Vasicentric tracheids present in all species, the cells

irregular in shape and with conspicuous bordered pits. Fibres with indistinctly

bordered pits, walls moderately thin in the Selangan group but very thick in Giam.

Intercellular Canalis. Only axial canals present, in continuous tangential rows,

variable in size, usually equal to or smaller than that of the vessels but sometimes

larger; canals usually blocked with chalky deposits.

Crystals. Always present, rhomboidal in shape, in ray cells or parenchyma or

both, occasionally chambered or sub-divided, sometimes in idioblasts.

Silica. Wanting.

Growth Rings. Absent.

PHYSICAL PROPERTIES

Sapwood lighter in colour than the heartwood, about 1-2 inches wide, often

with a greyish tinge, poorly differentiated from the heartwood.

Heartwood yellow or light yellow-brown when freshly cut, darkening to red-

brown on exposure.

Timber weighing over 60 Ib. per cu. ft. air-dry, i.e. Giam, is hard and durable:

the lighter wood (40-60 Ib. per cu. ft. air-dry), i.e. Selangan, moderately hard, less

durable.

Grain, usually shallowly interlocked, sometimes wavy or spiral; texture fine

or moderately fine, even.

DISTINGUISHING FEATURES

(i) The presence of axial intercellular canals in continuous tangential series

separates the timber from non-dipterocarps.

(ii) The same feature serves to distinguish Hopea from Keruing (Dipterocarpus

spp.), Mersawa (Anisoptera spp.), Resak (Cotylelobium spp. and Vatica spp.) and

Anatomical Features of the Dipterocarp Timbers of Sarawak 105

Penyau (Upuna borneensis), all of which have axial canals in a diffuse arrangement

or in short tangential series.

(iii) Giam can be distinguished from Selangan Batu (Shorea spp.) by its finer

texture.

(iv) Selangan resembles Yellow Meranti (Shorea spp.), but can be distinguished

from it by the absence of radial canals in its rays and the even distribution of its

pores. In Yellow Meranti, pores have a radial arrangement.

Parashorea

FIELD CHARACTERISTICS

Four species of Parashorea (White Seraya) are known to occur in Sarawak,

usually growing on damp sticky clay on hillsides near rivers and streams or on

well drained clay spurs and ridges. The genus has not been recorded from west

of the Batang Lupar. All species grow to large trees, maximum height 150-180

feet, with a girth of about 12 feet. The trees have tall, straight and cylindrical boles,

with large, rounded and slightly concave buttresses. Bark surface is distinctly

mauve-grey to purplish, with narrow, shallow fissures, broad, smooth or flaking

flat ridges, and numerous conspicuous large pale corky lenticels. Dammar is creamy

to yellowish, exuding from rows of resin canals on cut surfaces.

WOOD ANATOMY

Pores/Vessels. Solitary and sometimes in radial multiples of 2-4, generally

round to oval, moderately few. medium-sized to very large (size 4-6), mostly

moderately large (size 5); intervascular pitting alternate, moderately coarse, pits

vestured; perforation plates simple; tyloses present.

Parenchyma. Paratracheal, vasicentric, aliform, occasionally confluent; diffuse

and diffuse-in-aggregates; forming continuous tangential layers surrounding the

axial intercellular canals.

Rays. Uniseriate (fine) and 2~7 cells wide (medium-sized); low, less than 2 mm.

in height; heterogeneous (Kribs’s type III, occasionally type I1), commonly with 1

marginal row of square or upright cells; sheath cells sometimes present.

Vessel-ray Pitting. Simple. round, with large aperture. |

Tracheids and Fibres. Vasicentric tracheids always present, the cells irregular in

shape and with conspicuous bordered pits. Fibres with simple pits, walls thin in

P. macrophylla and P. malaanonan but thick in P. parvifolia and P. smythiesii.

Intercellular Canals

(i), Axial canals in continuous tangential rows, often rather widely spaced,

diameter considerably less than that of the vessels; canals usually filled with chalky

or yellow-white contents.

(ii) Radial canals absent from all the Sarawak specimens but observed in P.

smythiesii of Sabah origin.

Crystals. Always present, usually rhomboidal, in parenchyma, chambered,

usually in long strands; in ray cells, sometimes more than one per cell.

Silica. Wanting.

Growth Rings. Absent.

106 Gardens’ Bulletin, Singapore — X XVII (1974)

PHYSICAL PROPERTIES

Sapwood paler in colour than the heartwood, usually 2-3 inches wide, com-

monly stained with fungus; fairly distinct from the heartwood.

Heartwood pinkish or straw-coloured, darkening to light brown on exposure,

with a distinctive brownish tinge.

The timber of Parashorea may be divided into two classes, based on its density:

(i) P. malaanonan and P. macrophylla are of medium weight, averaging between

35 and 42 lb. per cu. ft. air-dry; (ii) P. parvifolia and P. smythiesiti are heavy

timbers, averaging between 48 and 52 Ib. per cu. ft. air-dry. Both groups of timber

are moderately hard.

Grain interlocked: texture rather coarse, even.

DISTINGUISHING FEATURES

(i) The presence of axial intercellular canals distinguishes the timber of Para-

shorea from non-dipterocarps. :

(ii) The presence of axial canals in long tangential series serves to separate

the timber of Parashorea from Keruing (Dipterocarpus spp.), Mersawa (Anisoptera

spp.), Resak (Cotylelobium spp. and Vatica spp.) and Penyau (Upuna borneensis).

(iii) In Parashorea, the pores are often in radial multiples of 2-4, an arrange-

ment quite different from Kapur (Dryobalanops spp.) in which the pores are almost

exclusively solitary.

(iv) The timber is unlikely to be confused with Selangan and Giam (Hopea

spp.) since those timbers are of a fine texture. "

(v) Confusion with Shorea —see under Shorea.

Shorea

This is the largest genus within the family Dipterocarpaceae with at least one

hundred and twenty species known to occur in Sarawak. All are trees capable

of yielding commercial timbers but some are of no commercial importance because

of inaccessibility or scarcity. Symington (1934) pointed out that, in West Malaysia,

groups of species within the genus could usually be recognised in the field from

bark and slash characters combined with the gross timber morphology. This group-

ing of species corresponds closely with local practice in the Sarawak timber industry.

In Sarawak, the accepted grouping of species within the genus generally cor-

responds to anatomical features or timber properties and the differences between

the groups are both constant and distinct. The four groupings recognised in Sarawak

are: —

I THe RED MERANTI GROUP

Sub-group 1. Light Red Meranti

Sub-group 2. Dark Red Meranti

Sub-group 3. Red Selangan (Red Balau)

If THE SELANGAN BaTU (BALAU) GROUP

If THE WHITE MERANTI GROUP

IV THE YELLOW MERANTI GROUP

_—————o“e ©

Anatomical Features of the Dipterocarp Timbers of Sarawak 107

FIELD CHARACTERISTICS

The Red Meranti Group

Botanists recognise sixty-two species of Shorea in the Red Meranti group which

is the most important group of timber trees in Sarawak. Red Merantis are to be

found in all types of forest, except the mangroves, from sea-level to about 3,000

feet altitude. All species develop into medium- to large-sized trees with, according

to species, a maximum height of 120 feet to over 200 feet. Trees with a girth of 24

feet are not unknown, although 12-14 feet is more common. The trees usually have

clean, straight boles, with large, thick, rounded buttresses. The bark surface is very

variable, but the inner bark is red, pink-brown, orange-brown, rarely coffee coloured,

and usually fibrous and relatively soft. Dammar exudations are commonly present,

of variable colour when fresh but oxidising from cream to yellow on exposure.

The Selangan Batu (Balau) Group

Twenty-five species of Selangan Batu (in West Malaysia under the name of

Balau) are known to occur in Sarawak. Some are abundant locally, usually on

well-drained sandy soils. They are common in the lowland forests but do not

occur in the swamp and mangrove forests. The trees are usually small to medium-

sized, maximum height 100-150 feet according to species, but giants of about

220 feet with a girth exceeding 20 feet are occasionally found. Boles are variable.

Buttresses are prominent, narrow and sharp. Bark surface is longitudinally cracked,

thinly or shaggily flaked; dull grey-brown, yellow-brown, tawny or chocolate and

indistinctly lenticellate. Dammar is frequently present as pale yellowish to brown

smears on the bole.

The White Meranti Group

Of the eight species of White Meranti known to occur in Sarawak, seven

are fairly common in the lowland forests and one in the heath forests; they are

not found in the peat swamp forests. The trees usually grow on well-drained or

sandy clay soils on ridges and hillsides, on relatively fertile soils derived from

igneous rocks, or on shallow podsolic sandy soils. Most species develop into

medium- to large-sized trees, maximum height 150-165 feet, with a girth of about

12-15 feet. The trees usually have tall, straight, cylindrical boles except S.

ochracea which is frequently twisted or crooked. Buttresses are prominent, thick

and rounded. Bark surface is pale or dark, irregularly fissured. Dammar exuda-

tion is pale yellow.

The Yellow Meranti Group

Twenty-five species of this group are found in Sarawak, occurring from

lowland to hill forests in a wide variety of habitats. Yellow Meranti is usually

found on well-drained soils or on soils derived from calcareous shales or igneous

rocks, but never on deep peats. With some exceptions, they are large trees reach-

ing a maximum height of over 200 feet with a maximum girth in excess of 25 feet.

Boles are variable. Buttresses are small or prominent, of medium thickness, nar-

rowly rounded. Bark surface is at first smooth, chocolate and grey dappled,

becoming pale or dark tawny-brown, rarely chocolate-brown, irregularly and

longitudinally cracked or flaked, closely and deeply fissured. Dammar is charac-

teristically dull, dark grey-brown to black, readily exuding on cut surfaces:

stalactitic exudations are usually abundant on the bole.

108 Gardens’ Bulletin, Singapore — X XVII (1974)

WOOD ANATOMY

Pores /Vessels. Solitary and sometimes in radial multiples of 2-4, in charac-

teristic groups of 3-5 in the White Meranti; generally round to oval in shape,

moderately few to moderately numerous, usually moderately small to medium-

sized (size 3-4) in Light Red Meranti, Selangan Batu and Yellow Meranti, but in

Dark Red Meranti, Red Selangan and White Meranti, medium-sized to moderately

large (size 4-5); intervascular pitting alternate, moderately coarse, pits vestured;

perforation plates simple; tyloses usually abundant.

Parenchyma. Paratracheal, vasicentric, aliform, occasionally confluent; dif-

fuse and diffuse-in-aggregates; forming continuous tangential layers around the

axial intercellular canals.

Rays. Uniseriate (fine) and 2-8 cells wide (medium-sized); low, less than 2

mm. in height; heterogeneous (Kribs’s types IT and III) with 1-3 marginal rows of

square or upright cells; sheath cells occasionally present.

Vessel-ray Pitting. Simple, round, with large aperture.

Tracheids and Fibres. Vasicentric tracheids always present, the cells irregular

in shape and with conspicuous bordered pits. Fibres with simple pits, walls thin

in Light Red Meranti, White Meranti and Yellow Meranti; thick in Dark Red

Meranti, Red Selangan and some species of Yellow Meranti, but in Selangan

Batu, very thick.

Intercellular Canals

(i) Axial canals in continuous tangential rows, diameter considerably less than

that of the vessels except in Dark Red Meranti where the canals are often as

large as the vessels; canals usually blocked with chalky or yellowish deposits.

(ii) Radial canals present in all species of Yellow Meranti and the following

species of Red Meranti: —Shorea leprosula, S. ovata, S. parvistipulata, S. scabrida

and §. teysmanniana.

Crystals. Occasionally present in all groups except White Meranti; in idio-

blasts, usually rhomboidal in shape, in parenchyma and ray cells.

Silica. Present in White Meranti in ray cells, but totally absent from other

groups.

Growth Rings. Absent.

PHYSICAL PROPERTIES

Red Meranti Group

Sapwood distinct, light yellow-brown or light grey-brown; about 1-2 inches

wide.

Heartwood red or red-brown in Light Red Meranti; in Red Selangan and

Dark Red Meranti, deep purple-red or dark red-brown, with prominent white or

yeilow-white lines of resin canals on all surfaces.

The timber of Light Red Meranti is soft to moderately hard, light to medium

in weight averaging about 35 Ib. per cu. ft. (range: 25-45 Ib. per cu. ft.) air-dry;

Dark Red Meranti and Red Selangan are harder and heavier, with the average

air-dry densities of 45 lb. per cu. ft. (range: 42-55 Ib. per cu. ft.) and 55 lb. per

cu. ft. (range: 48-60 Ib. per cu. ft.) respectively.

Grain straight or shallowly interlocked; texture even, rather coarse in Dark

Red Meranti but finer in Red Selangan and Light Red Meranti.

Anatomical Features of the Dipterocarp Timbers of Sarawak 109

Selangan Batu (Balau) Group

Sapwood fairly distinct from, and lighter in colour than the heartwood; about

1-2 inches wide.

Heartwood yellow-brown, grey-brown or brown, on exposure darkening to

dark brown or dark red-brown.

The timber is very hard and heavy, averaging about 63 Ib. per cu. ft.

(range: 58-70 lb. per cu. ft.) air-dry.

Grain usually interlocked and wavy; texture moderately fine and even.

White Meranti Group

Sapwood lighter in colour than the heartwood and fairly distinct from it;

about 2 inches wide.

Heartwood almost white when freshly sawn, later darkening to yellow-brown

or buff-coloured.

The timber is moderately hard and medium-heavy, with an average air-dry

density of 41 lb. per cu. ft. (range: 36-47 Ib. per cu. ft.).

Grain usually shallowly interlocked; texture moderately coarse and even.

Yellow Meranti Group

Sapwood lighter in colour than the heartwood, with a greenish tinge and fairly

distinct from the heartwood; 1-24 inches wide.

Heartwood light yellow-brown, darkening on exposure.

The timber is moderately hard and medium-heavy, average weight 44 lb.

per cu. ft. (range: 36-56 lb. per cu. ft.) air-dry.

Grain usually shallowly interlocked and sometimes wavy: texture moderately

coarse and even.

Key to the timber groups in Shorea

1. (a) Silica present in the ray cells*; pores in characteristic groups of 3-5; wood

CEES, ater ee eee me Pare, er ee, ML RYE WHITE MERANTI

eM SCM. WOO MOU AS AWOME 3.7.0... cease cress eas cuies ssn vesens uncer» gedoe ad

2. (a) Wood over 60 Ib. per cu. ft. air-dry; wood very hard ......... SELANGAN BATU

(b) Wood below 60 Ib. per cu. ft. air-dry; wood never hard ................002.002. 3

3. (a) Wood yellow; ray tissue bright yellow; radial intercellular canals always

ode SUS arg aac ai 9 Seale een oe YELLOW MERANTI

(b) Wood red or red-brown; ray tissue not bright yellow; radial intercellular

canaispresent onlyan aifewsspecies of Red Meranti .,....,.-.¢. 26 .020....' -

4. (a) Wood parenchyma prominent, tending to short lines between rays; axial

intercellular canals -prominent:on all surfaces .. ccs cc... vews ices sedis. .cvessenis 5

(b) Wood parenchyma not prominent; axial intercellular canals distinct on all

SORTA COR OE <UVL WEMEOIMUIBO RC fet ee secchcgs « «carpe y pected LicguT RED MERANTI

fe) cend-cut elistening..or. waxyet iy iPM. AE. a RED SELANGAN

en Bar -cumiliie cient om) Stilw. -elibo 8 oe. fend): DaRK RED MERANTI

*Microscopic feature.

110 Gardens’ Bulletin, Singapore — X XVII (1974)

DISTINGUISHING FEATURES

(i) The presence of axial intercellular canals arranged in long tangential series

precludes the possibility of confusing the timber of Shorea with non-dipterocarp

timbers and with Keruing (Dipterocarpus spp.), Mersawa (Anisoptera spp.), Penyau

(Upuna borneensis) and Resak (Cotylelobium spp. and Vatica spp.). In those four

Dipterocarps, the axial canals are diffuse or in short tangential series.

(ii) The timber of Shorea, with pores in radial multiples of 24 is readily

distinguished from that of Kapur (Dryobalanops spp.) in which the pores are

almost exclusively solitary. Further, unlike Shorea which is odourless, Kapur has

a camphor-like odour when freshly cut.

(iii) Superficially the wood of Hopea species (Selangan and Giam) may

resemble that of Yellow Meranti and Selangan Batu (Shorea spp.), especially in

colour. The absence of any radial arrangement in the vessels and of radial canals

in the rays can be used to separate Selangan from Yellow Meranti. Although

both Giam and Selangan Batu are very heavy timbers, Giam may be recognised

by its finer texture.

(iv) When White Seraya (Parashorea spp.) is confused with Red or White

Meranti, it may sometimes be distinguished by its coarser texture and distinctive

brownish tinge. In doubtful cases, a microscopic examination of White Seraya

will reveal distinctive, chambered, crystalliferous parenchyma in long strands;

these strands do not occur in Red or White Meranti although silica is to be

found in the ray cells of White Meranti and Red Meranti may have some crystal-

liferous parenchyma.

Upuna

FIELD CHARACTERISTICS

Upuna is a monotypic genus with U. borneensis as the only species. U. bor-

neensis, locally known as Penyau, is scattered through the lowland forests, on well

drained ridges and deep yellow sandy clay soils. In Sarawak, it is widely distri-

buted but never abundant. This species develops into a large tree, attaining a

height of 160 feet and a girth of 18 feet. The tree has a tall, straight, cylindrical

and well-shaped bole, with buttresses varying from small and stout to very large.

Bark surface is dark purple-brown to chocolate, irregularly scaly. Dammar is pale

yellow.

WOOD ANATOMY

Pores/Vessels. Exclusively solitary, generally round to oval in outline,

moderately few, moderately small to medium-sized (size 3—4); intervascular pitting

alternate, moderately coarse, pits vestured; perforation plates simple; tyloses present.

Parenchyma. Paratracheal, as incomplete borders to the vessels; apotracheal,

diffuse and diffuse-in-aggregates; around the axial intercellular canals, occasionally

forming more or less continuous tangential layers.

Rays. Uniseriate (fine) and 2-7 cells wide (medium-sized); low, less than 2

mm. in height; heterogeneous (Kribs’s types II and III) with 1-3 marginal rows

of square or upright cells; sheath cells occasionally present.

Anatomical Features of the Dipterocarp Timbers of Sarawak 111

Vessel-ray Pitting. Simple, round to elongated, with large aperture.

Tracheids and Fibres. Fibres with moderately conspicuous bordered pits,

walls very thick.

Intercellular Canals. Axial canals scattered singly and in pairs throughout the

fibres, sometimes in more or less continuous tangential rows, diameter usually

smaller than that of the vessels.

Crystals. Wanting.

Silica. Wanting.

Growth Rings. Absent.

PHYSICAL PROPERTIES

Sapwood pale to light brown and distinct from heartwood; about }—1 inch wide.

Heartwood brown, later darkening to dark brown.

The timber is very hard and very heavy, with an average air-dry density of

65 Ib. per cu. ft. (range: 60-70 lb. per cu. ft.).

Grain shallowly interlocked and wavy; texture fine and even.

DISTINGUISHING FEATURES

(i) The presence of axial intercellular canals, arranged singly and sometimes

in more or less continuous tangential series distinguishes the timber from non-

dipterocarps and from most other dipterocarps.

(ii) The timber of Upuna borneensis resembles Resak (Cotylelobium spp. and

Vatica spp.), but the comparatively fewer pores in Upuna borneensis preclude the

possibility of confusing it with Resak. In Resak, the pores are numerous to very

numerous. Upuna borneensis which has no silica is readily distinguished from

Cotylelobium spp.

(iii) The timber can be separated from Mersawa (Anisoptera spp.) and

Keruing (Dipterocarpus spp.) by its higher density and much finer texture.

Vatica

FIELD CHARACTERISTICS

A widely distributed genus consisting of about twenty-eight species in Sarawak.

A few species occur in the peat swamps, and the remainder in the lowland and

hill forests. All species of Vatica, as well as Cotylelobium, are commonly known

as Resak. Vatica is usually found on clay soils on hillsides and ridges, white

sandy soils on terraces, rentzinas on limestone hills, peat soils, clay alluviums or on

shale, basalt and dacite slopes at higher altitudes. The trees are mostly rather

small, maximum height between 80 and 100 feet, with a girth of about 3-5 feet.

Boles are frequently sinuate. Buttresses are thick, rounded, concave and usually

small. Bark surface is usually grey-mottled, smooth, hoop-marked, becoming

patchily flaked and occasionally scroll-marked in large trees. Dammar is rare on

bark surfaces but clear when it exudes from cut surfaces.

112 Gardens’ Bulletin, Singapore — XXVIII (1974)

WOOD ANATOMY

Pores/Vessels. Mostly solitary with occasional radial, tangential and oblique

pairs, evenly distributed, generally round, numerous to very numerous, moderately

small to medium-sized (size 3-4); intervascular pitting alternate, rare, tending to

opposite or scalariform in V. granulata, V. havilandii, V. mangachapoi, V. sara- —

wakensis and V. umbonata, pits vestured; perforation plates simple; tyloses present.

Parenchyma. Paratracheal, as incomplete borders to the vessels, sometimes

abaxially aliform, confluent; diffuse and diffuse-in-aggregates; around the axial

intercellular canals.

Rays. Uniseriate (fine) and 2-10 cells wide (medium-sized); usually low, but

sometimes up to 2 mm. in height; heterogeneous (Kribs’s types II and III) with

1-3 marginal rows of square or upright cells; sheath cells occasionally present.

Vessel-ray Pitting. Simple, round to elongated, with large aperture.

Tracheids and Fibres. Fibres with moderately conspicuous bordered pits, walls

very thick.

Intercellular Canals. Axial canals scattered singly and sometimes in pairs

throughout the fibres, diameter less than that of the larger vessels but as large as

some of the smaller vessels; Canals usually packed with chalky deposits.

Crystals. Sometimes present, usually rhomboidal in shape, in chambered paren-

chyma or ray cells or both, rarely more than one per cell.

Silica. Wanting.

Growth Rings. Absent.

PHYSICAL PROPERTIES

Sapwood pale brown or yellow-brown, usually !-3 inches wide, sharply dif-

ferentiated from the heartwood when green, but less so when dry.

Heartwood red-brown, with a pink tinge, darkening to deep or dark reddish

brown.

The timber of Vatica varies in hardness, density and durability. The local

people often distinguish Resak Batu, the harder, heavier and more durable species,

from Resak Bunga which is lighter and of lower durability. An average air-dry

density of 60 Ib. per cu. ft. may serve as an useful criterion to separate the two

classes. However, it should be noted that the timber of a given species will be

either Resak Batu or Resak Bunga according to density of individual trees, although

V. mangachapoi and Cotylelobium usually produce timber of only the Resak

Batu Class.

DISTINGUISHING FEATURES

(i) The timber of Vatica resembles that of Penyau (Upuna borneensis) from

which it can usually be distinguished by its more numerous and smaller pores.

In Upuna the pores are exclusively solitary; with Vatica this is not so.

(ii) Separation from other timbers — see under Cotylelobium.

TABLE 1 SUMMARY OF ANATOMICAL CHARACTERISTICS OF 9 GENERA OF THE DIPTEROCARPACEAE OCCURRING IN SARAWAK

Vessels Parenchyma Rays Intercellular Canals Crystals

Inter- ; 2

vascular Paratracheal Apotracheal Hiclerorencous, o oe 3

aes ribs’s Types « =

z Pitting vo 5 7) >

Genus is 2 Ey ea ere Diameter iS a

And g iS ae| 3 | es ele

Group e | & P g = o&| 2 | 3e = | £ | 6

jaa SE] S = 5 a S & 3 8 es = Es a) c

8 o) a) < 3 |B aS Z } s2| é ‘eo 8 é 5 Fes

ts) n _ el q o a es io on a Zz 2 aw lal <a4\|8 Fa I 2 & 2 () 2

B & ge |ee| S |e 58 |e el | i | im fim | OC as} se VS P 16%le [68 el She | lh Ay Ee eg

2 3 | gs |e) 6 lBol a | Ea Tee) o [ese es [oo | Ge lee e. Os Neate co eb so aes

SG | BE LBB 2 eel Bye eel] ay ie Sens ee cueat| acs sele | els les |ae | 2 | s

K & fais) 2 K v= BSI (6) =| 8 = 2a a as) a |e 22 |S0 g = a & 2

3 | Se (22s (ea & |S lee| @ | eal as asa leo pee Vag SS isos ele Ma la |) 6 |S | &

, (1)

Anisoptera 4 | 88—96 | 192-226 | + + ar + + + + ar = |1=7 |) = + + = ar — ae llse) |] ap + +

Cotylelobium .. 3 95—98 | 124—137 oP ap qr + + oP ap (+) = 1—8 | (—) oF ar (+) + = + = + + + — — — }

: (2)

Dipterocarpus .. | 30 | 87—97 | 187-264) + + + + aF = + = = || 8 |) Ge) |] 46 ae | (Ge) |) aro) GP) | ae = + + — - - —- - -- —

(1) (2)

Dryobalanops .. 6 | 89—95 | 180—232 | + + + + + + = = — |1-6|()}] + NG) |) -E _ + en = + E)are) = = ats

Hopea 5.0 28 §2—84 | 102—172 + + + oP + + + ap + 1—5 | (4) + + (+) aP + = oF = + + — (+) ar IP (+) = =

Parashorea 0'6 4 78—86 | 239—307 + a + + + AP ar = = = 7/ = (+) dP (4) oF + = ar I ar = = = + te + (+) =

Shorea

—Red Meranti 60 50—89 | 179—275 + + + oP + ap + (-) = 1—7 = + + (+) + + = ap = + ©) | © | G) | G@ | G) | ©) ae =

—Selangan Batu 24 49—82 | 134—217 + + oF + + + + 1—7 + + (—) + + = + = + = = (+) | G) (+)

—White Meranti 7 $3—71 | 153—254 + + ar + oP Ste + 1-6 + AF = oF + = te = t = re ne ar

—Yellow Meranti| 22 | 46—66 | 146—233 | + + + + + + + - — |}1-8} — + + |}()/] + =F = + — -- — + = 165) | (GD) | (Go) i (ay

Upuna oral) oat 6B | i@=oy |} es || ee se | ae eee i= a fa fs Pe fe eee es) eee ae | a Se SSS

Vatica -. | 21 | 47-90 | 88-147} + a cA a A. ae + = = | 1—=10) = + + |G] + — + = + + + = = | Gd) Ge) | Gey | C) |) =

+ = Present.

— = Absent.

(+) Usually present, majority of species conforming.

(—) Usually absent, present in some species.

(1) & (2) See text under generic description.

113

Anatomical Features of the Dipterocarp Timbers of Sarawak

115

Taste 2 THE DENSITY OF THE DIPTEROCARP TIMBERS OF SARAWAK

Botanical Name

Anisoptera (4 spp.)

—grossivenia V. SI.

—jlaevis Rid. . .

—marginata Korth.

—reticulata Ashton

Cotylelobium (3 spp.)

—burckii Heim.

—malayanum V. SI. *.

—melanoxylon (Hook f.)

Pierre ae

Dipterocarpus (30 spp.)

—acutangulus Vesque

—applanatus V. SI.

—apterus Foxw.

—borneensis V. Sl.

—caudiferus Merr.

—confertus V. SI.

—conformis V. SI.

—coriaceus V. Sl.

—costulatus V. SI.

—crinitus Dyer

—cuspidatus Ashton

—eurynchus Miq.

—exalatus V. Sl.

—fagineus Vesque

—geniculatus Vesque

—globosus Vesque

—gracilis Bl. . .

—humeratus V. SI.

—lowii Hook f.

—muundus V. Sl.

—nudus V. SI.

—oblongifolius BI.

—pachyphyllus Meijer

—palembanicus V. Sl.

—penangianus Foxw.

—rigidus Ridl.

—sarawakensis V. Sl.

—stellatus Vesque

—tempehes V. Sl.

—verrucosus V. Sl.

Dryobalanops (6 spp.)

—aromatica Gaertn. f.

—beccarii Dyer

—fusca V. SI.

—lanceolata Burck

—oblongifolia Dyer

—rappa Becc.

Hopea (28 spp.)

—aequalis Ashton

—altocollina Ashton

—andersoni Ashton

—argentea Meijer

—beccariana Burck

—bracteata Burck

Density —\b. per cu. ft.

Local or Vernacular — - 12% iy

Name

is nat Mean Range

Mersawa |

Mersawa kunyit 6 49.2 | 44.4—56.2

Mersawa durian 4 44.8 | 40.5—50.5

- | Mersawa paya .. 5 45.3) |. 41.5—49.7

| — 1 48.4 —

Resak

| Resak durian 5 66.6 | 62.0—71.1

Resak bukit 3 67.3 63.7—72.4

Resak hitam 5 58.3 55.9—61.0

Keruing

Keruing beludu 6 51.0 | 49.3—56.7

| Keruing arong . - 6 48.3 | 43.1—51.8

Keruing latek 5 39.3 | 38.2—39.9

Keruing sindor . - 5 50.5 47.4—54.7

Keruing puteh .. 6 44.1 41.8—48.7

Keruing kobis - . 5 53.3. | 48.2—58.0

Keruing beludu kuning 3 49.0 44.8—54.0

Keruing paya 5 42.8 40.7—45.1

_ Keruing kipas 2 50.5 | 46.9—54.0

| Keruing mempelas 6 59.2 54.9—61.8

Keruing runching 4 50:1 48.0—53.7

Keruing baran .. 2 46.1 | 44.6—47.6

Keruing kuntum puteh 5 41.8 | 38.7—45.0

Keruing pipit 1 50.2 —

Keruing kerubong 4 47.9 45.4—50.9

| Keruing buah bulat 5 52.0 47.8—55.8

-- | Keruing kesat 5 48.1 43.4—54.0

Keruing latek bukit 2 48.7 45.5—51.8

Keruing sol 3 49.3 |, 44.1352

— a 49.7 49.5—49.9

Keruing lichin . . 5 559 . | 605—567

Keruing neram > 47.8 40.6—51.2

Keruing sol padi 5 52.2 47.4—55.2

Keruing ternek 5 45.8 42.2—48.8

Keruing gasing 5 55.8 | 53.3—58.0

Keruing utap 5 49.9 43.7—53.7

Keruing layang. - 5 49.4 «|. 45.1-—55.2

Keruing daun nipis 5 56.1 | 49.3—60.0

Keruing tepayan = 41.0 | 37.7—45.9

Keruing merah 5 53.6 52.4—56.0

Kapur

Kapur peringgi . . 5 55.0 53.5—56.2

Kapur bukit 5 49.0 46.9—50.0

Kapur empedu 5 54.0 S2:7=35:5

Kapur paji 5 45.0 42.1—48.3

Kapur kelansau 5 54.0 47.4—59.9

Kapur paya 5 45.0 40.1—48.4

Giam, Selangan(merawan), etc.

= M ii Maw + ast

Luis gunong.. soetyell | 2 49.0 | 46.4—51.6

Luis somit ca PS 59.1- ~| 56.4—62.4

Luis timbul 2 58.6 56.8 — 60.4

Selangan penak 4 SAF 56.2—59.3

Merawan padi . - 5 42.3

| 39.7—45.8

116

Gardens’ Bulletin, Singapore — X XVII (1974)

TaBLE 2 THE DENSITY OF THE DIPTEROCARP TIMBERS OF SARAWAK—continued

Dense ate per cu. ft.

Botanical Name Local or Vernacular a fat 12% me)

Name | Trees

Mean Range

—centipeda Ashton — 4 47.2 40.5 —50.9

-cernua T. et B. — 3 58.0 55.6—62.3

— dasyrrhachis VS. — 1 70.5 pi

—dryobalanoides Miq. Mata kuching hitam 4 48.6 AAD 53,3

—dyeri Heim Merawan palit . . 4 52.8 50.9 —57.2

—enicosanthoides Ashton Luis selukai 1 56.6 adit

—fluvialis Ashton . | Merawan Ayer .. 2 69.1 68.6 —69.6

—griffithii Kurze Luis jantan ; 1 67.2 -

—latifolia Sym. Merawan daun bulat 2 58.7 54.0 —63.4

—megacarpa Ashton = 1 60.5 —

—mesucides Ashton = 1 59.3 eis

—micrantha Hook f. Merawan kerangas 2 47.4 46.6—48.1

—nervosa King Merawan Jangkang 3 48.8 43.3 —54.9

—nutans Ridl. Giam 1 63.0 —

—pachycarpa (Heim.) Sym. Merkoyong a 5 52.2 47.8—56.1

—pedicellata (Brandis) Sym. | Mata kuching aaa 1 50.5 mes 3

—pentanervia Sym. .. | Mang i 72.9 68.5—76.1

—pterygota Ashton — 1 62.7 ts

—sangal Korth. Gagil 2 46.9 43.3 —50.5

—semicuneata Sym. Sama rupa chengal 2 57.3 55.3 —59.3

—tenuinervula Ashton — 1 63.7 bes

—treubii Heim Merawan daun tebal 2 58.6 58.5—58.7

Parashorea (4 spp.) WHITE SERAYA

—macrophylla Wyatt- -Smith

ex Ashton Peran ‘| 42.3 38.4—45.4

—malaanonan (Blanco)

Mer. .. .. | Urat mata 4 35.8 32.0—42.4

—parvifolia ada ae

ex Ashton Urat mata bukit 5 48.8 46.4—54.5

—smythiesii Wyatt- -Smith

ex Ashton Meruyun 6 51.9 47.4—54.0

Shorea (56 spp.) RED MERANTI GROUP

—acuta Ashton .. | Meranti kawang tikus 1 34.3 Tz

—albida Sym. .. | Alan bunga 5 28.3 25 MSL

Alan | 51.0 41.8—57.0

—almon Foxw. es = 5 34.4 31.3—38.1

—amplexicaulis Ashton .. | Meranti kawang pinang lichin 5 35.9 30.38 37.8

—andulensis Ashton Meranti daun puteh 1 43.1 Sie

—argentifolia Sym. Meranti binatoh 6 42.5 38.2 —45.6

—beccariana Burck Meranti langgai 6 40.1 36.4—43.1

—builata Ashton Meranti melechur e) 49.9 47.4—52.3

—carapae Ashton Abang uloh 1 42.2 Tae

—coriacea Burck Meranti tangkai panjang 5 50.6 46.7—54.3

—cristata Brandis Meranti kawang pinang 3 38.8 38.1 —40.3

—curtisii Dyer Seraya 5 50.1 48.2 —52.1

—dasyphylla Foxw. Meranti batu 3 32.8 29.8 —36.8

—elliptica Burck Meranti jang 3 43.6 40.1 —48.5

—fallax Meijer Engkabang layar 5 39.4 35.3 —43.2

—ferruginea Dyer ex Brandis Meranti menalit 6 42.0 34.5—47.4

—flaviflora Wood ex Ashton | Selangan merah bukit 5 47.0 43.7—51.1

—fiemmichii Sym. Meranti raya 1 48.0 eee

—jnaeguilateralis Sym. Semayor 5 56.4 53.1—60.6

—kunstleri King Damar laut merah 3 58.3 56.0—60.7

—leprosula Miq. Meranti tembaga 5 37.9 36,2—40.4

—leptoclados Sym. Meranti Majau . . 3 34.0 32.7—35.6

—macrantha Brandis Engkabang bungkus 4 51.5 46.1—55.5

Anatomical Features of the Dipterocarp Timbers of Sarawak 117

TasLe 2 THE DENSITY OF THE DIPTEROCARP TIMBERS OF SARAWAK—continued

| Density —Ib. per cu. ft.

| Number (at 12% m.c.)

Botanical Name Local or Vernacular of 9 aa

Name Trees |

Mean Range

—macrophylla (De Vr.) |

Ashton .. .. | Engkabang jantong 4 29.9 2A i — 35,3

—miacroptera Dyer .. | Meranti melantai 5 41.2 3/.1—44.4

—mecistopteryx Ridl. .. | Engkabang larai 5 35.4 31.8 —38.6

—monticola Ashton .. | Meranti gunong 3 49.0 45.4—54.1

—myrionerva Wood ex

Ashton .. .. | Meranti sepit undang 2 41.1 | 40.1—42.1

—ovalis (Korth) BI. .- | Meranti kepong > 3 549, ug oho — 41.9

—ovata Dyerex Brandis | Meranti sarang punai bukit 5 53.2 49.6—55.5

—pachyphylla Rid!. ex Sym. | Meranti kerukup - 3 54.8 50.4—57.6

—palembanica Miq. .. | Engkabang asu . . 3 42.3 | 40.0—46.7

—pailidifolia Ashton — — 1 a7 aa os

—parvifolia Dyer .. | Meranti sarang punai 6 35.6 w 32.6—40.6

—parvistipulata Heim .. | Engkabang pinang bersisek 6 45.5 41.2—49.3

—pauciflora King .. | Engkabang cheriak if 46.3 43.7—52.4

— pilosa Ashton .. | Kawang bulu 4 32.4 28.2—39.0

—pinanga Scheff. .- | Meranti langgai bukit 5 33.0 29.0 —38.7

—platycarpa Heim .. | Meranti paya 5 47.5 45.7—49.1

—platyclados V. SI. ex Foxw.| Meranti bukit 5 50.2 | 45.2—54.5

—praestans Ashion ae — 2 46.1 41.2—50.9

—pubistyla Ashton -. | Meranti bulu merah 1 46.8 --

—quadrinervis V. SI. -- | Meranti sudu si 6 41.2 33.8 —43.1

—retusa Meier -- | Meranti daun tumpul Sui 3 35.6 33.6 — 38.9

—rubella Ashton .. | Meranti laut puteh 4 39.7 36.1—43.3

—rubra Ashton .- | Meranti merah kesumba 7 39.2 35.9—45.6

—rugosa Heim -- | Meranti buaya hantu 5 46.5 | 42.2—52.8

—sagittata Ashton | Merantiluang . 3 37.0 35.6—39.0

—scaberrima Burck .. | Meranti paya bersisek 5 422 38.4—45.6

—scabrida Sym. -- | Meranti lop 6 45.5 41.9—48.i

—gslooteni Wood ex Ashto Meranti kepong kasar 5 51.1 45.9—55.9

—smithiana Sym. .- | Meranti rambai Be) 35.4 31.2 —39.3

—stenoptera Burck -- | Engkabang rusa ) 42.9 41.0—45.7

—teysmanniana Dyer -. | Meranti lilin : 5 42.3 39.1 —46.9

—uliginosa Foxw. .. | Meranti buaya - . cele S. 45.1 | 41.5—47.8

—venulosa Meijer .. | Meranti tangkai panjang padi 6 50.3: , |. 44.0—355

Shorea (22 spp.) .- | SELANGAN BATU(BALAU)GROUP

—atrinervosa Sym. .- | Selangan batu hitam Z, 61.8 60.6 —63.0

—biawak Ashton .. | Resak biawak 2 64.0 62.4—65.7

—brunnescens Ashton .. | Selangan batu tinteng 6 63.1 59.7—67.1

—crassa Ashton .- | Selangan batu daun tebal 4 66.9 63.0—72.4

—domatiosa Ashton .. | Selangan batu lobang idong 6 65.4 | 62.4—68.6

—exelliptica Meijer .-. | Selangan batu tembaga 1 64.9 —

—flava Meijer .. | Selangan batu kuning 5 67.3 64.9—70.5

—foxworthyi Sym. .- | Selangan batu bukit 3 64.5 63.7 —65.5

—geniculata Sym. ex Ashton| Upun penyau 1 61.2 oa

—glaucescens Meijer .. | Selangan batu daun nipis 6 57.5 53.0—63.0

—havilandii Brandis .. | Selangan batu pinang 3 70.3 68.6 —72.4

—hypoleuca Meijer .- | Selangan batu kelabu 2 | 64.0 58.7 —69.3

—inappendiculata Burck . . ee 2, Mien eta 65.3 — 68.6

—isopiera Ashton .. | Selangan batu main buluayam | 4 62.9 | 62.4—63.7

—ladiana Ashton .. | Selangan batu kilat 1 68.5 —

—laevis Ridl. .. | Kumus ; 6 61.5 59.3 —66.1

—lunduensis Ashton a — 1 .) Se —-

—maxwelliana King -. | Kumus hitam 5 65.3 ay Lo o35-09.9

—obscura Meijer .. | Selangan batu padi 4 66.5 64.3 —72.4

—scrobiculata Burck .. | Selangan batu zang seit oi 64.50 — ==

—seminis (De Vr.) V. SI. | Engkabang terendak This See, 58.8 56.8 — 63.0

—superba Sym. ex Wood. . | Selangan batu tulang ikan .. | 5 63.9 61.2—66.8

118

Gardens’ Bulletin, Singapore — X XVII (1974)

TABLE 2 THE esac aoe OF THE DIPTEROCARP TIMBERS OF woah oeg'c

Botanical Name

Local or Vernacular

Name

Shorea (7 spp.) -

—agami Ashton

—bracteolata Dyer

—cordata Ashton

—lamellata Foxw.

—ochracea Sym.

—resinosa Foxw.

—virescens Parijs

Shorea (20 spp.)

—acuminatissima Sym.

—angustifolia Ashton

—collaris V. SI.

—cuspidata Ashton

—dolichocarpa V. S!.

—faguetiana Heim

—faguetioides Ashton

—gibbosa Brandis

—hopeifolia (Heim.) Sym. |

—ijliasii Ashton

—induplicata V. Sl.

—longiflora (Brandis) Sym.

—macrobalanos Ashton .

—mujongensis Ashton

—multiflora (Burck) a

—obovoidea V. SI.

—patoiensis Ashton

—resina-nigra Foxw.

—subcylindrica V. Sl.

—xanthophylla Sym.

Upuna (1 sp.)

—borneensis Sym.

Vatica (21 spp.) . -

—badiifolia Ashton

—borneensis Burck

—brunigi Ashton

—coriacea Ashton

—dulitensis Sym.

—endertii V. SI.

—globosa Ashton

—granulata V. SI.

—havilandii Brandis

—maingayi Dyer

—mangachapoi Blanco

—miicrantha V. SI.

—nitens King

—oblongifolia Hook f.

—odorata (Griff) Sym.

—papuana Dyer

—rynchocarpa Ashton

—sarawakensis Heim

—umbonata (Hook f.) Burck

—venulosa BI.

—vinosa Ashton

WHITE MERANTI GROUP

Meranti puteh timbul

Meranti pa’ang. -

Meranti lapis

Raruk

Meranti belang .

Meranti sulang sulang

YELLOW MERANTI GROUP

Lun runching

Damar hitam bukit

Lun kelabu

Lun runching padi

Damar hitam gondol

Lun siput se

Damar hitam daun nipis

Lun gajah

Lun siput jantan

Lun siput daun besar

Lun puteh

Damar hitam paya

Engkabang melapi

Damar hitam

Damar hitam padi

Lun meranti

Lun barun

Penyau

Resak

Resak kemudi ..

Resak daun tebal

Resak tiong

Resak ranting bersisek

Resak degong

Resak lidi

Resak julong

Resak hijau

Resak daun panjang

Resak membangan

Resak runting kesat

Resak daun besar

Resak ayer

Resak letop

Resak tangkai ungu

Density — Ib. per cu. ft.

Number (at 12% m.c.)

of as

eee Mean Range

6 39.4 | 36.2—43.7

4 46.9 42.1—50.0

l 39.3 —

1 47.0 =

4 39.0 38.1—41.8

i 39.5 ae

4 36.3 32.5—39.4

3 45.0 34.3 —53.5

5 49.0 40.4—54.7

1 44.3 —

1 44.9 —

6 Bp) 50.5—59.9

6 41.0 34.3 —44.3

| 45.9 43.4—48.7

4 45:5 41.8—49.7

4 46.5 43.7—48.7

6 41.6 38.0—44.9

1 37.4 —

2 44.6 | 42.4—46.8

1 39.9 —

1 54.3 —

6 AZ 3 38.5 —46.6

2 33:6 34.3 —36.8

6 44.4 40.6 —48.7

5 39.2 38.1—40.6

3 46.7 44.9—49.1

2 40.5 39.9—41.2

5 64.5 60.1 —70.4

1 61.3 —

1 70.5 —

1 59.3 a

2, 70.7 66.5 —74.9

4 54.4 52.1—57.9

Z 51.5 51.2—51.8

2 60.7 57.5—63.8

5 52.8 50.2—57.8

3 55,7 53.7—57.4

1 56.8 —

5 59.3 55.5—67.4

5 58.3 55.5—63.3

5 57.7 53.4—64.7

5 52.8 48.4—55.5

5 49.1 46.0—52.9

z 57.4 53.6—61.1

Z SHS 49.9—52.7

3 47.8 44.5—55.8

5 46.8 43.7—51.0

3 45.7 42.8 —48.5

3 58.1 54.4—61.8

Anatomical Features of the Dipterocarp Timbers of Sarawak 119

References

Ashton, P. S., 1964— Manual of the Dipterocarp Trees of Brunei State, Oxford

University Press.

. 1968 — A Manual of the Dipterocarp Trees of Brunei State and of

Sarawak — Supplement, Borneo Literature Bureau.

Brown, H. P., Panshin, A. J. & Forsaith, C. C., 1944——-Textbook of Wood

Technology, Vol. 1, McGraw-Hill Book Company, Inc.

Browne, F. G., 1955 — Forest Trees of Sarawak and Brunei, Government Printing

Office, Kuching.

Burgess, P. F., 1966—— Timber of Sabah, Sabah Forest Records No. 6.

Chu, F. Fei-Tan, 1969 — A Preliminary Study of Fibre Characteristics of Timbers

of the Peat Swamp Forests of Sarawak, Malayan Forester, Vol. XXXII,

No.3:

Committee on Nomenclature International Association of Wood Anatomists,

1957 — International Glossary of Terms Used in Wood Anatomy, Tropical

Woods No. 107.

Desch, H. E., 1957— Manual of Malayan Timbers, Vol. 1, Malayan Forest

Records No. 15, Reprint.

Forest Department, Sarawak, 1964 Common Sarawak Timbers, Borneo Litera-

ture Bureau.

Forest Products Research, 1960 — Identification of Hardwoods (A Lens Key),

Bulletin No. 25, Her Majesty’s Stationery Office, London.

Forest Products Research, 1961 — Identification of Hardwoods (A Microscope

Key), Bulletin No. 46, Her Majesty’s Stationery Office, London.

Kauman, W. G. & Kloot, N. H., 1968 — Technological Appraisal of Little-Known

or Unused Timber Species, C.S.I.R.O., Division of Forest Products, Australia.

Meijer, W. & Wood, G. H. S., 1964—Dipterocarps of Sabah, Sabah Forest

Records ‘No. ‘5.

Menon, P. K. Balan, 1956 — Siliceous Timbers of Malaya, Malayan Forest

Records No. 19.

Metcalfe, C. R. & Chalk, L., 1950— Anatomy of the Dicotyledons, Vol. I &

Il, Oxford University Press.

Murthy, L. S. V., 1965 — Silica in Sarawak Timbers, Malayan Forester, Vol.

XXVIII, No. 1.

Symington, C. F., 1934— Notes on Malayan Dipterocarpaceae, Gardens’ Bul-

letin II.

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Review

T. C. WuHiTMorRE, Palms of Malaya

Oxford University Press, Kuala Lumpur, Malaysia, 1973. Pp. v +

132, 106 text figures and 16 photographic plates.

Price: Malaysian $35.00

Most people do not seem to realize that among flowering plants, the palm

family is, in economic importance, only next to the grass family to which

numerous cereals belong. For instance, the climbing rattans supply cane for

furnitures, the trunk of some palms is strong enough for building huts or simple

houses while the pith of the sago palms yields starch which was at one time the

first substitute for rice. The leaves supply material for thatch, baskets, etc. The

large terminal buds of several palms were, in some cases wastefully consumed as

vegetables while the sap of the sugar palms yields sugar, and that of the coconut,

on fermentation, forms an alcoholic drink known as toddy. The fruit of many

palm species is edible, the coconut oil, extracted from the solid endosperm,

is used in cooking and for industry; the coconut milk which is the liquid endo-

sperm is an invaluable medium for tissue culture besides being a popular drink.

Moreover, the fibrous, fleshy outer-wall or mesocarp of the oil palm fruit yields

palm oil which is fast becoming one of the most important economic products

in this part of the world.

The flora of the Malay Peninsula and Singapore is extremely rich in palm

species. According to Dr. Whitmore, there are almost as many palms recorded

from Singapore Island (18 genera, 46 species) as on the whole of the African

continent (15 genera, 5O species). In addition, there are a large number of

introduced species of palms which has been naturalised and become a part of

our flora.

Even so, there were no general accounts available on Malayan palms. Under-

standably they were excluded from Wayside Trees of Malaya by E. J. H. Corner

and Malayan Wild Flowers by M. R. Henderson, as palms, rather like bamboos

and pandans, are in a strict sense neither tree nor herb but represent categories

of their own kinds. Abundant information on Malayan palms, nevertheless, is

scattered in the famous Dictionary of Economic Products by |. H. Burkill. Also a

series of valuable taxonomic treatises on various groups of palms by C. X. Furtado

were published in the Gardens’ Bulletin, Singapore and elsewhere.

It is for this reason, that Dr. Whitmore’s new book on palms is especially

welcome. Inside the green, glossy jacket, a general introduction and descriptions

of common Malayan palms form the two major parts. The first is so simple and

clear that readers with elementary knowledge on botany can comprehend. In

the second, taxonomic descriptions are kept to a minimum but there is ample

information on the history, economic use, botanical interest and legends, presented

121

122 Gardens’ Bulletin, Singapore —X XVII (1974)

in such a way that it is thoroughly enjoyable to read. It is Dr. Whitmore’s intention

to show the important part that palms play in our cultural and natural heritage.

Not only has he achieved this but has done so remarkably well.

Regrettably the author resisted the use of the name Johannesteijsmannia.

The name is indeed long and cumbersome but does not appear more “of a

mouthful” than Bougainvillea, Dieffenbackia and Mesembryanthemum. Besides,

it is a legitimate name. Binomials in the text are not complemented by authors’

names though they are in the Checklist of Malayan Palms. The three indices

to vernacular and English palm names are somewhat incomplete. At least one

reference is inexact, i.e. “‘rattan’’ on page 101 really refers to its section “The

Rattan Industry” and should have been page 98. It being a principle reference

should have been in boldface.

Apart from these minor defects, the book is a most welcome publication

and is highly commendable. Production of the book is of high quality and text

is well written. The keys are easy to use. Of the text figures, many are extremely

instructive and the photographs are excellently reproduced. It is the reviewer’s

sincere hope that the publisher will soon put up a cheaper edition to enable this

book to reach a much broader spectrum of readers as it is well deserved.

HsuaN KENG

Department of Botany

University of Singapore

New Plant Disease Records for Sarawak for 1972

KUEH TIONG-KHENG

Department of Agriculture, Sarawak, E. Malaysia

This list of new plant disease records for Sarawak is a follow-up of those

given by Johnston (1960), Turner (1963, 1964, 1966, 1967, 1969 and 1971) and

Kueh (1973). It consists of fungal diseases together with diseases caused by algae

and plant parasitic nematodes, collected and investigated by the writer during

the year 1972. Sixteen of these records appear in the Annual Report of the

Research Branch, Department of Agriculture, Sarawak, for the year 1972.

The microorganisms are arranged under their respective host plants which

are listed alphabetically under their botanical names. For each host plant a common

name and a Malay name are given where available. The frequency of occurrence

of a disease is given together with the Commonwealth Mycological Institute Her-

barium serial number, where identification has been carried out by the Institute.

Identifications performed by the Royal Botanic Gardens, Kew, and Commonwealth

Institute of Helminthology, Herts, are indicated by Kew and CIH respectively.

In the list that follows, One, Occ., Comm. stand for one record, occasionally

and common respectively.

Acacia auriculaeformis A. Cunn. ex Benth. (Black Wattle)

Pink disease Corticium salmonicolor Berk. & Br. One —

Ageratum conyzoides Linn. (Rumput Tahi Ayam)

Leaf rot and Choanephora cucurbitarum (Berk. &

Dieback Rav.) Thaxt. Occ. 171127

Thread blight Marasmius pulcher (Berk. & Br.) Petch One —

Allium cepa L. (Onion, Bawang)

Leaf blight Alternaria tenuissima (Kunze ex Pers.)

Wiltshire Occ. 168326b

Leaf blight Colletotrichum circinans (Berk.) Vogl. Occ. 168326c

Leaf blight Curvularia eragrostidis (P. Henn.)

J. A. Meyer Occ. 168326a

Leaf blight Curvularia penniseti (Mitra) Boedijn Occ. 168326e

Leaf blight Glomerella cingulata (Stonem.) Spauld.

& Schrenk Occ. 168326g

Leaf blight Nigrospora sacchari (Speg.) Mason Occ. 168326f

Leaf rot Choanephora cucurbitarum (Berk. &

Rav.) Thaxt. One —

Ananas cosmosus Merr. (Pineapple, Nanas)

Leaf blight Botryodiplodia theobromac Pat. One 174347a

Leaf blight Phomopsis sp. One 174347b

Areca catechu Linn. (Betel Palm, Pinang)

Frond base decay Marasmiellus inoderma (Berk.) Singer One Kew

124 Gardens’ Bulletin, Singapore — XXVII (1974)

Artocarpus heterophyllus Lam. (Jackfruit, Nangka)

Inflorescence rot Rhizopus artocarpi Racib. One —

Asparagus officinalis Linn. (Asparagus, Saparu Keras)

Stem blight Phoma herbarum Westend. One 168321

Bassia latifolia Roxb. (Indian Butter Tree, Mowa Tree)

Red rust Cephaleuros virescens Kunze Occ. 171125a

Sooty mould Trichomerium? crotonis Bat. One 171125b

Sooty mould Tripospermum sp. One 171125b

Sooty mould Phaeosaccardinula? tenuis (Earle)

Seaver & Chardon One 171125b

Begonia sp.

Root knot Meloidogyne incognita (Kofoid &

White, 1919) Chitwood, 1949. Occ. Ci

Root knot Meloidogyne javanica (Treub, 1885)

Chitwood, 1949. Occ... CEH

On root Helicotylenchus dihystera (Cobb, 1893)

Sher, 1961 Occ. Clo

Bixa orellana L. (Anatto, Kesumba Keling)

Stem lesion Cephaleuros virescens Kunze One —

Borreria latifolia Schum. (Broad Leaved Button-Weed, Rumput Setawar)

Leaf blight Curvularia borreriae (Viégas) M. B. Ellis Occ. 173903

Brassica chinensis Linn. (Chinese Cabbage, Pak Choy, Sawi Putih)

Leaf spot Peronospora parasitica (Pers. ex Fr.) Fr. One 168330

Brassica oleracea var. capitata Linn. (Cabbage. Pow Chai)

Circular spot Alternaria brassicicola (Schw.) Wiltshire One 171119a

Circular spot Leptosphaerulina trifolii (Rostr.) Petr. One 171119b

Canangium odoratum Baill. (Kenanga)

Sooty mould Meliola canangae Stev. Occ. 171124

Carica papaya Linn. (Papaya, Betik)

Fruit anthracnose Cladosporium oxysporum Berk. & Curt. One 166644b

Celosia argentea Linn.

Leaf speckle & Cercospora celosiarum Kar & Mandal Comm. 168915

Inflorescence mould

Celosia cristata Linn. (Cockscomb)

Leaf spot Cochliobolus geniculatus Nelson Occ. 168907a

Leaf spot Phyllosticta celosiae Thiim. Occ. 168907b

Chrysanthemum sp.

Leaf blight Stemphylum lycopersici (Enjoji)

Yamamoto One 173904

Leaf blight Cochliobolus geniculatus Nelson One 174346a

Leaf blight Phomopsis sp. One 174346b

Citrullus vulgaris L. (Water Melon, Semangka)

Leaf spot Cercospora citrullina Cooke Comm. 168317

Fruit rot Cochliobolus geniculatus Nelson Oce, 172113

Fruit rot Curvularia senegalensis (Speg.) Subram. Occ. 167057

Plant disease records, Sarawak, 1972

Citrus aurantifolia (Christm.) Swingle (Lime, Limau Masam)

Red rust Cephaleuros virescens Kunze

Citrus grandis (L.) Osb. (Pomelo, Limau Besar)

Dieback Fusarium coccophilum (Desm.) Wr.

Velvet blight Septobasidium lepidosaphis Couch

Gummosis Phytophthora nicotianae B. de Haan var.

parasitica (Dastur) Waterh.

Citrus nobilis Lour. (Mandarin Orange, Limau Cina)

Leaf rot Corticium solani (Prill. & Delacr.)

Bourd. & Galz.

Citrus sinensis Osb. (Sweet Orange, Limau Manis)

Dieback Botryodiplodia theobromae Pat.

Cocos nucifera L. (Coconut, Kelapa, Nyiur)

Leaf mould Pseudoepicoccum cocos (F. L. Stevens)

M. B. Ellis

Cucurbita pepo DC. (Pumpkin, Buah Labu)

Leaf spot Cercospora citrullina Cooke

Curcuma domestica Valeton (Turmeric, Kunyit)

Sclerotial wilt Corticium rolfsti Curzi

Desmodium umbellatum (L.) DC. (Sea Parkia, Petai Laut)

Leaf spot Cercospora sp.

Dieffenbachia sequina L. (Dumbcane)

Leaf blotch Macrophomina phaseolina (Tassi) Goid

Digitaria marginata Link

Leaf spot Pyricularia grisea (Cooke) Sacc.

Dracaena sanderiana Hort. Sand

Leaf blight Stemphylium lycopersici (Enjoji)

Yamamoto

Leaf blight Glomerella cingulata (Stonem.) Spauid.

& Schrenk

Durio zibethinus Murr. (Durian)

Sooty mould Capnodium moniliforme Fraser

Elaeis guineensis Jacq. (Oil Palm, Kelapa Sawit)

Frond base rot Porogramme ravenalae (Berk. & Br.) Pat.

Elettaria cardamomum Maton (Cardamom)

Leaf spot Phaeotrichoconis crotalariae (Salam &

Rao) Subram.

Sooty mould ? Asbolisia sp.

Sooty mould ? Microxyphium sp.

Sooty mould ? Cladosporium sp.

Erechtites hieraciifolia Rafin.

Leaf spot Colletotrichum sp.

Erechtites valerianifolia DC. (Malayan Groundsel)

Leaf spot Cercospora erechtitis Atkinson

Comm. —

One

ice.

Occ.

One

Comm. 166649

Occ.

One

Occ.

One

Occ.

One

Comm. 168909 &

173899

163736

168911

167053

1666Sic

171121

168912

173900

173902

171117a

171117b

174350

Kew

171115

171116

168908a

125

126 Gardens’ Bulletin, Singapore —X XVII (1974)

Eugenia aromatica Kuntze (Clove, Cengkih)

Red rust Cephaleuros virescens Kunze Comm. 168328

Ficus sp.

Thread blight Marasmiellus scandens (Mass.) Dennis

& Reid One —

Gerbera jamesonii Bolus (Barberton Daisy)

Leaf spot Cercospora gerberae Chupp & Viégas One 174344

Glycine max (L.) Merr. (Soybean, Kacang Bulu Limau)

Leaf blight Negrospora sphaerica (Sacc.) Mason One 173907a

Leaf blight Botryodiplodia theobromae Pat. One 173907¢g

Leaf blight Cladosporium cladosporioides (Fresen.)

de Vries One 173907f

Leaf blight Cochliobolus lunatus Nelson & Haasis One 173907d

Leaf blight Cochliobolus geniculatus Nelson One 173907e

Leaf blight Phaeoseptoria sp. One 173907h

Pod spotting Cercospora kikuchii (Mat. & Tom.)

Gardner One 168325

Root disease Marasmiellus inoderma (Berk.) Singer One Kew

Hibiscus esculentus L. (Ladies’ Fingers, Kacang Lender)

Root knot Meloidogyne incognita (Kofoid &

White, 1919) Chitwood, 1949. One CIH

Root lesion Practylenchus brachyurus (Godfrey, 1929)

Filipjey & Schuurmans Stekhoven,

1941 One CIH

Hibiscus syriacus Linn.

White root disease Fomes lignosus (Klotzsch) Bres. One —

Hydrangea hortensia Siebold

Leaf spot Corynespora cassiicola (Berk. & Curt.)

Wei One 171126a

Leaf spot Cercospora hydrangeae Ell. & Ev. One 171126b

Impatiens balsamina L. (Balsam)

Stem blight Glomerella cingulata (Stonem.) Spauld.

& Schrenk Occ. 168906a

Imperata cylindrica Beauv. (Lalang)

Thread blight Marasmius pulcher (Berk. & Br.) Petch One Kew

Ischaemum digitatum Brongn.

Leaf spot Colletotrichum graminicola (Ces.) Wilson One 166640

Ischaemum magnum Rendle

False smut Sphacelotheca hainanae. Zundel Occ. 166687

Lycopersicum esculentum Mill. (Tomato)

Leaf rot Choanephora cucurbitarum (Berk. &

Rav.) Thaxt. One —

Mangifera indica Linn. (Indian Mango)

Premature fruit drop Glomerella cingulata (Stonem.) Spauld.

& Schrenk One 166650

Plant disease records, Sarawak, 1972

Mangifera foetida Lour. (Horse Mango, Bacang)

Leaf spot Coniothyrium fuckelii Sacc.

Grey leaf blight Pestalotiopsis sp.

Leat spot Glomerella cingulata (Stonem’) Spauld.

& Schrenk

Leaf spot Phomopsis mangiferae Ahmad

127

One 166646c

One 166646d

One 166646b

One 166646a

Melastoma malabathricum L. (Singapore Rhododendron, Engkodok)

Sooty mould Trichomerium sp.

Michelia alba DC. (Cempaka Putih)

Red rust Cephaleuros virescens Kunze

Morus alba Linn. (Mulberry)

Leaf mould Cercospora mori Hara

Leaf spot Cercospora sp.

Cutting necrosis Colletotrichum capsici (Syd.) Butler

& Bisby

Nephelium lappaceum L. (Rambutan)

Root rot Sphaerostilbe repens Berk. & Br.

Rim blight Pestalotiopsis cruenta (Syd.) Steyaert

Rim blight Pestalotiopsis sp.

Oryza sativa L. (Rice, Padi)

On panicle Tetraploa aristata B. & Br.

Passiflora edulis Sims (Passion Fruit)

Flower blight Choanephora cucurbitarum (Berk. &

Rav.) Thaxt.

Pennisetum purpureum Schumach. (Napier Grass)

Leaf spot Curvularia leonensis M. B. Ellis

Leaf spot Nigrospora sphaerica (Sacc.) Mason

Phaseolus atropurpureus (Moc. & Sesse) ex DC.

Leaf & Stem rot Corticium rolfsii Curzi

Piper colubrinum L.

Black leaf spot Phytophthora palmivora (Butler) Butler

Piper nigrum L. (Black Pepper, Lada)

On berries & leaves

Root knot

Corticium rolfsii Curzi

1919) Chitwood, 1949

Curvularia eragrostidis (P. Henn.)

J. A. Meyer

Leptosphaeria suffulta (Fr.) Niessl.

Leaf blight

Pogostemon cablin Benth. (Patchouli)

Slow decline

Leaf spot

Fusarium solani (Mart.) Sacc.

Cercospora sp.

Colletotrichum capsici (Syd.) Butler &

Bisby

Psidium guajava L. (Guava)

Leaf blight Pestalotia psidii Pat.

Meloidogyne incognita (Kofoid & White,

Occ. 168318

Occ. —

Comm. 168319

One 163735

One 167054a

One —

Comm. 166648

Occ. 173905a

One —

One 166645a

One 166645b

Oce..° =

One 172112

One —

Occ. CIH

One 168910a

One 168910b

Comm. 172115

Occ. 173901b

Occ. 173901la

Occ. 166641

Raphanus sativus L. Var. hortensis Backer (Chinese Radish, Lobak)

Circular leaf spot

Alternaria brassicicola (Schw.) Wiltshire

Occ. 168329

128

Ricinus communis L. (Castor Oil Plant)

Thread blight Marasmiellus scandens (Mass.) Dennis

& Reid

Corticium solani (Prill. & Delacr.)

Bourd. & Galz.

Leaf rot

Rosa sp. (Rose)

Sooty mould Trichomerium sp.

Sanseviera laurentii (N.E. Br.) De Wild.

Rim blight Botryodiplodia theobromae Pat.

Rim blight Cochliobolus geniculatus Nelson

Rim blight Fusarium oxysporum Schlecht.

Sauropus androgynus Merr. (Cangkok Manis)

Cercospora phylianthicola Shakil &

Kamal

Glomerella cingulata (Stonem.) Spauld.

& Schrenk

Alternaria sp.

Leaf spot

Stem blight

Leaf rot Alternaria macrospora Zimm.

Leaf rot Choanephora cucurbitarum (Berk. &

Rav.) Thaxt.

Leaf rot Corynespora cassiicola (Berk. & Curt.)

Wei

Stenolobium stans Seem.

Sooty mould

Trichomerium sp.

Podoxyphium sp.

Stylosanthes gracilis H.B.K. (Stylo)

Leaf & stem rot Corticium rolfsii Curzi

Theobroma cacao L. (Cocoa, Koko)

Tripospermum gardneri (Berk.) Speg.

ex Hendrickx

Sooty mould

Vanilla planifolia Andr. (Vanilla)

Leaf blight Curvularia leonensis M. B. Ellis

Leaf blight Cochliobolus lunata Nelson & Haasis

Leaf blight Myrothecium gramineum Lib.

Vigna sesquipedalis (L.) Fruw. (Long Bean, Kacang Panjang)

Leaf anthracnose Colletotrichum lindemuthianum (Sacc. &

Magn.) Bri. & Cav.

Associated with leaf Ascochyta phaseolorum Sacc.

anthracnose

Leaf rot Choanephora cucurbitarum (Berk. &

Rav.) Thaxt.

Voandzeia subterranea Thouars (Kacang Bogor)

Leaf & stem rot Corticium rolfsii Curzi

Xanthophyllum aff. pulchrum King

Sooty mould Trichomerium sp.

One

Occ.

One

Occ.

One

Occ.

One

Gardens’ Bulletin, Singapore — XXVII (1974)

171120

166642a

166642b

166642c

168320

168905a

168905b

174345a

174345b

174345c

168913

166639

168322a

168322b

168322c

168327a

168327b

168914

tt tt EL

Plant disease records, Sarawak, 1972 129

Zingiber officinale Rosc. (Ginger, Haliya)

Leaf blotch Glomerella cingulata (Stonem.) Spauld.

& Schrenk One 168902b

Leaf tip dieback Mycospharella sp. One 168916b

Leaf scorch ? Pseudocercosporella sp. One 171122b

Leaf scorch Curvularia eragrostidis (P. Henn.)

J. A. Meyer Occ. 172114b

Leaf spot Phoma eupyrena Sacc. Occ. 172114a

Leaf scorch Curvularia senegalensis (Speg.) Subram. One 167055

White root disease Fomes lignosus (Klotzsch) Bres. One —

Acknowledgements

I am grateful to the Director of Agriculture, Sarawak. for granting permission

to publish this list and to the Directors and staff of the Commonwealth Mycological

Institute and the Royal Botanic Gardens, Kew, for their prompt assistance in

identifying many specimens. My thanks are also due to the Director and staff

of the Commonwealth Institute of Helminthology, Herts, where plant parasitic

nematodes were identified.

The assistance rendered by Mr. Soo Tan Kui and Mr. Bi'ee anak Jagat in

the field is also gratefully acknowledged.

References

Annual Report, (1972). Research Branch, Department of Agriculture, Sarawak.

(In press).

Johnston, A. (1960). A preliminary plant disease survey in Sarawak. F.A.O. Rome

(Mimeographed).

Kueh Tiong-Kheng, (1973). New plant disease records for Sarawak for 1970-1971.

Gardens’ Bull. Singapore, 26: 263-268.

Turner, G. J. (1963). New records of plant diseases in Sarawak for the years

1960-1961. Gardens’ Bull. Singapore, 20: 285-288.

. (1964). Ibid. 1962. Gardens’ Bull. Singapore, 20: 369-376.

. (1966). Ibid. 1963 & 1964. Gardens’ Bull. Singapore, 21: 393-402.

. (1967). Ibid. 1965. Gardens’ Bull. Singapore, 22: 123-128.

. (1969). Ibid. 1966 & 1967. Gardens’ Bull. Singapore, 24: 181-184.

——— —. (1971). Fungi and plant diseases in Sarawak. Phytopath. Papers, No. 13,

Cwealth Mycol. Inst., Kew, Surrey, England.

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Review

Animal Life and Nature in Singapore

edited by S. H. CHUANG

Singapore University Press, Singapore 1973.

Pp. xiv + 302, 34 text figures including 10 line drawings,

9 maps, 12 charts and diagrarns, and 11 coloured plates.

Price: Singapore $40.00

The arrival of this book on the Singapore scene is most welcome. For too

long, keen naturalists in Singapore have had to be content with books on natural

history which treated Singapore merely as a minor part of the Malay Archipelago.

It is true that the fauna of Singapore is basically an impoverished version of

the Malaysian but, there are many interesting phenomena such as the adaptation

of the fauna to artificial environments which can be better studied in Singapore

than elsewhere.

The range of subjects included in this volume is wide. Besides chapters

dealing with the animal life of Singapore, there are others which discuss the

vegetation, climate, geclogy and the sea around the Island.

The change in the composition of the fauna of Singapore in the recent past

is something which many of the writers have noted. They record the gradual

replacement of some species by others, and mention these changes among both

the vertebrates and invertebrates. As can be expected of a book compiled by ten

different authors, the coverage given to each subject differs. Some have chosen

to give broad general pictures of their fields, while other have chosen to highlight

only certain aspects of their subjects. But whichever approach they chose to their

subjects, the writers have all managed to produce articles which were factual at

the time of publication. D. S. Johnson in particular has written a well researched

piece about the changes in the composition of the bird life of Singapore in the

recent past. In it he mentions the disappearance of the magpie robin and the

increasing abundance of the mynah. He also makes observations on changes in

the populations of crows, egrets, orioles and munias, and expounds some credible

theories to explain these changes. He says that the widespread use of insecticides

has led to a decrease in the numbers of insect feeders, and, modern grain handling

methods which result in less spillage have led to a corresponding decline in the

numbers of the birds which feed on spilled grain.

Similar changes in the sea life around Singapore have been recorded by Tham

Ah Kow and S. H. Chuang in their chapters. In one, Chuang gives an interesting

mathematical table showing the changes in the species of shells off Tanah Merah

beach. Both Tham and Chuang write that the depletion of the marine life around

Singapore is attributable to the increased pollution of the water.

ba Gardens’ Bulletin, Singapore —- X XVII (1974)

R. E. Sharma has contributed a chapter on noxious and toxic animals. This

chapter contains useful information for anyone venturing out to study nature in

Singapore. In it, Sharma has mentioned practically every animal which could

conceivably be a danger to human beings, and in many cases he outlines preventive

measures which can minimise the danger.

Unfortunately, data for the book was collected up to 1969, and the authors

could not possibly have foreseen the rapid pace of urbanisation in Singapore. Since

that time, swamps have been filled in and forested areas cleared. The end result

is that, while the book is currently still accurate as far as the various species of

animals mentioned in it are concerned, their actual numbers should be regarded

as an optimistic record. An example is the chapter by R. C. R. Morrell on

butterflies and moths. In it he mentions that hawk moths are still fairly well repre-

sented in Singapore. This might have been true five years ago, but today, with the

decrease in scrublands, increase in buildings and improved street lighting which

attracts moths and makes them easy prey for bats and birds, hawk moths are

certainly few and seen at longer intervals. His account of the butterflies here also

tends towards optimism. While the species which he mentions are still to be found

here, there is no doubt that their numbers have been greatly reduced.

The standard of photography is not as high as could be desired. Birds on

Plate 9 are not shot artistically. Little attention is paid to background in Plates 3

to 9. As a result the photographs portray specimens in the laboratory rather than

animals in life.

The treatment of “The Exploitation of Animals’ is rather cursory. There

are a few misnomers, e.g. “jambols’’ on page 253 are referred to “‘crested jays”

which should have been ‘“‘bulbuls’’.

Considered as a whole, this book is by no means comprehensive in its coverage

but it does serve to give an accurate general picture of animal life in Singapore

today. However, conditions here are changing so fast that it will come as no

surprise if the material contained in it becomes outdated five years from now.

Despite the shortcomings, the book serves a very useful function. In it the

authors have pointed out areas where there is plenty of scope for further study

and research. It should therefore be considered more than merely a record of

the fauna of Singapore and serve as the starting point from which students can

undertake a serious study of animal life and nature in Singapore.

LEE Cuu SAN

Grant Advtg International Inc.

Singapore

The Genus Horsfieldia (Mvristicaceae)

in and outside Malesia I:

H. sabulosa and H. whitmorei J. Sinclair spp. nov.

+ by J. SINCLAIR

‘Botanic Gardens

Singapore

The present paper comes from a manuscript which at the time of the author’s

demise was at Kew Herbarium, sent there in advance for his proposed completion

of the work on the genus Horsfieldia. The Botanic Gardens in Singapore gratefully

acknowledge the co-operation from individuals and herbaria who have loaned their

specimens for the study by the late Mr. J. Sinclair and are grateful to Mr. J. P. M.

Brennan, Keeper of the Kew Herbarium and Library for promptly taken steps in

returning the script. The manuscript contains complete descriptions of 33 species

and many varieties, and citations only of specimens under H. macrocoma and

H. irya. Regretfully, the classification system, many keys, distribution maps, illustra-

tions and the collector’s index had not been written up. This article describes the

two new species named by the author and is the first in the series. Ed.

Horsfieldia sabulosa J. Sinclair, sp. nov.

Species affinis H. wallichii a qua ramulis crassioribus, teretibus (nec apice

leviter compressis), foliis multo angustioribus, apices ramulorum versus dense con-

fertis, polystichis nec distichis, petiolis longioribus et nervis plerumque minus dis-

tinctis differt.

Arbor excelsa 10-37 m alta. Cortex ferrugineus longitudinaliter sulcatus; latex

ruber, copiosus. Ramuli in innovationibus ferrugineo- vel griseo-ferrugineo-tomen-

telli, 0.7—1 cm crassi, in partibus inferis glabri, nigro-grisei et cum multis cicatricibus

foliorum delapsorum spiraliter tecti, distaliter longo intervallo simplices ut videtur,

forsitan multo inferne ramos primarios versus furcati, cortice interdum excidente.

Folia polysticha, apices ramulorum versus dense conferta, coriacea, angustissime

oblonga, marginibus fere parallelis minute revolutis, supra nitida, modice viridia,

subtus glauca, siccitate supra griseo-viridia, hic illic nigricantia, subtus moribundo-

brunnea, utrinque acuta, basi in petiolum paullo decurrentia, 15-18 cm longa,

Saepissime 16 cm longa, 4-6 cm lata, vulgo 4.5 cm lata; costa supra in sulcam

depressa, subtus convexo-elevata ibique primo minute puberula deinde glabra;

nervi 15—18—jugati, fere paralleli, supra depressi, tenues, similes incisuris apertis

cultro tonsorio inflictis, subtus minus distincti et in partibus subevanidi; reticula-

tiones invisibiles; petioli (pro laminis comparati longi) 3.5—S cm longi, 3 mm crassi,

minute tomentelli, glabrescentes. Inflorescentia mascula et flores masculi ignoti.

Inflorescentia feminea immatura, nondum evoluta, 2 cm longa, bracteis pluribus

bene munita: bracteae | cm longae, 2-4 mm latae, pilis dendroideis 1-1.5 mm

longis dense indutae, dactyloideae, supra flores inflexae et eos celantes: flores

minutissimi, enimvero multo immaturi, glabri, breviter pedicellati; ovarium

134 Gardens’ Bulletin, Singapore — XXVII (1974)

glabrum. Fructus flavus, glaber, ovoideus, in sicco 4-4.5 cm longus, 3.5-4 cm

latus, ei H. wallichii similis sed probabiliter paullo minor; pericarpium in vivo

camosum, in sicco lignosum, 5 mm crassum, non nitidum (opacum); stipes (pedi-

cellus) 0.5-1 cm longus, 5 mm crassus. Arillus aurantiacus. Semen 3 cm longum,

2.7 cm latum.

Tall tree 10-37 m high (30-120 ft). Bark reddish brown, longitudinally fur-

rowed; sap red, copious. Twigs in the innovations rusty- or rusty-greyish-tomen-

tulose, 7 mm -l cm thick, lower down glabrous, blackish grey and _ spirally

covered with many scars of fallen leaves, apparently simple distally for some

distance, probably branched far down near their junction with the main branches,

the bark sometimes peeling off. Leaves in several rows, bunched towards the ends

of the twigs, coriaceous, very narrowly oblong with the sides nearly parallel and

minutely revolute, medium green and glossy above, glaucous beneath, drying greyish

green above with some blackish patches and a faded brown beneath, acute at

both ends with the base slightly decurrent on to the petiole, 15-18 cm long,

many of them constantly 16 cm long, 4-6 cm broad, average 4.5 cm broad: the

midrib sunk in a groove above, raised and convex beneath and there at first

minutely puberulous, later glabrous; nerves 15-18 pairs, nearly parallel, sunk

above, very slender and like open slashes with a razor blade, less distinct and

vanishing in parts beneath; reticulations invisible; the petioles 3.5-5 cm long

(long in proportion to their blades) 3 mm thick, minutely tomentulose becoming

glabrous. Male inflorescence and male flowers unknown. Female inflorescence im-

mature, not yet open, 2 cm long and well protected by bracts; the bracts 1 cm

long and 2-4 mm broad, densely covered with 1-1.5 mm long dendroid hairs,

finger-shaped, curving over the flowers and concealing them; flowers very small and

indeed very immature, glabrous and shortly stalked; ovary glabrous. (The flowers

are so small that there is no point in giving measurements and further detatis.)

Fruit yellow, glabrous, ovoid, 4-4.5 cm long and 3.5—4 cm broad (measurements

from dried material), similar to that of H. wallichii but probably a little smaller,

the pericarp fleshy when fresh, woody and brown when dry, 5 mm thick, dull

having the usual mat or parchment-like surface of dried Horsfieldia fruits; stalk

(the pedicel portion) 0.5—-1 cm long and 5 mm thick. Avil orange. Seed 3 cm

long and 2.7 cm broad.

BORNEO

SARAWAK:

Ist Division:—Gunong Gaharu, Serian, Sinclair 10248 (A, E, K, L, SAR, SING):

Sungei Sabal Tapang, Serian, SAR Nos 12671 (SAR) and 12691 (SAR).*

3rd Division:—Aup, Sibu, J. Wright 632 as FA 412 (SAR, SING).

4th Division:—Niah-Jelalong Protected Forest, sia Meluang, Bintulu, Briinig

SAR 956 (K, L, SAN, SAR, SING).

BRUNEI:

Andulau Forest Reserve, Belait District, Ashton & Whitmore BRUN 579 (BO, K,

KEP, L, SING); Smythies BRUN 828 (BO, K, L, KEP, L, SAR, SING); cpt 6.

Ashton, Smythies & Wood SAN 17560 (KEP, L) and ditto (north part) Sinclair &

Kadim bin Tassim 10437 (A, BM, E, FI, K, L, SAR, SING); Bukit Labi at the

51 milestone: Sinclair & Kadim 10491 (A, B, E, K, L, NY, SAR, SING).

-* Author intended to check the identity of 12671 & 12691.

Horsfieldia sabulosa and H. whitmorei spp. nov. 135

SABAH:

Interior Residency:—Mengalong Forest Reserve, Sibubu River, 34 miles south-

south-west of Sipitang, Wood SAN 15146 (L, SAN, SING).

DISTRIBUTION: Borneo (as above).

TYPE MATERIAL: Sinclair & Kadim 10491 (A, B, E, K holotype, L, NY, SAR,

SING) Bukit Labi F.R., Brunei.

A tree of sandy soil or sand with a little peat; in Brunei sometimes growing

with Agathis alba ssp. borneensis. It is closely allied to H. wallichii from which

it should be distinguished vegetatively by the growth habit of its twigs and the

narrower leaves with longer petioles. It is a pachycaul whereas wallichii is a

leptocaul. Thus in sabulosa the leaves are all closely bunched towards the ends of

the twigs in a spiral, non-distichous fashion. The twigs are much stouter than

those of wallichii and are not flattened or compressed at their extremities. Below

the innovations they bear numerous scars of fallen leaves also in several ranks.

Such portions of twigs as conveniently fit a herbarium sheet are not branched;

some of them probably branch far down near where they join on to the main

branches. In contrast similar portions of the same length in wallichii are usually

branched once on a herbarium sheet if not two or three times. The narrower leaves

of sabulosa have their sides nearly parallel and do not broaden out just above

the base. The 3.5-5 cm long petioles at once catch the eye as being long in pro-

portion to the blade. They are 2 cm long on the average in wallichii with an overall

range of 1-3 cm. The veins on the undersurface of the leaves are much fainter

than those of wallichii and sometimes fade out altogether, but, as is the general

tule in Myristicaceae thin-leaved specimens with more prominent veins not yet

collected will probably turn up. A greyish brown indumentum on the young

petioles, lower midrib and innovations may also help in identifying this species.

Of the two, sabulosa seems to be the taller for the examples I collected in Brunei

were from trees 100 and 120 feet high. Ashton also collected from trees 100 feet

in height. Along with H. ridleyana all three have the same reddish brown, longi-

tudinally furrowed bark and a rather similar firm fleshy fruit which has a minutely

rough parchment-like surface when dry. The fruit of sabulosa is probably slightly

smaller than that of wallichii, ridleyana having the smallest but this information

requires confirmation.

It seems strange that with so different a vegetative habit the fruit of both

species should be the same. It may be that sabulosa is after all only a variety or

subspecies of wallichii, yet this seems rather unlikely. I have been waiting patiently

since 1955 for male flowers to turn up, hoping that they may finally settle this

problem. The female flowers from Wood SAN 15146 collected near Sipitang.

Sabah in 1955 are so very young as to be of no value in deciding. Even their

inflorescences have not yet unfolded from the bud stage.

Horsfieldia whitmorei J. Sinclair, sp. nov.

H. novoguineensis (non Warb.) A. C. Smith in J. Arn. Arb. 22, 1 (1941) 62

pro parte quoad Kajewski 2022 (altera pars = H. irya et H. spicata): H.

palewensis (non Kanehira) Whitmore, Guide to the Forests of the British

Solomon Islands (1966) 131 & 186 in check list, correct original spelling is

palauensis).

136 Gardens’ Bulletin, Singapore —X XVII (1974)

Species endemica affinis H. iryae et H. spicatae et cum eis in Insulis Salomonis

crescens. A priore dimidia parte floribus masculis longioribus et laxius dispositis in

inflorescentia breviore et minus evoluta, fructibus oblongis, reticulationibus foliorum

subtus scalariformibus nec retiformibus differt; ab altera floribus masculis minoribus

globosis nec lateraliter elongatis, foliis pro rata angustioribus, costa in canaliculo

petioli elevata, reticulationibus subtus saepius visibilibus, lenticellis minoribus

recedit; ab ambabus floribus pallidioribus (cremeis), ramulis rubro-brunneis (non

nigro-griseis ut in irya nec stramineis ut in spicata), subtilius striatis cum lineis

duabus ex petiolo ad petiolum carentibus vel subnullis, nervis foliorum pro rata

magnitudini laminae plus numerosis, inter se magis approximatis obliquis, in arcibus

singulis, raro duplo-anastomosantibus cum arcibus marginalibus ipsis prominentio-

ribus distinguitur.

Arbor 7-22 m alta, vulgo 12 m, sine radicibus adventitiis. Cortex fibrosus

extus fuscus, intus rubro-brunneus vel subroseus, verticaliter fissuratus demum

papyraceo-squamulosus; latex ruber copiosus. Ramuli 3-4 mm crassi rubro-brunnei

(lateritii) teretes, in innovationibus cum pilis brevissimis stellato-dendroideis tomen-

telli, in partibus inferis glabri, striolati, nonnunquam lenticellati, hic illic in quibus-

dam internodis excavati. Folia chartacea, infrequenter coriacea, supra in sicco

griseo-brunnea vel virido-brunnea, subtus pallidiora, nervis exceptis leviter rubro-

brunneis, glabra, anguste oblonga cum lateribus fere parallelis, basi acuta, apice

acuminata; costa supra sulcata, plana vel convexa, et itidem in sulco petioli con-

spicue elevata; nervi (16)—22—26—(30)-jugati, vulgo 23 paria, in foliis parvis 16

paria, inter se proximi, 7 mm —1.5 cm distantes, vulgo 8 mm, recti, paralleli vel

saepe curvati, angulo 45-70° orti, supra impressi, subtus prominentes, prope

marginem in arcibus singulis perspicue anastomosantes (tantum in foliis latissimis

duplo conjuncti); reticulationes supra invisibiles, subtus subtiliter scalariformes;

lamina 12-30-(42) cm, vulgo 21 cm longa, 3-6.5-(9) cm, vulgo 4.5 cm lata,

petiolus 0.7-1-1.5 cm, vulgo 1 cm latus, 2 mm crassus. /nflorescentia mascula cum

pilis minutis dendroideis ferrugineis vel pallido-brunneis vel griseis pubescens vel

breviter tomentosa, gracilis, 2-8 cm longa, 1 cm in diam.; ramuli secundarii ac

tertiarii 0.5-3 cm longi, primo adscendentes, deinde patentes vel reflexi, ultimi in

cymas subracemosas terminantes. Flores masculi fragrantes, cremei vel flavidi

semper coloris pallidi nunquam aurantii vel intense flavi, in textura variabiles,

tenues vel coriacei, glabri vel saepe in partibus inferioribus tenuiter pilosi, globosi,

in sicco 1.5 mm in diam., in vivo 2 mm, primum in lobos duos rotundos 3-fissi,

denique fissura perianthii deorsum suturam secus fere ad basin floris attingens,

sutura ea plerumque prominens elevata, non-nunquam in sulcam circumferentialem

depressa. Columna_ staminalis compresso-globosa, 1 mm in diam., brevissime

stipitata vel fere sessilis, eius depressio apicalis stomatibus similis, primum angusta

et fere clausa cum lateribus arcte compressis, postea ad plenum anthesin in lati-

tudine augens, cavitas nunc profundior, } partem totae columnae aequialta; antherae

10 in cupulam staminalis inflexae, apicibus interdum sublilberae; pedicelli 1.5—1.8

mm longi, aliquando 2-(2.5) mm longi, 0.2 mm crassi, pilis ut in inflorescentia

pubescentes. Inflorescentia feminea 1.5—5 cm longa, ramuli eius quam ei inflore-

scentiae masculae breviores pauciores, 0.5—-1 cm longi; inflor. fructifera 10 cm

longam attingens. Flores feminei ovoideo-globosi, 2 mm in diam. (in sicco), aliter

ut in masculi; pedicelli 1 mm longi, 0.5 mm crassi; ovarium tomentosum, 1.5 mm

longum, 1-1.2 mm latum. Fructus pallido-flavus (probabiliter in maturitate rube-

scens) in sicco rubro-brunneus, glabrus, oblongus, utrinque rotundatus, 2—2.3 cm

longus, 1.5 cm latus cum linea suturali prominenti; stipes 1 cm longus, 2-3 mm

crassus. Arillus aurantiacus.

——————————

Horsfieldia sabulosa and H. whitmorei spp. nov. 137

Tree 7-22 m high, average 12 m, stilt-roots absent. Bark fibrous, dark brown

on the surface, reddish brown to pink inside, vertically fissured and finally flaking

in small papery scales; sap red, abundant, free flowing. Twigs 3-4 mm thick,

reddish brown (brick-red) terete, tomentulose in the innovations with very short

stellate-dendroid hairs, lower down glabrous, finely striate and sometimes lenticel-

late, hollow here and there in certain internodes. Leaves chartaceous, infrequently

coriaceous, drying greyish brown or greenish brown above, paler beneath except

the reddish brown nerves, glabrous, narrowly oblong with the sides nearly parallel,

the base acute, the apex acuminate; the midrib sulcate above, flat or convex and

moreover in the same way conspicuously raised in the channel of the petiole; the

nerves (16)-22—26-(30) pairs, mostly 23 pairs, 16 pairs in small leaves, close to

each other, 0.7-1.5 cm apart, usually 8 mm, straight, parallel or often curved,

arising at an angle of 45—70°, impressed above, prominent beneath, very clearly inter-

arching near the margin in single loops (only in double loops in very large leaves):

reticulations invisible above, very finely scalariform beneath; blade 12—30-(42) cm

long, usually 21 cm long, 3-6.5—(9) cm broad, average 4.5 cm broad; petiole 0.7-—

1-1.5 cm long, average 1 cm long, 2 mm thick. Male inflorescence pubescent

or shortly tomentose with minute rusty or pale brown or greyish dendroid hairs,

slender, 2-8 cm long and 1 mm in diam.; the secondary and tertiary branches 0.5

—3 cm long, at first ascending, later spreading and reflexed, the ultimate ones

ending in subracemose cymes. Male flowers sweet scented, cream-coloured or pale

yellow, always of a pale colour, never orange or a deep yellow, variable in

texture, thin or coriaceous, glabrous or often thinly pilose, globose, 1.5 mm

in diam. in dried specimens, 2 mm in diam. in fresh ones, at first split down

4-way into the two rounded lobes, finally the split of the perianth reaching

downwards along a suture almost to the base of the flower, the suture

usually prominent and raised, sometimes sunk in a_ groove girdling

the circumference. Staminal column globose but slightly flattened laterally,

1 mm in diam., very shortly stalked or almost sessile, its apical depression like

that of stomata, at first narrow and almost closed with the sides tightly drawn

together, later increasing in width at the peak of flowering, the cavity now

deeper, + as deep as the whole column; anthers 10, bent over into this staminal

cup, sometimes slightly free at their apices; pedicels 1.5-1.8 mm long, occasionally

2+(2.5) mm long, 0.2 mm thick, pubescent with hairs as in the inflorescence.

Female inflorescence 1.5-5 cm long, its branches shorter and fewer than those

of the male inflorescence, 0.5—1 cm long; fruiting inflorescence reaching 10 cm

long. Female flowers ovoid-globose, 2 mm in diam. (when dry), otherwise as in

the male; pedicels 1 mm long and 0.5 mm thick; ovary tomentose, 1.5 mm long

and 1-1.2 mm broad. Fruit pale yellow (probably reddening at maturity), reddish

brown when dry, glabrous, oblong, rounded at both ends, 2—2.3 cm long, 1.5 cm

broad with the line of suture prominent; stalk 1 cm long, 2-3 mm thick. Aril

orange.

SOLOMONS

BOUGAINVILLE:

Koniguru, Buin, Kajewski 2022 (A, BM, BO, BRI, G, L, NSW. S).

SHORTLAND ISLAND:

North-east end, T. C. Whitmore’s collectors BSIP 5905 (L, SING).

East end opposite Bembalama Island, T. C. Whitmore BSIP Nos 4045 (L, SING)

and 4046 (L, SING).

138 Gardens’ Bulletin, Singapore —-X XVII (1974)

WAGINA ISLAND:

Whitmore’s collectors BSIP 5529 (L, SING).

NEW GEORGIA GROUP:

Baga Island:—Whitmore’s collectors BSIP Nos 2811 (L, SUNG); 3052 (L, SING)

and 5569 (L, SING).

Gizo Island:—Whitmore’s collectors BSIP Nos 3035 (L, SING) and 5617 (L,

SING).

Kolombangara Island:—North coast, Rei Cove, Whitmore 1537 (L, LAE, SING);

east coast in swampy forest area, Whitmore BSIP 4096 (L, SING); west coast,

Merusu Cove, Whitmore BSIP 1405 (L, LAE); Kape Harbour, flat land behind

camp along Lever’s enumeration Line, Womersley & Whitmore BSIP 803 (L, LAE,

SING).

New Georgia Island:—All from north-west part, Vaimbu River, A. W. Cowmeadow

BSIP 3679 (SING); near Jela, Whitmore’s collectors BSIP 3745 (L, SING); Kim-

bukimbu River, Cowmeadow’s collectors BSIP 3218 (SING); Lae River, Cow-

meadow’s collectors BSIP 4834 (SING).

Vangunu Island:—Ridge in forest, near Merusu Island, Whitmore BSIP 970 (L,

LAE, SING): Sosole River, J. W. P. Chapman 427 (K, SING).

Rendova Island:—Zaimane River, west coast, Whitmore BSIP 1848 (LAE, SING).

SANTA ISABEL (YSABEL):

Maringe Lagoon, near Tiratona Village in dense forest over limestone, Whitmore

BSIP 2273 (L, LAE, SING); ultrabasic ridge half a mile due west of Tatamba,

Whitmore BSIP 2582 (L, SING).

MALAITA:

Are Are District, Kiu west coast, Z. Lipageto BSIP 3406 (L, SING); 3 miles in-

land from Kiu, tributary of Wairaha River, Lipageto & Whitmore BSIP 3481 (L,

SING).

ULAWA ISLAND:

Between Haraina und Mwadoa R. Teona BSIP 6230 (SING).

GUADALCANAL:

Rere River, 3 miles inland, Lipageto BSIP 3318 (SING); forest adjacent to Tina

River, 12 miles from the coast, Whitmore & Womersley BSIP 1124 (L).

SAN CRISTOBAL:

Wairaha River, 5 miles from north coast, Whitmore BSIP 4230 (L, SING).

DIST RIBUTION. Widely distributed throughout the Solomons, endemic.

TYPE MATERIAL. T. C. Whitmore BSIP 1848 (K, LAE, SING lectotype)

Rendova Island, New Georgia Group.

The author has indicated in the M.S. that were he io find the KEW holding unsuitable

as a holotype, the SING would be preferential. Since selection is not indicated, the latter is

designated here as lectotype, Ed.

ECOLOGY. On a variety of habitats, but not in mangrove. Mostly in primary forest

on flat land, ridge tops, and river banks, dry and swampy, also in secondary

forest in well sheltered valley bottoms. Apart from alluvial, the kind of soil is not

Horsfieldia sabulosa and. H. whitmorei spp. nov. 139

often stated but limestone, ultrabasic, igneous rock and red soil have been recorded.

Therefore it may be concluded that this species is tolerant of a wide range of

habitats and is not very “‘choosy’’. Flowers during all months from September

to May with peak periods in December and April. Fruiting is from October to

May with peak periods in January and May.

VERNACULAR NAMES. Aininiu (Kwara’ae language) the usual spelling. Other

Variations are aininu, ainunu and ainynu; kisu-kisu (Buin, Bougainville). The name

aininiu is also the one in common use for H. irya.

It seems that H. whitmorei arose in the isolation of the Solomons from a

gene pool supplied by only two species, namely H. irya and spicata for certain

morphological features of both these ancestors reappear in this endemic. The

resemblances are too close for it to have been in the Solomons before the arrival

of irya and spicata or conversely also too close if we suppose that there was no

connection. These two are the oldest members of the flora and must have been

there first. They are more numerous in individuals than our present species and

also seem to have a greater geographical range occurring in nearly all the islands

of the Solomons. In fact irya has a very wide distribution throughout Malesia and

not only there but it extends from Indo-China, Siam, Burma, the Andamans and

Ceylon to Palau in the Carolines. It is the most widely collected species and the

type of the genus. It has more synonyms and references in the literature than any

other Horsfieldia. H. spicata begins in the Moluccas and also ends in Palau. It

has been recorded from numerous islands in the Bismarck group as well as in

New Guinea and it will have a wider range still if one unites it with parviflora.

One may argue that the genetical complex of whitmorei is drawn from more

species than two and that it could have come from species now extinct in the

Solomons. This is not very likely as the flora of the Solomons is poor in species

in comparison with New Guinea and Malesia. Island floras are always poorer than

those of adjacent mainlands. Whitmore on the very first page of his book, “‘Guide

to the Forests of the British Solomon Islands” deals with this subject and also

remarks about the poverty of the flora of Melanesia.

Our present species is much more than a hybrid between irya and spicata.

It is a distinct species with quite a number of differences. It probably first arose

with fertilization in the female tree of spicata by the pollen from irya. It has

retained the oblong fruits of spicata; these are identical. The male flowers are

like those of irya because of their globose shape, but being slightly larger, they

take after spicata in size; yet none are as large as those of spicata. Whitmorei

differs from them both in having no raised lines on the twigs but a faint indication

of the two lines on more than one occasion has been seen.

Sterile or atypical specimens are often troublesome to identify so it is better

to be thoroughly acquainted with details of the differences between the three.

Besides the points mentioned above the following may be helpful.

Our present species has very finely striate brick-red twigs and the lenticels (not

always present) are rather small. H. irya has blackish twigs with larger, more

numerous lenticels and the two lines from petiole base to petiole base are raised

and very distinct. H. spicata has pale straw-coloured twigs but it can have other

colours as well such as medium to dark brown and greyish shades. The striations

are generally coarser, the two lines present or absent and not usually so distinct

as those of irya. The leaves are narrow with nearly parallel sides and thus look

more like those of irya. There are some narrow-leaved specimens of spicata which

140 Gardens’ Bulletin, Singapore — X XVII (1974)

at limes are very close also to those of our present plant but they may be distin-

guished by their slightly different venation and the lack of a raised midrib in the

groove of the petiole. Whitmorei has the nerves more strictly parallel and closer

spaced than those of the other two. The nerves interarch at the margins with

greater regularity and distinctness, forming single loops. Only in very large leaves

are there double loops. They are much impressed on the upper surface of the

leaf. The other two often have double loops while the primary loops may be

faint, broken or indistinct in parts. The nerves of irya tend to leave the midrib

at a much greater angle, sometimes almost at right angles. In such cases they

curve more, ascending more gradually. Those of spicata are often more oblique

and parallel but not quite so strictly parallel and equidistant as those of our

present species. The reticulations, present on the lower surface, are scalariform.

Those of the other two form a lax network. In spicata they are not so distinct and

are often absent or seen only with a hand-lens. The lamina of all three may, at

times, show whitish marks, patches or streaks when dry, but this feature is much

commoner in irya than in the others. The inflorescence especially the male is

much branched and well developed in irya. That of the other two is simpler.

perhaps spicata would come next in order and then whitmorei, but in many cases

there is not much difference in complexity or simplicity between these two and

it would be hardly correct to put the one before the other. The globose male

flowers are half as big again as those of irya but as already pointed out not so big

as the turbinate or obtriangular ones of spicata. They are fewer in the inflorescence

when compared with an inflorescence of irya but this is only natural: being larger

they take up more room. They are cream coloured and therefore much paler than

those of irya and spicata. Finally the fruit is oblong like that of spicata, irya having

a perfectly spherical fruit. Such oblong fruits will match ones in spicata having

the same size but the fruit is of various sizes in spicata and sometimes flattened,

so it should be possible in some cases to identify the larger ones with spicata.

I must confess that without Dr. Whitmore’s valuable Solomon Islands collec-

tions I should never have properly understood this species or been aware of its

existence. Before the advent of his specimens I had only seen Kajewski 2022 to

which I had given various preliminary names including novoguineensis, irya and

spicata and Chapman 427 which I had wrongly dismissed at the time as something

different. Incidentally the Chapman specimen is a very good example of whitmorei

with very fine male flowers but it has coriaceous leaves that have dried a pale

brown. Just because of the thickness of the leaves and their unusual colour it

looks at first quite puzzling. The student will recall several instances of this sort

of thing in Horsfieldia where chartaceous leaves are replaced by coriaceous ones

of a darker or lighter colour, thus altering considerably the normal appearance

of the species e.g. glabra, polyspherula, reticulata, spicata, subglobosa and others.

A. C. Smith also named Kajewski 2022 H. novoguineensis. In fact he referred

the Brass and Kajewski Horsfieldia numbers from the Solomon Islands to this

species; but actually his novoguineensis included all three of the Solomons species,

namely, irya spicata and whitmorei.

I had therefore come to the conclusion that the Brass and Kajewski specimens

were irya and spicata but there was some doubt about the latter as I thought some

of those with narrow leaves might be palauensis. I should probably have finally

concluded that Kajewski 2022 was a narrow-leaved form of spicata but, as pointed

out, I should not from a single gathering have recognized it as a new species. The

next stage was the arrival of Whitmore’s collection and his information that there

Horsfieldia sabulosa and H. whitmorei spp. nov. 141

were three species of Horsfieldia in the Solomons. At first we both thought that the

third species might be palauensis because of its oblong fruit but I had not then

examined flowering material of the latter nor had I any available in Singapore with

which to make a comparison. In the meantime Whitmore’s book was _ published.

I did not pursue the matter any further at that time since I had then only just

started to write up Horsfieldia. In the final stages when I did eventually come

to examine whitmorei and palauensis | saw that they were not identical. The latter

is not different from the wide-spread spicata. | am grateful to the herbaria of

BISH, TI and TNS for the loan of material of palauensis.

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7 -Hottrum, R. E.: Asplenium Linn., sect. Thamnopteris Presl - - - 143-154

—:A Commentary on Comparative Morphology in Zingiberaceae - - 155-165

\ _ —: The Tree-ferns of the genus Cyathea in Borneo - . - - 167-182

_ Grex, Stew YIN ET KENG, HsuaN: Morphological studies on some _ inland

\9 _ Rhizophoraceae - - - - - - : : 183-220

y, Report of the Botanic Gardens for the year 1973 - . - - : 221-245

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Asplenium Linn., sect. Thamnopteris Presl

R. E. HOLTTUM

Royal Botanic Gardens, Kew

Summary

The status of this section (also known as a genus Neottopteris J. Sm.) is discussed.

A key to all known species is provided, with synonymy and a brief description for each.

The importance of growth-habit (not shown by herbarium specimens) is stressed, and the

need for more observation of living plants, especially on the islands of the Pacific Ocean.

Two new species (A. pacificum and A. spirale) are described. Names of species which have

been included by various authors in Neottopteris but which do not belong to this section

are listed.

This section was established by Pres] in 1836 (Tent. Pterid. 105); the sole

species was Asplenium nidus L. and the only distinctive character described was

the anastomosis of veins near the margin of the lamina. John Smith raised the

section to generic rank in 1841 (Hook. Journ. Bot. 3:409 and 4:175) with

citation also of other species. He chose a new generic name Neottopteris, and gave

the superfluous new name N. vulgaris to the type species, listing A. midus Linn.

as a synonym; this was corrected when Hooker described and illustrated the

genus in his Genera Filicum (1842) t. 113B. Presl later used the generic name

Thamnopteris (Epim. Bot. 68; 1851) and in this was followed by Beddome (Handb.

Ferns Br. India, 1883) but at the generic level Neottopteris has priority; it should

be noted also that in 1849 Brongniart published the generic name Thamnopteris

for a fossil fern stem of the family Osmundaceae.

The group of species which conform to Presl’s description is certainly a

natural one, but a decision on its status should be deferred until a scheme for a

natural division of the large genus Asplenium (estimated 700 species) is established.

On this subject Copeland made the following comments (Gen. Fil. 167; 1947): “in

a genus so large, there are of course natural groups of species, some of them easily

recognizable ............ these groups are remarkably indefinable”. He did not refer

to John Smith’s Historia Filicum (pp. 316-321; 1875) in which Asplenium is

divided into many sections and subsections. In spite of Copeland, and without

agreeing in detail with John Smith’s arrangement, it seems to me probable that

Asplenium could be subdivided into natural groups of species if someone would

take time to make in intensive survey of the whole of it, noting characters not

hitherto incorporated into specific descriptions.

John Smith (lc. p. 316) cited A. marinum L. as type species, but Copeland

(Gen. Fil. 164) cited A. trichomanes L. without mentioning Smith, whose selection

should have priority. It appears to me that a considerable number of tropical

Species including the widely distributed A. tenerum Forst. are related to A.

marinum; and in my book on the Ferns of Malaya (p. 415) I have noted the

possibility that A. nidus belongs to this alliance. Therefore it is possible that if

the genus Asplenium as now recognized were divided into several distinct genera,

143

144 Gardens’ Bulletin, Singapore — X XVII (1974)

A. nidus and near allies would remain as a section of Asplenium s. str. For this

reason I do not here recognize Neottopteris as a genus. I think it important that

a full survey of Asplenium should be undertaken, so that natural groups of species

can be found and delimited. Whether or not such groups are regarded as distinct

genera, their recognition would provide a basis for comparative studies of a kind

which are not now possible, and the distribution of the various groups would help

towards an understanding of the possible history of Asplenium sens. lat. As Cope-

land has remarked, there is a tendency within Asplenium towards the reduction of

fronds from a much-divided to a less divided condition, with the result that simple

fronds, like those of A. nidus, have originated on various evolutionary lines.

Paired sori facing each other have developed in some of these simple fronds; this

is the Phyllitis (Scolopendrium) condition. Hooker (Spec. Fil. vol. 3) put all these

species into a suborder Scolopendrieae and arranged them in four sections of

Scolopendrium (others have since been discovered); but they are the ends of

several evolutionary lines within Asplenium s. 1. and we shall not well understand

them until the species within Asplenium related to each are recognized. Section

Thamnopteris is undoubtedly a distinct section, as shown by its venation and its

distribution; it is in fact the only section of Asplenium which can at present

be clearly delimited.

The characters of sect. Thamnopteris are: caudex erect, slow-growing, un-

branched, bearing at its apex a series of almost sessile simple fronds in a very

close spiral so that together they form a nest-like structure (except A. spirale and

possibly A. longum); veins slender, parallel, close together, often forking once,

less often twice, their tips always united in a series of short arcs just within the

cartilaginous margin of the lamina; sori elongate, with narrow indusium; spores

usually with a narrow translucent wing and some anastomosing cross wings.

Distribution: tropics and subtropics of the Old World from East Africa to

Hawaii; mostly epiphytes, but a few species specialized to a habitat on limestone

rocks.

Cytology. In northern India Bir found n = 72 (tetraploid) for both A. nidus

and A. phyllitidis (Current Science 29: 445; 1960); Abraham, Ninan and Matthew

made the same observation for both species in southern India (Ind. Journ. Bot.

41 : 381; 1962). Vazart reported that a plant of A. nidus of unspecified origin,

cultivated at Paris, had n = 144 (i.e. octoploid; Rev. Cyt. Biol. Veg. vol. 17, 1956)

but his statement is not clearly expressed and needs confirmation.

The opposite key covers all species I have been able to discover. It needs

to be critically considered by people who have access to living plants, especially

on the islands of the Pacific Ocean. An important character in plants of this

section is habit of growth, especially the angle each frond makes at its base with

the vertical line of the caudex, and the curvature of the midrib. This character

is not shown by herbarium specimens, and I have seen no reference to it in taxo-

nomic works on ferns. Works on cultivated ferns sometimes describe habit, but

may be unreliable in the use of names. Schneider’s “Book of Choice Ferns” (3

volumes, 1892-1894) is still the most comprehensive of these. His published figure

of A. nidus (vol. 1 p. 618) shows a plant with the habit of A. australasicum; but

under A. nidus var. australasicum he describes the growth-habit typical of A.

nidus, though associating it with the character of a keeled midrib, a condition I

have not seen. Under A. nidus var. musifolium he states that the fronds “take an

upright direction from the start” and that the sori extend nearly to the edge;

neither statement is correct (the second probably copied from Hooker: see note on

Asplenium, sect. Thamnopteris 145

A. musifolium below). I have seen no plants in cultivation in Britain which cor-

respond to Schneider’s description of var. musifolium. Under A. nidus in Nichol-

son’s Dictionary of Gardening (vol. p. 132; 1884) and in the R. H. S. Dictionary

of Gardening (vol. 1 p. 205; 1951) are published almost identical habit-drawings

which correspond to A. australasicum. Bailey’s Cyclopaedia of Horticulture (vol. 1

p. 414, edition of 1939) gives a different habit-drawing, also corresponding to A.

australasicum but with the name A. nidus. In Exotica 3, pp 822 and 824 (1962)

A. B. Graf publishes photographs of plants which have the habit of A. austral-

asicum, though the fronds have wider basal halves than those now cultivated in

Britain and those illustrated in the British publications quoted above.

The shape of fronds, even on the same plant, is somewhat variable, especially

the shape of the apex, which is often damaged during the rather long period in

which it remains in a tender actively-growing state (it is the last part of the frond

to become hardened). Width of fronds certainly varies according to light intensity

in plants of A. nidus in Malaya, as I have proved experimentally by taking a

plant with narrow fronds from the weak light in which it was growing to a stronger

light; the next frond was more than twice as wide as preceding ones. This is

also true of plants in cultivation at Kew. But my impression is that in some other

species (A. phyllitidis, and probably A. australasicum) this response to stronger

light does not occur; in Malaya, A. phyllitidis does not grow in such exposed

positions as A. nidus.

As herbarium specimens do not clearly indicate the curvature of the base of

the frond, even if the base is present (often it is not), one object of the present

account is to call attention to the need for field observation of the habit of

growth. John Smith recognized this a century ago when he wrote: “The cultivated

examples of four forms known to me, although difficult to recognize as distinct

when put in the Herbarium, are however readily seen to be distinct species in

the garden” (Historia Filicum, p. 329). He was implicitly criticizing W. J. Hooker,

who did not pay sufficient attention to the living plants under John Smith’s care

at Kew.

KEY TO THE SPECIES OF SECT. THAMNOPTERIS

1. Fronds gradually narrowed from their widest part to the base

2. Sori on almost all veins, including both branches of a vein which

forks near costa; indusia less than 4 mm wide

3. Fronds nest-forming

4. Fronds ascending steeply; nest narrowly funnel-shaped

5.) 0@ostasharply keeled. of lower surface ............2.....0:+

1. A. australasicum

5. Costa not keeled on lower surface ...... 2. A. pacificum

4. Fronds spreading horizontally at base and then curved

upwards

6. Fronds to c. 20 cm wide; apex gradually attenuate

Bee a Sue RM he cod Wa ceeds SEMA Meets Baia Sc ae 3. A. nidus

7. Fronds unlobed and fiat

146

| pe

i &

Gardens’ Bulletin, Singapore — X XVII (1974)

8. Sori usually much more than 10 mm long

9. Sori on basal 4-4 (rarely %) of each vein

Be iets Sao, Lot geht ee var. nidus

9. Sori on middle part of each vein .........

“AMAA bGinilte wate HUES MIGETS Merah REREAD var. biseriale

8. Sori not. more .than-10 mm long. .....)........

b eapee totaee tks phi coceree Mee ones var. curtisorum

7. Fronds much lobed, or plicate

£0: . “Bronds’ miteh Jobed 7th var. multilobum

tO!’ SS rons “puCale te. = kere errr var. plicatum

Fronds 25-35 cm wide; apex broadly rounded .........

FA NI RR hak Orlin he yt et eB <a e 4. A. musifolium

Fronds not nest-forming

Fronds:toat least! 70)x010%enn 2). MIRC Oe. 5. A. longum

Fronds tO C2 2) KD Cl ance ete es renee 6. A. spirale

Sori usually on one branch of each vein, indusium more than 4 mm

wide

12.

Fronds to 20 cm or more wide with dilated bases; indusia

more:tuan i-nim wide)... c.a6.: eee 7. A. cymbifolium

Fronds not over 12 cm wide, bases not dilated; indusia less

than 1 mm wide

13.

Fronds of mature plants to at least 9 cm wide; costa little

or not prominent on lower surface

14. Wing of spores with short broad projections; sori

not more than-half leneth' of vems s:.2524 ee

PRIA, DAA 8 | QOS at Biro ate 8. A. carinatum

14. Wing of spores uniformly narrow; sori ? length of

VOUS ois cak meee es eee 9. A. antiquum

Fronds of mature plants in many cases narrower; costae

distinctly prominent on lower surface

15. Fronds 3-4 cm wide; sori lying in a depression in

the dried lamina, the depression with a distinct

Outer edge™~), BLN), . 81207 sean 10. A. colubrinum

15. Fronds in some cases wider; sori not lying in a

depression

16. Fronds to 3 cm wide; scales on stipe not more

than 5 mm long; sori at c. 45° to midrib

Casas alee toe. al ob bao 11. A. simonsianum

16. Fronds commonly 4-8 cm wide (exceptionally

wider); scales 10 mm or more long; sori

usually at a wider angle ... 12. A. phyllitidis

17. Wings of spores of uniform width;

fronds never more than 8 cm _ wide

Ss SAPP Fs, ALO subsp. phyllitidis

Asplenium, sect. Thamnopteris 147

17. Wings of spores bearing many slender

projections equal in length to more than

4 width of spore; fronds sometimes to

PORCTO OP MIOGS WIDE 6. acs: is <oLgsmeen stone’

1. Fronds with basal half or more narrow and sterile, abruptly widened to

fertile distal half

18. Rhizome-scales c. 5 x 14 mm: costa keeled on lower surface

Oe le ee Pek sah «Weiss org a POS ea eentnnhenbeur 13. A. grevillei

18. Rhizome-scales 3-4 mm wide, little longer than wide: costa not

keeled on lower surface

19. Sterile base of frond 14 cm wide at base, widening to 24 cm

ae PB A shia ask 14. A. antrophyoides

19. Sterile base of frond a very narrow wing, grading impercep-

tibbyl-tel da Stipe sialisse. . coed .prses,.. zhi. 15. A. humbertii

1. Asplenium australasicum (J. Sm.) Hook., Fil. Exot. (1859) t. 88; Mett., Farn-

gatt. VI (1859) 85. — Neottopteris australasica J. Sm., Cult. Ferns (1857) 49;

Hist. Fil. (1875) 330. — Thamnopteris australasica (J. Sm.) Moore, Ind. Fil. (1859)

115. Type: cult. Hort. Bot. Kew., origin Australia (BM, in herb. J. Sm.).

Fronds to c. 100 x 15 cm, ascending at a small angle to vertical, together

forming a funnel-shaped nest; frond widest just above middle, gradually narrowed

towards base; midrib strongly and acutely keeled on lower surface: veins, sori

and spores as in A. nidus, but sori often occupying more than half of the length

of a vein. Hooker figures the base of the frond somewhat dilated, but this is not

always so.

Distribution: Queensland and New South Wales; Norfolk Isl., Lord Howe

Isl., New Caledonia, Fiji, Samoa, Tonga, Tahiti.

2. Asplenium pacificum Holtt., sp. nov.

Ab Asplenio nido differt: frondibus e basi suberectis, congerie frondium ut

in A. australasicum infundibuliformi, sed costis subtus latis, paulo elevatis, non

carinatis.

Largest frond seen 60 x 12 cm, widest about middle, gradually tapering to

rather wide base and more narrowly to apex; small scales on lower surface very

few, irregularly stellate with some dark-walled cells in the centre, terminal cells

of arms not elongate; costa dark, broad, slightly prominent and rounded on lower

surface, strongly prominent on upper surface; sori on all veins, from near midrib,

extending a little more than half-way towards margin.

Type: cult. Hort. Bot. Kew., accession no. 584/65, grown from spores

collected on Washington Island by W. A. Sledge (K).

3. Asplenium nidus L., Spec. Pl. (1753) 1079; Holtt., Rev. Fl. Mal. 2 (1955)

419. — Neottopteris vulgaris J. Sm. in Hook. Journ. Bot. 3 (1841) 409; 4 (1841)

175, nom. nov. superfl. — Neottopteris nidus (L.) Hook., Gen. Fil. (1842) t. 113B:

Brack. in Wilkes U.S. Expl. Exp. 16 (1854) 175. — Thamnopteris nidus (L.) Presl,

Epim. Bot. (1851) 68: Bedd., Handb. Ferns Br. India (1883) 137. Type: Osbeck

s.n., Java (LINN).

Asplenium ficifolium Goldm., Nova Acta Acad. Leop. Carol. 19, Suppl. 1

(1843) 461. Type: Meyen s.n., Manila (B, not seen).

148 Gardens’ Bulletin, Singapore — XXVII (1974)

Neottopteris stenocarpa Fée, Gen. Fil. (1852) 203. Type: Cuming s.n., Philip-

pines (not seen).

Neottopteris elliptica Fée, Gen. Fil. (1852) 203. Type: Cuming s.n., Philippines

(not seen).

Neottopteris mauritiana Fée, Gen. Fil. (1852) 204, — Thamnopteris mauritiana

Pres], Epim. Bot. (1851) 68, nom. nud. Type: Sieber, Fl. Maurit. 46 (isotype K).

Neottopteris rigida Fée, Gen. Fil. (1852) 203. Type: Henslow s.n., Macao

(not seen).

Neottopteris salwinensis Ching, Bull. Fan. Mem. Inst. Biol. Ser. 2, 1 (1949)

304. Type: T. T. Yu 19163, N. W. Yunnan, Salwin valley (PE, not seen).

Var. nidus. Fronds to about 150 x 20 cm, narrowed gradually to base and apex,

at base growing out horizontally and curved gradually upwards; costa broad but

only slightly prominent (not keeled) on lower surface, strongly prominent and

+ 2-angled on upper surface except near base; small scales on lower surface of

lamina of young fronds, near base, stellate with black-walled cells in centre,

end-cells of rays pale, short, thin-walled; sori on all veins, usually extending

from costa not more than 4 way towards edge; indusia less than 4} mm wide.

Distribution: E. Africa (Dar es Salaam, Tanzania Zanzibar); Madagascar,

Mascarene Islands, Seychelles, Ceylon & S. India, N. E. India; Indo-China & S. W.

China; Malesia; Solomon Islands to Tahiti and Hawaii.

Throughout much of its range, var. nidus is represented by abundant plants

which grow in fairly exposed positions on branches of trees; these plants vary

much in size, according to habitat conditions. Though I have not seen the types

of Fée’s species which are placed above as synonyms, I do not think that they

are distinguishable entities; they differ in size and shape of fronds, length of

sori etc, all very variable characters. In the Pacific region it appears that A. nidus

and A. australasicum both occur in Samoa, and there the variation is greater

than in the Malayan region. In the 1880’s Thomas Powell sent many specimens

to Kew, with annotations in which he attempted to distinguish several distinct

varieties or species, but not all his specimens were kept, and those at present in

the herbarium at Kew do not appear to me to show enough recognizable characters

to establish new names. I have described var. biseriale from one collection of

Powell’s (see below) which he pointed out as most distinctive. He also stated that

he regarded A. australasicum as distinct. It is possible that some of the variation

in Samoa is due to hybridization between A. nidus and A. australasicum; a new

local survey of these plants might yield interesting results. A similar survey is

also desirable in other island-groups of the Pacific.

Var. bieriale Holttum, var. noy.; a varietate typica differt soris media venarum

occupantibus, nec costam nec marginem accedentibus, lamina basi leviter dilatata.

Type: T. Powell 240, Samoa (K). See above discussion.

Var. curtisorum (Chr.) Holtt., var. nov. — Asplenium curtisorum Chr., Ann. Jard.

Bot. Btzg 19 (1904) 39. — Neottopteris curtisora Hosokawa, Trans Nat. Hist. Soc.

Formosa 31 (1941) 474. Type: Sarasin 2034, Gimpu, Central Celebes, 5 Sept.

1902.

The original specimens was described as having fronds 50 x 9 cm, short-

acuminate, with sori about 10 mm long; there is no information about width

of indusia. A report from the University Herbarium at Basel states that the type

is not with the earlier Sarasin collections from Celebes; it may be at Paris. In Kew

Asplenium, sect. Thamnopteris — 149

herbarium are specimens, as cited below, from Sabah and Sarawak which conform

to the above description, also a plant from Sabah in cultivation at Kew. But there

are other specimens from Borneo with sori 12-15 mm long, and I am not sure

that var. curtisorum is sharply distinct.

Price’s specimen from Mindanao has fronds 17 cm wide and sori 8-10 mm

long close to the midrib. The collector reports that when fresh the upper surface

of the midrib was prominent and rounded in section, not 2-angled as it usually

is in A. nidus, but the dried specimens do not show this clearly so that a com-

parison with the Bornean specimens cannot be effectively made.

SABAH. Elmer 20943, Tawao. Cult. R. B. G. Kew, accession no. 455/63,

origin Sabah (no locality) leg. Giles & Wolliams. SARAWAK. Clemens 21011, Mt

Matang. Brooks s.n. Aug. 1909, without locality (BM). JAVA. Donk s.n. 5.4.1950,

near Tjibodas (K).

PHILIPPINES, M. G. Price 2752, Mindanao, Agusan del Norte Prov., mountains

above Cabadbaran.

Var. multilobum F. M. Bailey, Queens]. Agric. Journ. 1 (1897) 370, with photo.

Type: Bailey s.n., Trinity Bay Ranges, N. Queensland (isotype K).

Fronds irregularly and deeply lobed; midrib pale in distal part (apparently

dark near base), rather broad and rounded on lower surface: sori irregular, in

some cases in pairs facing each other, as in Phyllitis.

It is possible that this should rank as a variety of A. australasicum; the Kew

specimen consists of the apical part of the frond, and has slight indications of a

keel on the lower surface of the midrib, which however is very broad, due probably

to the frequent branchings. I have seen no later specimens, and have no informa-

tion whether this variety persists.

Var. plicatum v.A.v.R., Handb. Malayan Ferns (1908) 440. Type: not cited.

apparently at BO, from a cultivated plant.

Plant rather small, with lamina on each side of the midrib plicate or crimped

so that it will not lie flat. A plant of this nature in cultivation at Kew has very

thick rigid fronds; its origin is not recorded.

4. Asplenium musifolium Mett., Farngatt. VI (1859) 86; Hook., Spec. Fil. 3

(1860) 78, p.p. — Thamnopteris nidus var. musifolia Bedd., Handb. Ferns Br.

India (1883) 139, p.p. — Asplenium nidus var. musifolium v.A.v.R., Handb. Mal.

Ferns (1908) 439, p.p.; Holttum, Rev. Fl. Mal. 2 (1955) 418. Type: Cuming 89,

Luzon p.p. (isotype K).

Asplenium nidiforme v.A.v.R., Bull. Jard. Bot. Btzg II, 7 (1912) 6; Handb.

Suppl. (1917) 281. Type: Cult. Hort. Bog. II K XIV 39, origin New Guinea (BO).

Fronds growing out horizontally at the base, then curved upwards as in

A. nidus, those of well-grown plants 25-35 cm wide, base gradually and then

abruptly narrowed to a black stipe, apex rather abruptly and broadly rounded

with a short mucro; costa rather strongly prominent on lower surface but not

keeled; sori as in A. nidus.

Extreme forms of this look distinct from A. nidus (small plants also have

proportionately broad fronds), but it is desirable to establish its distinctness by

growing plants from spores.

Cuming’s collection no. 89, on a specimen from which Mettenius based the

first published description, was a mixed gathering, part being as described above

150 Gardens’ Bulletin, Singapore — X XVII (1974)

and part being from a plant of A. cymbifolium (no. 7 below). Mettenius’s specimen

is probably lost with the destruction of the Leipzig herbarium, but his description

leaves no doubt as to its identity, matching exactly a specimen of no. 89 in the

Kew herbarium. Hooker’s description of 1860 covers both species, including the

statement that in one “form” the sori may reach almost to the edge of the lamina

(this “form” was A. cymbifolium), and this was copied by Beddome and van

Alderwerelt. John Smith first published the name as Neottopteris musifolia, citing

Cuming 89, but without a description; he wrote the name N. musifolia on a speci-

men of A. cymbifolium in his herbarium, and may have intended it for that species,

which is strikingly distinct, but the type of A. musifolium must be the specimen

described by Mettenius.

The type specimen of A. nidiforme, of which I have seen a photograph, has

fronds to 50 x 124 cm, and I judge that it was an immature plant; the shape of

frond is like that of A. musifolium.

A specimen from the Admirality Islands (Challenger Expedition, 1875) in

the BM herbarium consists of a frond of the form of A. musifolium, 22 cm wide,

but the midrib is decidedly keeled on the lower surface, as in A. australasicum.

The curvature of the base of the frond cannot be judged. This is another indica-

tion of the complexity of sect. Thamnopteris in the Pacific, and of the need for

more observation.

A plant in cultivation at Kew, from N. E. New Guinea (accession no. 247/70,

2351) has fronds of about the same size and shape as the type of A. nidiforme, but

it has small scales on the lamina as in A. phyllitidis, and the spores are abortive:

it is probably a hybrid.

5. Asplenium longum v.A.v.R., Bull. Jard. Bot. Btzg II, 28 (1918) 8. Type:

Biinnemaijer 2131, Bangka, Mt Mangkol (BO; phot. at BM). — Asplenium per-

longum v.A.V.R., lc. Type Biinnemeijer 1846, Bangka, Mt Maras (BO; Phot.

at BM).

“Not nest-forming” (v.A.v.R.); fronds of type of A. longum 70-80 x 10-15

cm, of A. perlongum to 145 cm long; sori of former not touching costa, of latter

touching costa; little other clear difference. The photographs of type specimens

do not indicate how the growth habit differs from that of A. nidus, nor is there

clear distinction in form of fronds and sori. I place this species here to call

attention to its description, in the hope that further information about plants on

the island of Bangka may be obtained.

6. Asplenium spirale Holtt., sp. nov.

Caudex elongatus, c. 8 mm diametro, paleis nigris angustis c. 8 mm longis

vestitus, frondes laxe spiraliter ferens. Stipes anguste alatus, 1-2 cm longus, lamina

rigida, 18-24 cm longa, 4.2-5.0 cm lata, leviter oblanceolata, basi subabrupte

ad alam angustata, apice late vel anguste acuta, non cuspidata; costa utrinque

paulo elevata; venae more A. nidi prope marginem anastomosantes, c. 1 mm

inter se distantes, sub angulo c. 45° a costa abeuntes; sori e costa dimidio marginem

versus producti, indusia vix 4+ mm lata; sporae alis angustis praeditae.

Type: Pulle 184, Lorentz River, W. New Guinea, 30 m (BM).

The fronds are shaped about as in A. musifolium but are small and rigid; they

are attached to the caudex in an ascending spiral arrangement so that about 8

fronds are attached to a length of caudex 5 cm long. Thus the habit is quite

distinct from that of A. nidus.

ec ttl ALLL LLL

Asplenium, sect. Thamnopteris 151

7. Asplenium cymbifolium Chr., Bull. Herb. Boiss. II, 6 (1906) 999; v.A.v.R.,

Handb. (1908) 440; Copel., Philip. Journ. Sci. 2C (1907) 130; Fern FI. Philip.

(1960) 451. — Neottopteris cymbifolia (Chr.) Tagawa, Journ. Jap. Bot. 22 (1949)

161. Type: Loher s.n., Mt Makiling, Luzon (P, not seen).

Fronds similar in size and shape to those of A. musifolium but broader and

dilated in basal part; veins about 2 mm apart in middle of lamina; sori not on

every vein, rather irregular in spacing and length, many extending almost to edge

of lamina: indusia a little more than 1 mm wide. Scales on rhizome up to 4 mm

wide: rather pale: small scales also abundant on edges of costa near base of frond.

Distribution: S. Sumatra, Borneo, Philippines, New Guinea, Samoa.

As noted above under A. musifolium, Cuming included specimens of this

species under his no. 89. Mr Michael Price informs me that the species grows in

the mossy forest near the summit of Mt Makiling. and that the bases of the fronds

overlap so closely that the “nest” holds water. Roots of the fern grow up into the

water, and tree-frogs lay eggs in it.

Forma lingganum v.A.v.R., Bull. Jard. Bot. Btzg III, 5 (1922) 184. Type:

Biinnemeijer 7388, Lingga Arch., P. Singkep (BO: isotype K). Fronds to 60 x 6

cm, base dilated as in type of species: sori short. This may be a young plant. It

appears to have been found in mangrove, where high average atmospheric humi-

dity encourages abundant epiphytic growth on certain species of trees in quite

bright light: some of these species do not occur in normal lowland forest in Malaya,

but occur further north on mountains (e.g. Ctenopteris moultonii (Copel.) Holtt.)

8. Asplenium carinatum v.A.v.R., Bull. Jard. Bot. Btzg II, 28 (1918) 9, Type:

Teysmann 12791, Maros, S. W. Celebes (BO).

Fronds 40-50 x 9 cm: apex rather abruptly short-pointed to cuspidate; lower

part of frond gradually narrowed towards base, a very narrow wing extending

almost to attachment of frond to caudex; costa strongly keeled on upper surface.

not on lower; veins at c. 60° to costa: sori from costa along basal half of veins, or

shorter, on alternate veins; indusium more than } mm wide; spores with wing of

irregular width, not produced into filaments as in A. p/yllitidis subsp. malesicum.

Only known from two collections from the type locality: the second is

C. J. Brooks 16856 (BM), found growing on limestone near Maros Waterfall. This

habitat is interesting, in view of the fact that species 13-15 of the present account

occur only on limestone in mainland Asia.

9. Asplenium antiquum Makino, Journ. Jap. Bot. 6 (1929) 32: Tagawa, Col. IIl.

Jap. Pterid. (1962) 147, 177, fig. 324. — Thamnopteris antiqua Makino, Journ. Jap.

Bot. 8 (1932) 7. — Neottopteris antiqua Massumune, Trans. Nat. Hist. Soc. For-

mosa 22 (1932) 215. Type: “Hab. Japan, southern”, not seen.

Fronds to c. 100 x 12 cm, narrowed gradually to base and more abruptly to

apex: costa broad and only slightly prominent on lower surface, little more so

On upper: veins in middle of lamina more than 1 mm apart: sori on alternate

veins, 4 or more of length of veins; indusia more than } mm wide; spores winged

as in A. nidus.

Distribution: Kyushu (Yakushima Isl.), Izu Isl., Okinawa.

10. Asplenium colubrinum Chr., Bull. Herb. Boiss. II, 6 (1906) 999; Copel., Fern

Fl. Philip. (1960) 451. Type: Loher s.n. March 1906, Angilog, Luzon (P?: isotype

K). Also cited Loher 18.4.1905, Mt Batay, 1380 m.

1$2 Gardens’ Bulletin, Singapore — XXVII (1974)

Fronds to almost 100 cm long, 3-4 cm wide, firm texture, rather uniform in

width for a great part of their length, narrowed gradually to both base and apex

which is sometimes short-acuminate; costa keeled on lower surface, not on upper:

sori on alternate veins, occupying } or more of length of veins, the sporangia lying

in a depression in a dried frond, the edge of the depression distinct; indusia a

little more than + mm wide; spores with a narrow wing.

Distribution: Luzon, Mindoro.

Copeland described a variety faenicphyllum (Philip. Journ. Sci. 2C: 131;

1907) based on a specimen collected by Merrill (no. 5900) on Mt Halcon, Mindoro.

He described the sori as occupying 3-2 of the veins, with black indusia. I have

not seen a specimen, but have seen another from Mt Halcon (Ramos & Edano

40623) which does not differ significantly from Loher’s Luzon specimens.

Mr M. G. Price has recently sent a specimen (no. 440) from Mt Polis, Luzon,

which agrees with the above description except that the sori do not lie in a

depression in the surface of the frond. The fronds of this specimen are thinner

than those of the type. The species is thus perhaps chiefly distinguished by its long

narrow fronds.

11. Asplenium simonsianum Hook., Ic. Plant. 10 (1854) t.925. — Thamno-

pteris sitmonsiana (Hook.) Moore, Ind. Fil. (1857) L: Bedd., Handb. Ferns Br.

India (1883) 141. — Neottopteris simonsiana J. Sm., Hist. Fil. (1875) 330. Type:

Simons 232 (err. typ. 432 in Hook. l.c.), Khasya (K).

Scales dark, firm, 2-3 x 1 mm; stipe c. 2 cm long; frond to 40 x 3 cm, widest

1 from apex, very gradually narrowed towards base, basal part a very narrow wing

on each side of costa, apex rather abruptly narrowed to a cusp 15-20 mm long,

15-2 mm wide; costa prominent on both sides near base, rounded on lower side,

on upper side grooved at base, above base raised and 2-angled when dry; veins at

c. 45° to costa, distinct but not or little prominent either side; sori from near

costa 3} towards edge, usually with a sterile vein between sori; indusia over + mm

wide when young; spores with translucent narrow somewhat crisped wing.

Distribution: Assam, Sikkim, Chittagong, Vishakapatanam, on hills at about

3000 ft.

12. Asplenium phyllitidis Don, Prodr. Fl. Nepal. (1825) 7; Hook., Spec. Fil. 3

(1860) 80. excl. syn. A. simplex Bl. — Neottopteris phyllitidis (Don) J. Sm. in

Hook. Journ. Bot. 3 (1841) 409. — Thamnopteris phyllitidis (Don) Pres], Epim.

Bot. (1851) 68; Bedd., Ferns S. India (1863) t. 123. — Thamnopteris nidus var.

phyllitidis Bedd., Handb. Ferns Br. India (1883) 139. — Asplenium nidus var.

phyllitidis v.A.v.R., Handb. Suppl. (1917) 282: Tard. & C. Chr. in Fl. Gen.

Indochine 7, pt. 2 (1940) 220. Type: Wallich, Nepal (not seen).

Thamnopteris orientalis Presl, Epim. Bot. (1851) 69; Holtt., Novit. Bot. Inst.

Bot. Univ. Carol. Prag. 1968 (1969) 52. Type: Wallich 198, Nepal ? (PRO).

In the Wallich Herbarium at Kew, there are 9 sheets of 198, not all from

Nepal; only two agree with Presl’s specimen.

Subsp. phyllitidis

Scales at least 10 mm long; fronds commonly 40-70 cm long, 4-8 cm wide,

widest above middle, gradually narrowed to base, apex rather narrowly pointed or

short-cuspidate; costa prominent and rounded on lower surface, on upper surface

grooved near base, above base slightly prominent and 2-angled; veins c. 14 mm

Asplenium, sect. Thamnopteris 153

apart mid-way between costa and edge, uniting 1 mm or more from edge; sori

from near midrib 4-3 length of veins, usually on acroscopic branch of a vein

which forks near costa, at an angle of more than 45° to costa; indusia more than

1 mm wide: spores with a narrow wing.

Distribution: S. India, N. E. India, N. Thailand, Yunnan, Hainan.

Subsp. malesicum Holttum, subsp. nov., a subspecie phyllitidi differt frondibus

saepe majoribus, usque 12 (-15) cm latis, sporis semper ciliatis. Type: Cuming 319,

Samar (K).

Asplenium nidus var. simonsianum sensu Chr., Bull. Herb. Boiss. 6 (1898) 151.

Asplenium oblanceolatum Copel., Philip. Journ. Sci. 9C (1914} 229. Type:

Brooks 28, Lebong Tandai, Benkoelen, Sumatra (BM).

Asplenium phyllitidis sensu Holtt., Rev. Fl. Mal. 2 (1955) 420; sensu Copel.,

Fern Fl. Philip. (1960) 449.

Asplenium simplex sensu Kunze, Bot. Zeit. 6 (1848) 145; sensu Mett., Farn-

gatt. VI (1859) 86; non B1.

Thamnopteris simplex sensu Presl, Epim. Bot. (1851) 69, non (B1.)

Fronds commonly to 100 x 8 cm, sometimes to 10-12 (—15) cm wide; sori

varying in length, usually 4-2 length of veins but sometimes shorter; spores always

with a translucent wing which is produced into many slender points equal in length

to more than half width of spores; scales on lower surface of young fronds, near

base, very small, middle cells with pale walls, fringed with elongate orange glan-

dular cells (these scales not seen on specimens of subsp. phyllitidis).

Distribution: throughout Malesia: in mainland Asia northwards to southern

Burma and Thailand and to Tonkin.

In New Guinea there appears to be more variation than elsewhere. Two

distinct forms are in cultivation at Kew, but herbarium specimens from New

Guinea are in some cases intermediate. In one cultivated plant the nest has a very

wide open base, fronds of a yellow-green colour, sori not over % length of veins

at a rather broad angle to costa, small scales on lower surface near base abundant:

the other cultivated plant has more erect fronds (forming a less open nest) which

are darker green, with sori almost the full length of the veins and at about 45°

to costa, superficial small scales fewer with glandular cells less conspicuous.

13. Asplenium grevillei Wall. ex Hook. & Grev., Ic. Fil. (1831) t. 228. Tard. &

C. Chr. in Fl. Gen. Indoch. 7, pt. 2 (1940) 217. — Neottopteris grevillei (Wall.)

J. Sm. in Hook. Journ. Bot. 3 (1841) 409. — Thamnopteris grevillei (Wall.) Moore,

Ind. Fil. (1857) L; Bedd., Handb. Ferns Br. India (1883) 139. Type: Wallich

1036, Tavoy, Burma (K).

Scales not well seen, apparently to 5 x 14 mm, firm but thin; stipe very short:

frond to 45 cm long, basal 20-25 cm a narrow wing on each side of costa, up to

1 cm wide or little more, rather abruptly widening to 44-6 cm wide in distal

part; texture thin; costa keeled on lower surface; veins c. 1 mm apart, at c. 60°

to costa; sori from near costa ? or more towards edge; spores translucent with

a narrow wing.

Distribution: Peninsular Burma and Peninsular Thailand: S. India and

Ceylon.

154 Gardens’ Bulletin, Singapore — XXVII (1974)

14. Asplenium antrophyoides Chr., Bull. Geogr. Bot. Mans 20, pt. 1 (1909) 170,

Tard. & C. Chr. in Fl. Gen. Indoch. 7, pt. 2 (1940) 218 — Neottopteris antrophyoides

(Chr.) Ching, Bull. Fan Mem. Inst. Biol. Bot. 10 1941) 7. Type: Cavalerie 1877,

Sept. 1907, Kweichow, Lo-fou (P).

Asplenium subspathulatum Rosenst. in Fedde. Rep. 13 (1913) 122. Type:

Cavalerie s.n. Kweichow (isotype BM).

Scales c. 4 x 4 mm; fronds to c. 40 x 7 cm, spathulate, abruptly cuspidate,

sessile; basal 15-17 cm gradually widening from base to 24 cm wide, then less

abruptly than in A. grevillei to the wide distal part: veins in broad part of frond

c. 14 mm apart: sori from 3-5 mm from costa to 3 distance to edge, at c. 60°

to costa; spores not translucent, with a fairly wide somewhat crisped wing.

Distribution: Kweichow, Tonkin, N. Thailand, on limestone.

15. Asplenium humbertii Tard., Aspl. du Tonkin (1932) 25, t. 2, f. 1, 2. Tard. &

C. Chr. in Fl. Gen. Indoch. 7, pt. 2 (1940) 218. — Neottopteris humbertii (Tard.)

Tagawa, Journ. Jap. Bot. 22 (1949) 161. Type: Balansa 68, Tonkin, Than Moi,

sur les rochers calcaires, Jan. 1886 (P: K).

Scales as in A. antrophyoides; basal part of frond very narrow, 15-20 cm

long, apical fertile part to 23 x 4.8 cm, apex abruptly cuspidate; sori from

near costa along 4—} of veins.

Distribution: Tonkin, North Thailand, on limestone.

This species is very near A. antrophyoides, but appears to be distinct in the

very narrow basal part of the frond. Specimens in Kew herbarium from N. Thai-

land are variable in the angle between sori and costa.

EXCLUDED SPECIES WHICH HAVE BEEN INCLUDED IN Neottopteris

Neottopteris squamulata (Bl.) Fée, Gen. Fil. (1852) 203 — Asplenium squamu-

latum B1., Enum. Pl. Jav. (1828) 174. — Thamnopteris squamulata (Bl.) Presl,

Epim. Bot. (1851) 260. Type: Blume, Java (L).

Neottopteris pachyphylla (Kunze) Fée, Gen. Fil. (1852) 203. — Asplenium pachy-

phyllum Kunze, Bot. Zeit. 6 (1848) 146. — Thamnopteris pachyphylla (Kunze)

Pres], Epim. Bot. (1851) 69. Type: Zollinger 2414, Java (isotype K). = A.

squamulatum Bl.

Neottopteris simplex (Bl.) Fée, Gen. Fil. (1852) 203. — Asplenium simplex Bl..

Enum. Pl. Jav. (1828) 174, non Kunze Bot. Zeit. 6 (1848) 145, nec Mett. Farngatt.

VI (1859) 86. — Thamnopteris simplex (Bl.) Presl, Epim. Bot. (1851) 69. Type:

Blume, Java (L.) — A. amboinense Willd. sensu Backer & Posth., Varenfi. Java

(1939) 133, but doubtfully identical with Willdenow’s species, the type of which

is a Sterile frond.

Neottopteris taeniosa (Kunze) Fée, Gen. Fil. (1852) 203. — Asplenium taeniosum

Kunze, Bot. Zeit. 6 (1848) 145. Type: Zollinger 2823, Java (isotype K). = A.

simplex BI.

Neottopteris stipitata J. Sm. and N. ovata J. Sm. are nomina nuda which have

been cited by other authors but have no status.

A Commentary on

Comparative Morphology in Zingiberaceae

R. E. HOLTTUM

Royal Botanic Gardens, Kew

Summary

A critical commentary is presented on the statement on comparative morphology of

Zingiberaceae in Malaya published in Gard. Buli. Sing. Vol. 13, part 1 (1950), and a

comparison made with the account of the family prepared by Bakhuizen for the Flora of

Java Vol. 3 (1968) where terms are differently used and a different view of some genera

(Achasma and Amomum) is maintained. An attempt is made to correct inconsistencies in

the use of terms in both accounts, especially in relation to inflorescence-structure which is

of basic importance in this family. Floral morphology is only considered in reference to

the genera Nicolaia (mis-named Phaeomeria in Holttum 1950), Achasma and Amomum.

Some comments are made on the status of some generic names but no proposals for change

of the scheme of 1950 are made. A suggestion is made that experimental work might throw

light on the structure of the condensed lateral cymes and in demonstrating the essential

uniformity of structure of ihe inflorescence in the family, A plea is made that taxonomic

statements on tropical plants should no longer be limited by the defects of dried specimens

in European herbaria.

Introduction

In Singapore, during World-war Il, I prepared a systematic study of the

family Zingiberaceae in Malaya, based on all available records in the herbarium

and library at the Botanic Gardens, on a considerable number of species in

cultivation and native in Singapore island, and on inflorescences of many species

preserved in alcohol, the majority collected, with elaborate field notes, by

E. J. H. Corner. In this study I was greatly helped by the published works of

Valeton, who also had (at Bogor) a wide range of living plants at his disposal

and was able to see clearly structures which are not well preserved in dried

specimens, and thus were not considered, or not understood, by herbarium

botanists, notably J. G. Baker (1892) and K. Schumann (1904). After the war, in

the years 1946-1949, I was able to examine a considerable number of living plants,

some of the species not seen previousiy, in various parts of Malaya, and thus to

improve my manuscript. I made drawings in all cases from living plants, with

use of a camera lucida.

In the Flora of Java Bakhuizen expressed disagreement with my opinions

(and those of Valeton on which mine were largely based), quoting a statement

of mine which he believed to be contradictory. This led me to re-read what I

wrote in 1950, and in so doing I found some inconsistencies in the use of terms.

I have therefore decided to re-write my statement on morphology, especially of

the inflorescence, to remove inconsistencies, and to show where I think Bakhuizen

155

156 Gardens’ Bulletin, Singapore — X XVII (1974)

has misunderstood not only my statement but also some specimens at his disposal.

I judge that a large part of his difficulty was due to the fact that he had little

or no living material available of most species. It is difficult or impossible to

understand the detailed structure of the inflorescence of most Zingiberaceae from

dried specimens unless such specimens have been specially prepared to demon-

strate the significant characters. I have rarely seen specimens which do this

effectively. However, the taxonomic study of tropical plants ought not to be

limited by the imperfections of dried specimens.

Since Bakhuizen’s publication, Burtt and Smith (1972) at Edinburgh have

produced a most valuable critical report on the early history of taxonomic study

of the family and on the significance of published generic names, accompanied

by an excellent series of drawings. Burtt and Smith did not arrive at a conclusion

as to the application of all the generic names, and in part such application

depends on how genera are delimited. As I pointed out in 1950, I had no means

of discovering the correct names for some of the groups I recognized as genera.

I used the generic names which involved the least name-changes, and was chiefly

concerned to characterize clearly the groups in question. In the present contribu-

tion I do not propose any changes in generic concepts, but I do offer some

comments on the status of Achasma and Nicolaia (Nicolaia should replace the

invalid name Phaeomeria of Holttum 1950).

Growth-habit

This is invariably sympodial, each new aerial stem arising from a bud at the

base of the preceding stem. The new growth is first, for a short or longer distance,

more or less horizontal and root-bearing, then erect. Erect leaf-bearing stems may

have a terminal inflorescence, or flowering and leafy stems may be distinct from

each other.

All leaves have a sheathing base, and a blade on a (usually short) stalk; the

sheath is produced upwards above the attachment of the stalk into a ligule, so

that the stalk appears to be a dorsal organ attached near the top of the sheath.

At the base of every stem, whether it bears fully developed leaves or not, are

sheaths lacking leaf-blades. These are successively longer, and they often bear

the rudiment of a blade just below the apex; the blade on the first blade-bearing

sheath is small, later ones successively larger. The difference between leafy stems

and flowering stems, where the two are different, lies solely in the fact that no

blade-bearing sheaths are developed on the specialized flowering stems, which are

covered with 2-ranked sheaths, often more numerous than those at the base of

leafy stems but not otherwise different. In my work of 1950 I used the term

sheath for these imperfect leaves. They might also be called cataphylls, but I

preter sheath because they represent exactly the sheaths of normal leaves. Bakhui-

zen calls them scales, but they are very different from the scale-like leaf-rudiments

which cover the resting buds of dicotyledons. Exactly similar sheaths are found |

at the bases of new stems in many other families of Monocotyledons, but they |

are not always so conspicuously sheath-shaped as in most Zingiberaceae.

Morphology in Zingiberaceae 157

A flowering stem bears either 2-ranked complete leaves, or 2-ranked sheaths,

below the flower-bearing part. The flower-bearing part bears bladeless sheaths

spirally arranged (except in Boesenbergia, which needs more study). These are

called the primary bracts of the inflorescence. If the inflorescence is at the top of

a leafy stem, it is protected when young by the sheaths of the leaves on the lower

part of the stem and emerges into free air only at a late stage of development.

Correlated with this is the fact that the primary bracts of such an inflorescence

are in many cases reduced (Alpinia and allied genera) or even absent; one may

reasonably assume that the bracts have become reduced because their protective

function is unnecessary. In cases where an inflorescence is borne on a short stem

bearing short bladeless sheaths only, the primary bracts of the inflorescence are

always well developed, and are exposed at an early stage of development.

The primary bracts, and the basal sheaths on all stems, whether flowering or

not, are homologous, and all are essentially leaves. The primary bracts of the

inflorescence are so called to indicate their position and function, not because

they are essentially in any way different from ordinary sheath-leaves. In most

inflorescences on short stems where the primary bracts are well developed, there

is a leaf-blade rudiment just below the apex of each bract, just as on the sheaths

at the base of a new stem.

At the transition from 2-ranked sheaths to spirally arranged primary bracts,

on non-leafy flowering stems, there is often a gradual transition of shape between

the uppermost 2-ranked sheaths and the lowest primary bracts. But the transition

from the 2-ranked to the spiral arrangement is distinct, as can be seen from fig.

19A, 21A, 23A, 26A in Holttum 1950. The 2-ranked sheaths of the short flowering

stems have no axillary buds, but the spirally arranged primary bracts normally

bear flowers, or short cymose branch-systems, in their axils. There are however

in Malaya and Java genera in which the lower primary bracts also are empty

(Nicolaia, Achasma, Hornstedtia); in these cases the empty lower primary bracts

are much larger than the upper ones (which have flowers in their axils) and

completely enclose the latter. Bakhuizen (1968, p.51) states that he cannot see

any difference between the 2-ranked sheaths and outer empty bracts, and therefore

does not agree with my treatment of Achasma as a genus distinct from A momum.

I will revert to this matter later.

Inflorescence-structure

The spirally arranged primary bracts of an inflorescence bear in their axils

either single flowers or more or less condensed monochasial cymes bearing several

Successive flowers. There is not a sharp distinction between the two conditions.

In the genus Catimbium (Holttum 1950, p. 149) the same inflorescence may have

3-flowered cymes in the axils of basal primary bracts and only single flowers in

the axils of upper primary bracts. As axillary monochasial cymes occur in other

families of Monocotyledons, it is probable that this was the primitive condition

in Zingiberaceae. The genera which have solitary flowers may thus be regarded as

separate developments of reduction from the cymose state. In the genus Zingiber

all species have solitary flowers except Z. clarkei King, which certainly belongs

to the genus, as judged by the peculiar floral structure which occurs in no other

member of the family.

158 Gardens’ Bulletin, Singapore — X XVII (1974)

TexT-FiGc, Diagrams showing branching of the inflorescence. In all cases the main axis of

the inflorescence is shown bearing one bract which has in its axil a single flower or a

monochasial cyme; P = primary bract, S = secondary bract. 7: the condition of Curcuma,

with an axillary cyme. The first secondary bract is at right angles to the primary bract, and

successive secondary bracts each at right angles to the preceding one. 2: the condition of

Amomum, the axillary cyme reduced to one flower. 3: the condition of Hornstedtia and

Achasma, with outer larger empty primary bracts and smaller inner ones each with one

axillary flower. 4: the condition of Scaphochlamys. The first secondary bract is a 2-keeled

prophyll backing the main axis, the rest as no. 1. 5: the condition of Kaempferia, with

deeply bilobed secondary bract backing the main axis, and one flower. 6: the condition of

Zingiber, with unkeeled secondary bract facing main bract, and a single flower.

The text-figure shows diagrammatically the postulated primitive type of

inflorescence in Zingiberaceae, with an extended monochasial cyme in the axil of

a primary bract. This cyme consists of a series of short axes each bearing one

lateral bract and ending in a flower; a bud in the axil of the bract repeats the

pattern. In this diagram for convenience, all branching is shown in the plane of

the paper, but each bract appears to be at right angles to the preceding one, so that

branching is not in one plane; the cyme is called a cincinnus. In practice, the cymes

are in most cases so condensed (the successive axes being very short) that the exact

relation of each bract to the preceding one is often difficult to see exactly. The

situation is most easily observed in Globba and Alpinia (sensu Holttum 1950,

p. 140); a condensed cyme of Curcuma is illustrated in Holttum 1950 fig. 4 B, C.

The detailed development of these cymes needs more precise morphological study

than I can attempt, especially in the case of Scaphochlamys (see below), by someone

with access to ample fresh material.

The nomenclature of the bracts of the cincinnus needs clarification, In 1950,

p. 7, I noted that these bracts could be called secondary bracts, to distinguish them

from primary bracts, but that it was “more usual and more convenient to call

them bracteoles’’, as Schumann (1904) did; however I was not consistent in this

terminology, using both terms in later descriptions of genera and species. In the

present statement all bracts on the cincinnus will be called secondary bracts only

(see figure). They are developed on successive axes of the cincinnus and, though

they are all regarded as having the same status, the later ones (at least when

immature) are completely enfolded by the larger earlier ones when axes of the

eae a

Morphology in Zingiberaceae 159

cincinnus are very short, for which reason I used the term “inner bracteole”’ on at

least two occasions for small later secondary bracts, a term taken up by Bakhuizen

in a way not clearly explained (see below).

Thus the first short axis of the cincinnus, with a flower at its apex, is in the

axil of a primary bract; later ones are all in the axils of secondary bracts. In 1950

(following Schumann) I sometimes used the term “‘floral bract”, which in some

cases referred to a primary bract (e.g. Achasma), sometimes to a secondary bract

(e.g. Geostachys). In Achasma I used it to distinguish the inner primary bracts,

each having a flower in its axil, from the outer empty primary bracts.

The condition of Amomum and other genera in which the cincinnus is

reduced to one flower is shown in fig. 2. The condition of Hornstedtia, Achasma

and Nicolaia is shown in fig. 3. In the latter case, the sterile primary bracts,

attached at a lower level on the main axis than those which have axillary flowers,

are larger and often completely cover the upper ones; the lower ones are thus

in practice “outer” bracts and the upper ones “inner” bracts, but all, being

attached to the main axis of the inflorescence, are primary.

In the genus Scaphochlamys (which does not occur in Java, and is thus not

dealt with by Bakhuizen) there is a peculiar development in the first secondary

bract, at the base of each cincinnus. This bract is 2-keeled, like the organ

commonly called a prophyll in Gramineae, Cyperaceae, Marantaceae and other

families (the German term is adossierte Vorbiatt). In these families a 2-keeled

prophyll occurs at the base of every new branch, whether large or small, and its

back is usually curved to fit the parent axis against which it is compressed. So

far as I know, such a 2-keeled prophyil does not occur elsewhere in Zingiberaceae

except in a simplified form in Kaempferia. The first secondary bract in Curcuma

is certainly lateral, not facing the primary bracts (Holttum 1950, fig. 4B).

Schumann (1904, p. 14) mentions an adossierte Vorblatt in his species Alpinia

orchioides and A. pterocalyx from Celebes, which I have not seen and do not

understand from his descriptions.

In Scaphochlamys the 2-keeled prophyll functions as an additional (basal)

secondary bract on the first axis of the cincinnus (fig. 4) and has either one or

two flowers in its axil; it is comparable to the utricle in Carex, which is a

prophyll in the axil of which a female flower is borne. The rest of the cincinnus

in Scaphochiamys has normal secondary bracts, but they are smaller than the

2-keeled one which at first enfolds them all (Holttum 1950, fig. 10). For this

reason I used the term “outer bracteole” (= outer secondary bract) for the

2-keeled bract, in the generic description of Scaphochlamys, and I believe nowhere

else (not in the descriptions of individual species, where I used the terms first

bracteole for the 2-keeled secondary bract, and 2nd, 3rd bracteoles for the rest).

Again, Bakhuizen has taken up this term and has given it another significance.

In Kaempferia (Holttum 1950, p. 117) there is only one flower in the axil

ot each primary bract. The only secondary bract is more or less deeply bilobed,

sometimes divided to the base (fig. 5); it appears to be comparable to the

2-keeled first secondary bract in Scaphochlamys, which is certainly a related

genus. In Zingiber the sole secondary bract faces its primary bract, in the same

way as the first secondary bract in Scaphochlamys, but it is neither keeled nor

bilobed (fig. 6).

160 Gardens’ Bulletin, Singapore — X XVI1 (1974)

The above statement is necessary to clarify and correct my terminology of

1950, and also to point out how Bakhuizen has used some of the same terms

confusedly in his opening statement on the family (1968, p. 41). He begins by

stating that the main axis of the inflorescence is “provided with spirally arranged

primary bracts bearing in their axil a lateral axis (cincinnus) with bracts of lower

rank”. Here it should be noted that a cincinnus is not a single axis but a branch-

system. Bakhuizen then adds a parenthesis stating that the secondary bracts are

taxonomically important, but he does not define them. One may however infer

that they are not on the primary axis of the inflorescence; yet in his description

of Nicolaia (see further comment below) he uses the term secondary bract for

the inner (upper) bracts of the primary inflorescence axis.

Next Bakhuizen writes: “the whole complex of the usually numerous bracts

which together with the branch-system and the floral bracts constitutes the

inflorescence consists in the most complete condition of 3 elements: 1° primary

bracts (bearing in their axil a lateral shoot, therefore sterile), often firmer or in

other respects more conspicuous than the internal bracts, sometimes called

involucre when they both distinctly surround the other ones and greatly differ

in size and/or shape; as a rule the difference is not remarkable, in most cases

the bracts gradually pass from one kind to another, which makes distinction

almost impossible, especially in herbarium material”.

As indicated in my own statement above on inflorescence-structure, primary

bracts bear in their axil either a cincinnus (sometimes reduced to one flower) or

nothing. Bakhuizen’s statement that they bear in their axil a lateral shoot and

are therefore sterile is to me unintelligible. His statement that inner bracts

(e.g. in Nicolaia) are sometimes smaller than outer ones, with no sharp distinction

is true, but they are all primary bracts.

He continues: “2° secondary bracts and those of lower ranks (outer brac-

teoles in the sense of Holttum; the outer ones sterile, the innermost fertile”. I

used the term “outer bracteole’ once only, in describing the non-Javan genus

Scaphochlamys, as above explained. The terms outer and inner “outer bracteoles”

are not intelligible to me.

Next follows: “3° bracteole (‘inner bracteole’ in the sense of Holttum) in

the axil of the innermost ‘outer bracteole’, entirely or partly surrounding the true

flower, sometimes absent”. So far as I can see, I only used the term inner bracteole

in two cases (generic description of Scaphochlamys, and fig. 20C of Hornsiedtia

leonurus) to describe the smaller secondary bracts which are enfolded by earlier

ones. They are not bracts of a third order, a concept never mentioned by me.

In the whole of the above discussion on the three types of bracts Bakhuizen

fails to mention to which axes the three classes of bracts or bracteoles are

attached, and therefore one cannot be sure what he means. One can only infer

from his later use of the terms in generic descriptions which are discussed below.

Comments on Bakhuizen’s Key to the genera (1968, p. 42).

As shown in fig. 1, there is a cyme, or branch-system, in the axil of each

primary bract of several genera. In one sense, this may be regarded as a branch

of the inflorescence, but the axillary branch-system does not duplicate the

structure of the main axis. In Malaya there are species in the genera Plagiostachys

(Holttum 1950, p. 160), Alptnia (p. 140) and Languas (p. 156) which have true

inflorescence branches in the axils of lower primary bracts (see Holttum 1950

fig. 16); these branches are identical in structure with the apex of the whole

Morphology in Zingiberaceae 16]

inflorescence (Holttum 1950 p. 6, last paragraph). But Bakhuizen, in his key to

the genera, under “axis of the inflorescence branched” includes Globba, Alpinia

and Catimbium where (in species of Java) the “branches” are not like the axis

of the inflorescence but are cymes in the axils of primary bracts.

Apart from the ambiguity of the phrase “axis of the inflorescence branched”,

there are errors in the later part of the key. The characters given under /0 a

and JO b are mixed: they should be corrected as follows:

10 a. Bracts distichous, the upper ones flowering first; apex of bracteole

entire; labellum often saccate, not strongly bilobed; axis of inflorescence distinctly

mutecte to short but not “discoid <2 .:2) 2... 11. Boesenbergia

10 b. Bracts not distichous, lower ones flowering first; bracteole 1 (2-dentate

or 2-partite) or 2 (unlobed, linear); labellum deeply bilobed, not saccate; axis

Meemerarescence, GISCOIG, flat OF CONVEX . 0:06.65 scu-asccninne eo ietenene ste 10. Kaempferia

The characters under JJ a and J// hb are also confused. // a@ should be

corrected to “bracteoles tubular’; // b needs the addition “‘bracteoles with free

margin”.

The genera Hornstedtia, Nicolaia and Achasma

Malayan species of these genera, so far as then known, were all included

in Amomum by Baker (1892, pp 233-243) but mixed in his various subgenera.

These genera all differ from Amomum by the fact that the lower primary bracts

are larger than the later ones, which they enfold; these outer primary bracts

are empty and serve for protection of the later-formed parts of the inflorescence.

The three genera also differ from Amomum in floral structure. The basic pattern

of the inflorescence is shown in fig. 3. As pointed out by Valeton (1904) and

as shown in Holttum 1950 fig. 21, 22, 23, the floral structure in Nicolaia and

Achasma is identical, and distinct from any species of true Amomum, but

Bakhuizen maintains Nicolaia as a distinct genus and unites Achasma to

Amomum. He distinguishes Nicolaia from Amomum in his key to the genera

(1968, p. 42, 43) solely by the fact that in the former the inflorescence is on an

erect peduncle, whereas in Amomum the peduncle is short and wholly or almost

wholly subterranean. But in Nicolaia solaris (B1.) Horan. the inflorescence is

sometimes only just raised above the ground (van Steenis 1972, pl. 57, fig. 5),

and in N. hemisphaerica (B1.) Horan. (Bakhuizen 1968 p. 63) the peduncle is

only 34-12 cm long. Thus Bakhuizen rates this single, not very distinctive

character as of greater importance than the distinctive and uniform characters

of floral structure. The coloured illustrations published by van Steenis (1972,

pl. 57) show the great similarity between N. solaris and Amomum (Achasma)

coccineum (B1.) K. Schum. in general aspect of the inflorescence quite apart from

details of floral structure.

Bakhuizen unites Achasma with Amomum because he cannot see, from her-

barium specimens, that there is a clear distinction between the “outer stalk-

scales [i.e. 2-ranked sheaths of the peduncle], involucral leaves [i.e. empty outer

primary bracts] and bracts, which pass into one another in dried specimens”.

He states that in Holttum 1950, p. 183, I wrote that “involucral leaves and bracts

pass into one another”. What I did write was: “involucral bracts 2-8, much

wider than the inner floral bracts ... Floral bracts with one flower to each, the

162 Gardens’ Bulletin, Singapore — X XVII (1974)

inner ones narrow, the outer often wider and showing a transition to the

involucral bracts”. The ‘floral bracts” of this statement are the inner primary

bracts which bear flowers in their axils and by this fact are distinguished from

the empty outer involucral primary bracts. In the case of Nicolaia, which has

an indentical inflorescence structure, Bakhuizen himself notes (1968, p. 62) that

there is a gradual change from involucral bracts to the inner ones which have

axillary flowers.

Bakhuizen further confuses the situation by using the term secondary bract

for the inner primary bracts of Nicolaia which have axillary flowers, but in his

description of Amomum coccineum (B1.) K. Schum. (which belongs to Achasma

and was so placed by Valeton) he does not. The basic distinctions between

Nicolaia + Achasma and Amomum are that in the former the filament of the

stamen and the base of the labellum are united in a separate tube beyond the

apex of the corolla-tube, and the anther is massive, more or less cleft at the

apex, never crested. In the key which is combined with the specific descriptions

in Amomum, Bakhuizen (1968, p. 55, para. 8a) fails to mention these as distinc-

tive characters; in his generic description of Amomum he includes “labellum .

usually not adnate to the filament, sometimes partly so”, not indicating that the

latter condition applies solely to Achasma, nor does he mention the distinctive

character of the anther of Achasma. In his description of Nicolaia he does

mention these identical distinctive characters. Thus he disguises the identity of

floral structure in Nicolaia and Achasma. To the above distinctions may be

added two others, mentioned by Valeton but not by Bakhuizen: in Nicolaia and

Achasma the base of the labellum is rolled spirally inwards on withering (a

conspicuous character on living inflorescences) and in both genera several flowers

open simultaneously, forming a circle with the labella radiating outwards (van

sreemis, 19/2, Dir 37,08.. 2.10:

In my judgement, Achasma and Nicolaia should be united. The only

differences are: (a) length of peduncle, (b) length of labellum, (c) fewer involucral

bracts in some species of Achasma, (d) outer involucral bracts spreading in

Nicolaia, not in Achasma (where they are prevented by the earth from spreading).

These are relatively trivial characters, and none of them are very sharp. This has

been noted above for the peduncle of Nicolaia solaris. Amomum maingayi Bak.,

which Schumann transferred to Phaeomeria (Holttum 1950, p. 180, fig. 21) is like

Nicolaia in having a fairly long erect aerial peduncle, but its involucral bracts do

not spread horizontally and its labellum is about intermediate in length between

typical Nicolaia and typical Achasma (Valeton 1904, p. 96, suggested that such

intermediates might occur.). Thus in characters a, b, and d there is no sharp

distinction between the two genera, and in the floral characters mentioned there

is quite uniform identity. As regards number of involucral bracts, in Achasma

they are usually sufficiently numerous to give a quite distinctive aspect in living

plants (which are very abundant locally in Malaya, though apparently not in

Java) as shown in Holttum 1950, fig. 23A). But in two Malayan species which

have only 1-3 flowers in an inflorescence the number of involucral bracts is

fewer, sometimes only two. It seems to me possible that plants described as

Achasma pauciflorum and A. subterraneum (Holttum 1950, p. 187) are only

depauperate (or immature) forms of A. macrocheilos and A. sphaerocephalum

respectively, with which species they agree in form of corolla and other details.

Morphology in Zingiberaceae 163

There is another complication to this situation, not mentioned in my work

of 1950 nor by Bakhuizen, namely the existence of species in the eastern part of

Malesia which were included by Valeton in the genus Geanthus (Valeton 1913,

pp 930-936, pl. 162-166: 1914, pp 43, 55-58). These species have a floral

structure as in Nicolaia and Achasma but have few or no involucral bracts. The

original publication of the generic name Geanthus by Reinwardt (see Burtt and

Smith 1972, p. 215) included species of Hornstedtia, Achasma and Nicolaia.

G. coccineus (BI.) Reinw. is recognized as the type of Geanthus; Achasma Griff.,

a later name, is certainly a synonym. But there is a much earlier generic name

which certainly applies to the Malayan species Achasma megalocheilos, namely

Etlingera Giseke (1792) based on a long and detailed description of Amomuin

littorale Koenig (1783; for references see Burtt and Smith 1972). It seems to me

clear therefore that the species which I included in Achasma and Phaeomeria in

1950 should be transferred to the genus Eftlingera. The problem is whether

Geanthus sensu Valeton 1913 and 1914, which does not include an original

species of Geanthus Reinw., should also be included.

Of the three Malayan genera recognized by me in 1950 which have large

involucral bracts, Hornstedtia remains to be considered. Ridley (1924) included

Achasma sensu Holttum 1950 in Hornstedtia because of the obvious similarity

of the form of the inflorescence (which he could see from living plants). But the

structure of flowers in the two genera (which Ridley did not observe carefully)

is different, and they should certainly be separated. On this I have no new

information, but I again call attention to the aberrant species H. leonurus

(Holttum 1950, p. 167), which should perhaps be made the type of a new genus;

excluding H. leonurus, Hornstedtia is very sharply distinct from Achasma.

Bakhuizen’s generic description of Hornstedtia (1968, p. 58) includes the follow-

ing: “each involucral bract ... bearing in its axil (very) numerous secondary

bracts (floral bracts), the outer ones of which (inner involucral bracts) still

resemble the involucral bracts, the inner ones narrower”. The fact is that all

involucral bracts are empty. Again, as in Nicolaia, he refers to the bracts which

subtend flowers as secondary, whereas they are primary bracts, attached to the

primary axis of the inflorescence.

Alpinia and allied Genera.

As regards the genera of the Alpinia alliance in Malaya, Burtt and Smith

point out that according to the present Code the name Alpinia should strictly

apply to the species I have included in Languas, and that there is an earlier

name which should replace Catimbium. But it is still not clear what name should

replace Alpinia sensu Holttum 1950; this was the reason why I retained the name

Alpinia for them (all but one had already names in Alpinia) and used Languas

for L. galanga and its near allies. In correspondence, Mr Burtt has pointed out

to me the great similarity between the flowers of Alpinia (Languas) galanga

(L.) Willd. and Alpinia allughas (Retz.) Roscoe of Ceylon (Burtt and Smith 1972,

fig. 3A, 3B); A. allughas clearly belongs to Alpinia sensu Holttum 1950, on

account of its funnel-shaped secondary bracts. Mr Burtt suggests that Languas

and Alpinia, of my arrangement of 1950, should be united, pending further study

of non-Malayan species; but I still think that my Alpinia represents a species-

group distinct from Languas, and that recognition of a distinct genus Languas,

164 Gardens’ Bulletin, Singapore — X XVII (1974)

though in an illegitimate sense, is the simplest solution for the present situation,

recognizing that there are extra-Malayan species which need further study and

might indicate a different concept or the use of another generic name. Merely

observing the rules of the Code does not ensure that one writes rational taxo-

nomy.

Experimental investigation of inflorescence-structure

D. L. Smith (1967, p. 25) treated the developing inflorescence of a species

of Carex with kinetin, and thereby induced the further development of a partial

inflorescence which is normally reduced to one female flower. It occurs to me

that treatment of young inflorescences of species of Zingiberaceae which have

solitary flowers might induce the development of a branch of the cincinnus in

the axil of the secondary bract which in such cases is normally empty, though

no doubt experimental technique would be difficult in Achasma and other genera

with tightly over-lapping primary bracts. If such an experiment were successful,

it could throw new light on the basic inflorescence-structure in the family. Such

treatment of Kaempferia and Zingiber might demonstrate that their secondary

bracts are homologous with the 2-keeled first secondary bract in Scaphochlamys;

in any event, a careful study of the development of cincinni in the group of

genera called Hedychieae in my paper of 1950 is very desirable.

I have the impression that a detailed comparison of inflorescence-structure

in Monocotyledons as a whole might throw much light on taxonomic problems,

and that experimental work like that of D. L. Smith could be of much help in

understanding relationships (perhaps in Dicotyledons also, but that is beyond

the scope of my detailed knowledge). In most general taxonomic works there is

detailed description of floral and fruit structure but very cursory information

on branching of inflorescences. McClure (1934) was the first person to show

that the bamboos of Asia show two distinct types of inflorescence-branching.

Until Lane’s paper of 1955 no description had ever been published of the

peculiar branching of the inflorescence in Orchidantha, which is of much interest

in relation to that of other families in Zingiberales (see Holttum 1970). Other

examples might be quoted.

As Valeton showed (1904) a clearly defined classification of Zingiberaceae is

impossible without reference to inflorescence-characters; Schumann’s attempt

(1904) to define genera on floral characters alone led to much confusion. I found

that he had species of both Boesenbergia and Scaphochlamys in his genus

Gastrochilus and also in Kaempferia, and some species of Scaphochlamys in

Curcuma. Hutchinson’s latest account of the family (1973) is still based mainly

on Schumann (little changed by Loesener in 1930) and incidentally retains

Ridley’s “genera” Conamumum and Carenophila. Ridley’s descriptions of these

were so inaccurate that Schumann placed Conamumum and Loesener the later-

described Carenophila in the tribe Hedychieae, whereas examination of the type

specimens in Singapore shows that they belong to Alpinieae. Conamomum in

Ridley 1924 consists of two species of Amomum and one of Geostachys; the sole

species of Carenophila also belongs to Geostachys. The full classification of the

family will not be understood until much further study, from fresh material, of

the species in the eastern part of the Malayan region has been undertaken. It

is very gratifying to know that such study is being pursued by Mr Burtt and

Miss Smith at Edinburgh, but there is much still to be done.

——

i a a

Morphology in Zingiberaceae 165

Literature Cited

BAKHUIZEN VAN DEN BRINK, R. C. Jr. 1968. Zingiberaceae, in Backer & Bakhuizen,

Flora of Java, vol. 3: 41-76. Wolters-Noordhoff, Groningen.

BAKER, J. G. 1892. Scitamineae, in J. D. Hooker, Flora of British India, vol. 6:

198 — 264. Reeve, London.

Burtt, B. L. AND R. M. SmiTH. 1972. Key species in the taxonomic history of

Zingiberaceae. Notes R. Bot. Gard. Edinburgh 31: 177 — 227.

Hoitrum, R. E. 1950. The Zingiberaceae of the Malay Peninsula. Gard. Bull.

Singapore 13: 1— 249, fig. 1— 33.

1970. The genus Orchidantha (Lowiaceae). Gardens Bull. Singa-

pore 25: 239 — 246.

HUTCHINSON, J. 1973. The families of Flowering Plants, 3rd edition. Oxford,

Clarendon Press.

LANE, I. E. 1955. Genera and generic relationships in Musaceae. Mitt. Bot.

Staatssamml. Miinchen 13: 114— 141.

LOESENER, T. 1930. Zingiberaceae, in Engler & Prantl, Nat. Pflanzenfam. 2 Aufl.

Bd. 15a.

McC ure, F. A. 1934. The inflorescence in Schizostachyum Nees. Journ. Wash.

Acad. Sci. 24: 541 — 548.

RImDvey, H. N. 1924. Flora of the Malay Peninsula, vol. 4. Reeve, London.

SCHUMANN, K. 1904. Zingiberaceae, in A. Engler, Das Pflanzenreich IV, 46.

W. Engelmann, Leipzig.

SMITH, D. L. 1967. The experimental control of inflorescence development in

Carex. Ann. Bot., N. S. 31: 19 — 30.

STEENIS, C. G. G. J. VAN. 1972. The mountain flora of Java. E. J. Brill, Leiden.

VALETON, T. 1904. Uber neue und unvollstandig bekannte Zingiberaceae aus West-

Java und Buitenzorg. Bull. Inst. Bot. Buitenzorg 20: 1— 99.

—_————. 1913. Zingiberaceae. Nova Guinea 8: 923 — 988, t. 162 — 179.

1914. Die Zingiberaceen Deutsch-Neu-Guineas. Bot. Jahrb.

52: 40 — 100.

The Tree-ferns of the genus Cyathea in Borneo

R. E. HOLTTUM

Royal Botanic Gardens, Kew

The most complete taxonomic account of the tree-ferns of Borneo is con-

tained in Flora Malesiana, Series II (Pteridophyta) vol. 1, part 2 (1963), which

covers the whole of Malesia and includes descriptions of 191 species. For identifi-

cation of the 29 species of Borneo therefore it is necessary to use elaborate keys

which were designed to cover a much wider range of species. The present paper

is based on the work of 1963, since which little new information has been

obtained, but the keys are revised and simplified to facilitate identification of

Bornean species. Some additional information about local distribution is also

given.

The present paper is written in the hope that it may help people who have

the opportunity of making new observations on these plants. The descriptions are a

summary of the characters of specimens seen by me, and anyone studying them

in comparison with living plants will find gaps in the information here provided.

I hope that some local naturalists will be able to take up this study, and [ will

do my best to help any such persons who will communicate with me. In particular,

it is desirable that the keys to species should be revised, to include more

characters which are easily observable in the field; many characters in the

present keys can only be seen by careful use of a hand lens. Such a new key

would help other locai naturalists, and perhaps help to ensure that all species

of this very interesting group of ferns are allowed to persist, in spite of increasing

destruction of natural forest.

Almost all the species occur within the Mount Kinabalu National Park in

Sabah or in neighbouring lowland forest, so that the present account covers

the special needs of field botanists in the national park. Six* species are only

known from Mt Kinabalu: C. longipes, C. acanthophora, C. havilandii, C. stipiti-

pinnula, C. discophora and C. megalosora.

Cyathea plants may be recognized in nearly all cases by the presence of a

trunk bearing moderately to very large bipinnate fronds which bear scales at the

bases of their stipes (petioles). There is one tree-fern of the genus Dicksonia

(D. mollis Holtt.) in forest on Mt Kinabalu at 5000-6500 ft: it has dense reddish

hairs on its stipes, not scales, and sori in marginal pouches. Four species of

Cyathea in Borneo have simply pinnate fronds and at most very short trunks;

their indusia, if present, identify them generically, also their scales on stipe-

bases which have minute oblique dark marginal setae.

Diagnostic characters for recognition of species

The most important characters for the main subdivision of the genus are

found in the large scales at the bases of stipes. These are best seen on young

fronds: old ones may lose most of them. When collecting herbarium specimens,

* The seventh, C. brachyphylla Holttum sp. nov. is described in the Addendum.

167

168 Gardens’ Bulletin, Singapore — XXVII (1974)

one can cut off a thin strip 5-10 cm long from the scale-bearing surface of the

stipe; this is much more easily dried than a whole stipe. An additional character

shown by the stipe is the distribution and shape of the areas,-on each side of the

stipe, where aerating tissue comes to the surface (called pneumathodes, or linear

aerophores). These areas are almost white, and their surface is broken when

old: they usually form a discontinuous line on each side of the stipe, or sometimes

a double line. When whole stipes are dried, they shrivel along these lines, so that

the shape and distribution of pneumathodes cannot be seen in most herbarium

specimens, and in most cases information about them has not been recorded. A

thin strip cut along the side of the stipe, to include several pneumathodes, is

easily dried and provides permanent information in the herbarium.

The length of the stipe and size of the lowest pinnae may be important. In

some species (e.g. C. loheri) the stipe is very short and the lower pinnae are

gradually smaller, the lowest quite short. In two Bornean species there are very

small pinnae at the base of the stipe, and then a long gap to the large pinnae:

these basal pinnae should be looked for.

The other characters are best shown by the largest middle pinnae of a frond.

Except in the simply pinnate fronds of a few species, these middle pinnae carry

pinnules of almost uniform size for the greater part of their length; it is these

pinnules which are described in the detailed descriptions of species, and their

size, shape, depth of lobing and venation are important. On the upper surface of

pinna-rachises and costae of pinnules there are always short curved hairs, but

the occurrence and character of hairs and scales on the Jower surface are much

more varied and must be observed carefully; they may be seen in sufficient detail

with a hand lens of 10 X magnification. These scales and hairs are in all cases

mentioned in the present paper.

Sori are also important diagnostically, and have indusia of various shapes,

Or sometimes no indusia. Some indusia are very small and can only be seen by

carefully removing the sporangia. The hairs which occur among the sporangia

(paraphyses) are sometimes distinctive. In one Bornean species (C. tripinnata)

there are false indusia formed by overlapping separate scales; these are not easy

to see with a hand lens, but fortunately the species is easy to recognize from

the fully tripinnate fronds which occur in no other species.

Classification

The classification here adopted is that of Flora Malesiana, with a change of

subgeneric name which must be explained. In Flora Malesiana I divided the genus

Cyathea into two subgenera, subg. Cyathea and subg. Sphaeropteris. The type

species of Cyathea, C. arborea (L.) Sm., is native in the West Indies; I regarded

the Malesian species placed in subg. Cyathea as closely related to it. But Dr

R. M. Tryon subsequently made a new study of tropical American tree-ferns

(Contr. Gray Herb. no. CC, 1970) and showed that C. arborea and some related

species have stipe-scales of a distinct type not found in any species of the Old

World. Dr Tryon also considered that subg. Sphaeropteris is sufficiently distinct

to rank as a separate genus (there are a few species in tropical America). Thus

he needed a new generic name for the Malesian species of Cyathea subg. Cyathea

of my treatment, and the oldest available name is Alsophila R. Br. (type species

A. australis R. Br. of Australia). He transferred all Malesian species of Cyathea

subg. Cyathea to the genus Alsophila, and all in subg. Sphaeropteris to the genus

Sphaeropteris. But in my view the differences are hardly of generic rank: this

Cyathea in Borneo’= | 169

is supported by the fact that species of both subgenera have the peculiar chromo-

some number 69. Therefore I retain the genus Cyathea for Malesian species, sub-

stituting subg. Alsophila for subg. Cyathea; the name subg. Cyathea should then

be restricted to the tropical American species related to C. arborea.

A further note on the generic name Alsophila is desirable because it was used

in the past in a quite different sense. The type species A. australis has no

indusia, and in the 19th century this was regarded as the prime distinguishing

character of the genus, which thus was made to include all then known species of

Cyathea (in the sense of the present paper) which had no indusia. But Copeland

pointed out (Philip. Journ. Sci. 3C: 353; 1909) that such a definition brings

together species which are not allied; in the classification of the present paper it

will be seen that exindusiate species occur in both subg. Alsophila and in subg.

Sphaeropteris, and in each case the exindusiate species certainly conform in all

other characters to their own subgenus. The genus Alsophila, as defined by Tryon,

includes species with and without indusia; thus it has a significance quite different

from that given to it by 19th century pteridologists (e.g. Beddome in his Handbook

to the Ferns of British India, 1883).

Prior to Flora Malesiana, the most important publication on the ferns of Mt

Kinabalu was in Gardens Bulletin S. S. vol. 7 (1934) 191 — 324, in which Carl

Christensen gave an annotated list, based on all known collections including those

made by J. C. and M. S. Clemens in 1931-32 and my own of 1931, which together

doubled the number of species known from the mountain, and included reference

to a few from other parts of Borneo. The paper includes an account of my

itinerary, with notes on ferns seen (pp. 185 + 206). The number of species of

Cyathea in that paper was 18. In some cases the nomenclature of the present

paper differs from that of Christensen of 1934; references are given to all such

cases.

When preparing my account of Cyathea for Flora Malesiana I examined type

specimens of all previously published species, few of which had been described

in sufficient detail for clear recognition, with the result that all comparative

taxonomic statements were in some measure confused. I found for example that

no two accounts of the species of Cyathea in the Nature Reserve adjacent to the

mountain garden at Tjibodas in Java agreed as to number of species and distinc-

tion between them, though this forest had been studied more carefully than any

other area in Malesia. This was due to the fact that scales on pinnules, and

indusia, had not been examined in sufficient detail. When re-describing type

specimens I discovered that many Malesian species had received more than one

name, so that many earlier names appear as synonyms in Flora Malesiana. |

have not cited all these synonyms, except those used by Christensen in 1934,

nor have I cited previous descriptions of the species, which are mostly inadequate

where they are not also inaccurate; they are not reliable as indicators of distribu-

tion of species. I have also not included Dr Tryon’s new names, which may all

be found in his publication cited above. References to the account of Cyathea in

Flora Malesiana, Series II, vol. 1 (1963), are given as F.M. with page number.

KEY TO THE SUBGENERA AND SECTIONS

Stipe-scales usually dark and rigid, always with fragile edges which are eroded

when old and may or may not bear irregular dark setae: hairs on lower

surfaces, if present, crisped and appressed; indusia in some cases attached only

on side of sorus remote from edge of pinnule-segment ......................c00ceceeeeeee.

170 Gardens’ Bulletin, Singapore — X XVII (1974)

Indusia present (sometimes very small, hidden by sorus); rachises usually

green when fresh, never dark chocolate-brown; fertile and sterile pinnules

of almost* uniform ‘size ee. PR eee ea eee sect. Alsophila

Indusia absent; rachises dark chocolate-brown; fertile pinnules usually

narrower, or with narrower lobes, than sterile ............ Sect. Gymnosphaera

Stipe-scales usually thinner and paler, of uniform texture, with very short

(usually darker) setae set obliquely on their edges: hairs on lower surface, if

present, stiff and spreading erect from surface; indusia completely covering

sorus and breaking at maturity, or lacking (in a few cases imperfect as an irregular

ring round base; of sorus); never attached ion:one) sider. 211. :3.20. anon ed eee

Pinnules commonly 10-15 cm long, lobed almost to costa throughout or

fully pinnate; costules not over 4 mm apart on pinnules 10 cm long; basal basi-

scopic vein of each group arising from costule above its base.....................

rasdapecdel fur eadess toiett t nerhen Tort Cmte eee ec a. a eee eat tne ee Sect. Sphaeropteris

Free tertiary leaflets few, sori indusiate or not, never covered by over-

lappines SCalesyo.5 «cates -asectens = pbs ater cobea th attnmce subsect. Sphaeropteris

Free tertiary leaflets present on all pinnules; sori covered by overlapping

scales which simulate an indusium ........................ subsect. Fourniera

Pinnules rarely to 10 cm long, lobed less deeply, or pinnae undivided;

costules usually 4 mm or more apart; basal basiscopic vein of each group

arising from costa near base of costule ...................0s.00005 Sect. Schizocaena

SuBG. Alsophila sect. Alsophila

1. Stipe conspicuously spiny in basal part, spines 4-5 mm long:

2. \Pinnulés sail’ stalkedexceptvdistalcones!. 72 nyae I 1. C. longipes

2. Pinnules not stalked, + jointed to rachis ......... 2. C. acanthophora

1. Stipe + warty at base, lacking slender spines:

3. Lower surface of costae densely and persistently scaly; indusium

conspicuous, covering sorus to maturity:

4. Pinnules commonly to 24 cm long; pinna-rachis densely scaly

vbesug eine ne “ula air acini ta toe ene oR ae 3. C. havilandii

4. Pinnules commonly to 74 cm or more long; pinna-rachis glabres-

cent:

5.* Stipe 50 cm or more long; basal pinnae not greatly

reducedy7iis. Vig? UE Veaine~aea! Sterne ee eee: 4. C. oosora

5.* Stipe much shorter: lower pinnae gradually reduced, lowest

CA 'cm Jone) are eae es ie ee) ee ee ee 5. C. loheri

3. Lower surface of costae not densely -scaly; indusium small, at

maturity reflexed against costule or covered by sorus:

6. Pinnules commonly more than 2 cm wide, with wide sinuses between

1Ob@6 47)... 2A2.. ATER ASE On, MOR TAN 6. C. incisoserrata

6. Pinnules less than 2 cm wide, sinuses between lobes narrow:

*See modification in Addendum, p. 181.

Cyathea in Borneo 171

7. Indusium reflexed against costule and usually visible at maturity of

menuns pirapnyses BHOTE y. LON CD, 80 1.2 sel)... 7. C. borneensis

7. Indusium very small, covered by mature sorus; paraphyses longer

than sporangia, 2-4 cells wide at base .................. 8. C. latebrosa.

1. Cyathea longipes Copel., Philip. Journ. Sci. 12C(1917) 54. Gard. Bull. S. S.

7(1934) 205, 222. F.M. 98.

Stipe slender, to 200 cm long, dark and copiously spiny near base, spines

to 5 mm long; basal scales early caducous, rather broad (they need further

examination). Pinnae to at least 70 cm long; pinnules all stalked except distal

ones, stalks of lowest on lower pinnae 10 mm long, on smaller pinnae 3-6 mm;

largest pinnules 10-13 xX 2.0-3.2 cm, acuminate, 1-2 pairs basal segments

free or connected by a narrow wing along costa; rest of pinnule lobed almost

to costa, lobes crenate, costules 44-6 mm apart; veins to 10 pairs. Sori near

costules; indusium rather thin, covering young sori, breaking irregularly at matu-

rity. Main rachis and pinna-rachises green, lacking scales beneath, with scattered

short spines: scales on lower surface of costae ovate-acuminate, thin, entire,

brown, distal ones shorter and + bullate; bullate acuminate scales on costules.

Only known from Mt Kinabalu; formerly abundant in ridge forest on

Penibukan and Marei-Parei ridge, at 4000-5500 ft.

2. Cyathea acanthophora Holtt., Kew Bull. 16(1962) 51. F.M. 93.

Sinilar to C. Jongipes in spines on stipe, and in scales and sori, differing as

follows: stipe to 80 cm long; basal scales to 20 X 1 mm (only seen on a young

frond); pinnules not stalked, more or less jointed to pinna-rachis, smaller

(83-10 cm X 1.5-1.8 cm), without free basal segments.

Only known from Mt Kinabalu, on the Kamborangah ridge, at 6000-7000 ft.

3. Cyathea havilandii Baker, Trans. Linn. Soc. II Bot. 4(1894) 249. Gard. Bull.

S. S. 7(1934) 202, 221. F.M. 96.

Trunk short; fronds mostly almost erect, to 100 cm long, densely scaly

throughout. Stipe 30-40 cm; scales near base to 15 X 1-2 mm, shining medium

brown with very narrow fragile edges, scales on upper part of stipe smaller.

Largest pinnae 10-16 cm long; pinnules to 2.5 cm X 7 mm, only a few free near

bases of largest pinnae, rest connected by a narrow wing along pinna-rachis;

largest pinnules, where fertile, lobed 4 way to costa, where sterile less deeply.

Sori in a single row on each side of costae of pinnules; indusium firm and dark,

breaking irregularly at maturity. Scales on costae bullate with flexuous hair-

tips | mm or more long.

Only known from Mt Kinabalu, in Leptospermum — Dacrydium forest on

ridges at 8000-10,000 feet.

4. Cyathea oosora Holtt., Kew Bull. 16(1962) 59. F.M. 101.

Stipe 50 cm or more long, warty near base; pneumathodes 14-20 mm long

in an almost continuous line, scales not seen. Largest pinnae 60 cm long; pinnules

to 9 x 2 cm, sessile or nearly so, lobed almost to costa, lobes rigid, crenate:

costules 34-4 mm apart, veins 9-10 pairs. Sori near costules; indusium firm,

brown, at first ovoid with a small apical aperture, later breaking irregularly.

Pinna-rachis bearing a few narrow brown scales 3-4 mm long; costae densely scaly,

scales uniformly brown, lower ones elongate, grading to hair-pointed bullate

scales distally and on costules.

92 Gardens’ Bulletin, Singapore — X XVII (1974)

Distribution: Mt Kinabalu at 7000-10,000 ft; also from Mt Rante Mario

in S. W. Celebes at 10,000 ft. The Celebes specimens differ in having paler scales

on costae.

5. Cyathea loheri Christ, Bull. Herb. Boiss. II, 6(1906) 1007. F.M. 104. —

C. korthalsii sensu C, Chr., Gard. Bull. S. S. 7(1934) 222, non Mett.

Trunk to 10 m tall. Stipe short; scales to 25 xX 14-3 mm, pale, firm, their

fragile edges bearing scattered long setae; pneumathodes to 11 mm long. Lower

pinnae gradually smaller, lowest 7 cm long, longest 40 cm or more; largest pin-

nules 743-9} X 14-2 cm, sessile, short-acuminate, lowest 1-4 segments +

contracted at base, rest of pinnule lobed almost to costa, segments deeply crenate

where fertile; costules 33-4 mm apart, veins 10-12 pairs. Sori near costules;

indusium firm, shining brown, almost covering sorus to maturity but open on side

remote from costule, breaking when old. Pinna-rachis at first densely scaly,

larger scales pale with some dark setae; scales at base of costae light brown

with marginal hairs or setae, grading to bullate scales distally and on costules.

Distribution: Mt Kinabalu, near streams in forest in deep valleys at 7000-

9000 ft; Philippines (Luzon, Negros, Mindanao); Taiwan.

This species apparently occurs at lower altitudes in the Philippines than on

Mt Kinabalu. Young plants found by me had long stipes, the lower pinnae not

greatly reduced. The stipe-scales are distinctive.

6. Cyathea incisoserrata Copel., Philip. Journ. Sci. 6C(1911) 361. F.M. 113, fig.

18. — Alsophila latebrosa var. ornata Ridl., Journ. Mal. Br. R. Asiat. Soc. 4

(1926) 8. .

Trunk to about 4 m tall; stipe to 85 cm, warty or with short conical spines

at the base; scales sparse, to 10 X 1 mm, dark with narrow quickly abraded

fragile edges; pneumathodes in a double row on each side of stipe, almost con-

tinuous. Lower pinnae slightly reduced, longest 70 cm long; pinnules commonly

10 X 24 cm, to 12 X 34 cm, sessile, deeply lobed, lobes crenate and separated

by wide sinuses; costules 44-54 mm apart, veins 12-15 pairs. Sori near costules;

indusium very small, on one side of base of sorus, covered by sporangia;

paraphyses longer than sporangia, flat and 2-3 cells wide at base. Lower

surface of pinna-rachis pale green, smooth; scales near bases of costae elongate.

flat, entire, grading to bullate scales distally and on costules.

Distribution: lowland forest in Malaya and Sarawak. This is very near

C. latebrosa (no. 8) but seems to be constantly distinct.

7. Cyathea borneensis Copel., Philip. Journ. Sci. 6C(1911) 135. F.M. 110, —

C. obtusata Rosenst., Med. Rijksherb. no. 31(1917)1; Holttum, Rev. Fl. Mal.

2 (19355) 121;

Trunk to 2 m or more tall; stipes c. 25 cm, dark and warty at base, scales to

15 xX 1 mm, dark, glossy, with narrow fragile edges; pneumathodes 12-18 mm

long. Lower pinnae rather irregularly reduced, lowest c. 10 cm long, longest 60

cm; largest pinnules 8-10 X 1.7-2.2 cm, deeply lobed, lobes almost entire;

costules 34-5 mm apart, veins to 10 pairs. Sori near costules; indusium rather

thin, on costular side, pressed against costule at maturity of sorus; paraphyses

short. Pinna-rachis green, + suffused with purple, bearing crisped hairs distally on

lower surface; scales on costae dark, entire, flat or with bullate base, grading

to dark bullate scales distally and on costules.

ee :

Cyathea in Borneo 173

Distribution: Sarawak and Sabah, in lowland forest and to 3500 ft, but noi

recorded for Mt Kinabalu; Malaya and Peninsular Thailand.

8. Cyathea latebrosa (Hook.) Copel., Philip. Journ. Sci. 4C(1909) 52. Gard. Bull.

S. S. 7(1934) 195, 198, 222. F.M. 115. — Alsophila latebrosa Hook., Spec. Fil.

1 (1844) 37.

Similar to C. incisoserrata (no. 6) but smaller, with smaller pinnules having

costules 3-4 mm apart and narrow sinuses between the lobes: pneumathodes on

stipe in a single row on each side.

Distribution: throughout Borneo; Sumatra, Malaya, Thailand to Hainan, in

forest or on edges of forest, in low country and to 4500 ft.

SuBG. Alsophila sEcT. Gymnosphaera

1. Base of stipe bearing reduced pinnae, widely separated from normal

pinnae:

2. Small basal pinnae with many narrow persistent segments, green when

poune.usoon. tuimme browns... 2.4.0...) 9. C. ramispina

2. Small basal pinnae stiffly spreading, with few slightly-lobed leaflets,

peach oh 2s ga] | jae es ee a ee 10. C. recommutata

meeemense Of Stipe lacking reduced ‘pinnae .-.....-..0..0)......... 11. C. glabra.

9. Cyathea ramispina (Hook.) Copel., Philip. Journ. Sci. 4C(1909) 36. Gard.

Bull. S. S. 7(1934) 200, 222. F.M. 117. — Alsophila ramispina Hook., Syn. Fil.

(1866) 42.

Trunk rather slender, to 2 m tall, persistently covered with the finely divided

basal pinnae which remain attached to the old leaf-bases (the youngest ones

green and covering the apex of the trunk). Stipe dark, almost covered with small

scales, basal scales very dark, glossy, to 10 xX 14 mm with pale fragile edges.

Normal pinnae to 45 cm long; pinnules slightly dimorphous (fertile smaller),

lowest with stalks 2-3 mm long, largest 7-9 x 14-2 cm, lobed to about 2 mm

from costa, lobes slightly crenate; costules 444 mm apart, veins to 8 pairs,

usually all simple. Sori without indusia, distal ones close to costule, lower ones

more distant from it. Scales on costae and costules narrow, dark, glossy with pale

edges, grading to pale bullate scales.

Distribution: Sarawak and Sabah, in ridge forest at 6000-7000 ft on Mt

Kinabalu; at 3000-4500 ft in Sarawak: also at 500 ft on a sandstone hillside in

Tawao, Sabah.

10. Cyathea recommutata Copel., Philip. Journ. Sci. 4C(1909) 36. Gard. Bull.

S. S. 7(1934) 198, 220. Holttum, Rev. Fl. Mal. 2 (1955) 125. F.M. 118, fig.

19a, b, 20d.

Habit of C. ramispina but basal small pinnae not so closely placed, rigidly

Spreading on each side of base of stipe, bearing few less deeply lobed leaflets

which often become detached so that the reduced pinnae are like stout spines

When old. Largest pinnae 40 cm long; pinnules rather strongly dimorphous,

Sterile to 1.6 cm wide with costules 444 mm apart, fertile 6-12 mm wide with

Closer costules.

174 Gardens’ Bulletin, Singapore — X XVII (1974)

Distribution: Borneo, Malaya, central and southern Sumatra; in forest,

usually at lower altitudes than C. ramispina, on Mt Kinabalu at 4500 ft; also

occurring in several parts of Borneo in lowland swamp forest on sandy ground.

11. Cyathea glabra (B1.) Copel., Philip. Journ. Sci. 4C(1909) 35. Holttum, Rev.

Fl. Mal. 2(1955) 127. F.M. 120. — Gymnosphaera glabra Bl., Enum, Pl. Jav.

(1828) 242. — Alsophila vexans Cesati, Atti Acad. Napol. 7, no. 8(1876) 4. —

Cyathea vexans (Ces.) C. Chr., Gard. Bull. S. S. 7(1934) 218.

Trunk rather slender; stipes very dark; basal scales dark, glossy with pale

fragile edges. Lower pinnae sometimes much reduced but not remote from rest;

largest pinnae 45—55 cm long; largest pinnules 9-12 X 14-2 cm, lowest on stalks

2-4 mm long, edges crenate to slightly lobed, not or little dimorphous; costules

4-5 mm apart; veins 3-5 pairs, simple. Sori without indusium. Scales on costae

few, narrow, dark with pale edges which often bear a few dark setae; no bullate

scales.

Distribution: West Java, Sumatra, Malaya, Borneo; in Jowland swamp

forest and in mountain forest to 5000 ft. There has been much confusion in the

use of the specific name glabra, and there are other synonyms.

SUBG. Sphaeropteris SECT. Sphaeropteris

|. Free tertiary leaflets few:

2. Stipe minutely warty at base; lower surfaces of costae copiously hairy

throushout; INGUSIa PRESENCE “lone hore tee 12. C. leucotricha

2. Stipe strongly spiny at base; lower surface of costae bearing a few

Hairs distally: NG Mansa no ae ce eee we teeeee 13. C. contaminans

1. Free tertiary leaflets present on all pinnules ............... 14. C. tripinnata

12. Cyathea leucotricha Christ, Ann. Jard. Bot. Btzg 20(1905) 135. F.M. 127.

Height of trunk not recorded. Stipe SO cm, minutely warty; basal scales early

caducous, dark brown with concolorous marginal setae. Largest pinnae 60 cm

long; largest pinnules 9-12 x 145-2 cm, 1-3 basal segments free or nearly so, rest

of pinnule lobed almost to costa, lobes crenate; costules 4-5 mm apart, veins

10-12 pairs. Sori nearer to costule than to edge, indusiate; indusium pale, thin,

breaking irregularly at maturity. Pinna-rachis glabrescent; costae and costules

bearing many stiff spreading pale hairs on lower surface; a few narrow pale

scales with dark setae near bases of costae.

Distribution: in lowland forest, recorded for several widely-spaced localities

in Sarawak, Brunei and Kalimantan. This species is interesting because very few

others in this section have indusia. More information about it would be welcome.

13. Cyathea contaminans (Hook.) Copel., Philip. Journ. Sci. 4C(1909) 60. Gard.

Bull. 7(1934) 196, 222. Holttum, Rev. Fl. Mal. 2(1955) 119. F.M. 135. —

Alsophila contaminans Hook., Spec. Fil. 1(1844) 52.

Trunk often very tall, much thickened by adventitious roots at base, when

old showing leaf-scars on upper part. Stipe to 100 cm long, pale glaucous, purplish

towards base which is strongly spiny and covered with pale brown scales varying

in size up to 45 X 3 mm, very thin with short dark marginal setae; main rachis

bearing many short spines throughout. Lowest pinnae somewhat reduced with

stalks to 10 cm long; largest pinnae 60 cm; pinnules to 15 X 3 cm, often smaller,

Cyathea in Borneo 175

lowest distinctly stalked, largest with 1-2 pairs basal segments + free, rest lobed

almost to costa, lobes crenate, glaucous beneath; costules 444 mm apart, veins

commonly 12 pairs. Sori near costules, no indusia. Lower surface of costae

bearing at first scattered narrow pale setiferous scales which soon fall; costular

scales small, ovate, pale-fringed, soon caducous; a few hairs pairs present on lower

surface of costae and costules near pinna-apex, variable in number; on Mt Kina-

balu erect hairs may also be presented on lower surface of veins.

Distribution: throughout Malesia, in clearings and open places in forest,

especially near streams, at 1000-5000 ft, often abundant. This is now by far the

most conspicuous tree-fern at lower levels in Mt Kinabalu National Park.

14. Cyathea tripinnata Copel., Philip. Journ. Sci. IC, Suppl. 4 (1906) 251. Gard.

Bull. S. S. 7(1934) 198, 222. Hoittum, Rev. Fl. Mal. 2(1955) 120. F.M. 140.

Trunk 4-5 m tall; stipes to at least 40 cm, dark, bearing scattered sharp

spines 1-3 mm long, covered almost throughout by a felt of very small setiferous

scales, basal scales to 25 X 1 mm, thin and soft, matted together. Lowest pinnae

20-30 cm long, largest 60 cm; pinnules 9-14 x 14-24 cm, fully pinnate; tertiary

leaflets to 15 X 34 mm, largest deeply lobed at base, lower ones distinctly stalked;

veins to 9 pairs, those in free basal lobes pinnate. Sori near midribs of tertiary

leaflets, covered at maturity by overlapping thin pale scales. Lower surfaces of

pinna-rachis, costae and costules bearing minute pale fringed scales.

Distribution: Pulau Tioman, Sabah, Philippines (Luzon to Mindanao),

Amboina. The specimens from West Java which I formerly identified with this

species appear to be distinct, matching recent collections from southern Sumatra;

these will be described elsewhere as a new species of subsect. Fourniera.

SuBG. Sphaeropteris SECT. Schizocaena

1. Fronds sinply pinnate, pinnae entire or crenate-serrate:

2. Apex of frond a deltoid deeply lobed lamina; pinnae sessile; basal

Woime aiisuoinesings) eri .. PEPIN IER isnt A sees. 15. C. capitata

2. Apex of frond pinna-like; pinnae usually stalked; veins free:

3. Pinnae not over 1.5 cm wide, stalks to 12 mm long; sori in 1-2

FOWS,-On-,eack Side, Of costa, fully -indusiate. 22... cai. . .cct encase ss

3. Pinnae 2-4 cm wide, stalked or not; sori on fully fertile pinnae

in more than 2 rows, indusiate or not:

4. Pinnae not long-acuminate, upper usually sessile ...............

17. C. moluccana

4. Pinnae long-acuminate, all stalked ......... 18. C. arthropoda

1. Fronds simply pinnate with deeply lobed pinnae, or bipinnate:

5. Fronds simply pinnate with deeply lobed pinnae, or largest pinnae

sometimes with free pinnules at their base:

6. Pinnae commonly 25 cm long; no long pale hairs on lower

SR EN RNC a sa ticle ct Rey Ss Ede 19. C. alternans

6. Pinnae sometimes shorter; main rachis bearing long pale hairs on

lower; Supfaced: 0). 21. Pha 20. C. trichophora

176 Gardens’ Bulletin, Singapore — X XVII (1974)

5. Fronds amply bipinnate:

7. Sori indusiate (indusium sometimes reduced to a disc hidden by

sorus):

8. Basal pinnules of middle pinnae with stalks to at least 4 mm

long:

9. Pinnules lobed less than 4 way to costa; no free basal

SEPMIEMUS VRGITs.. Reed besa tan dies taooene 8 21. C. stipitipinnula

9. Pinnules lobed to 1-2 mm from costa; larger pinnules with

1-2, Tree “Daal SEPMen Si cise .avdk. eens 22. C. assimilis

8. Basal pinnules sessile or nearly so;

10. Indusium a disc hidden by sorus; basal pinna-lobes not

TES AAW Ds rch Sale acho ip eto atette dots 23. C. discophora

10. Indusium complete, covering sorus to maturity ............

is). .S AGL Jintao). Jee. BIeY OO, 24. C. megalosora

7. Sori without indusia:

11. Long spreading hairs abundant on lower surface of rachis

and/or pinna-rachis, often also on costae:

12. Costules and veins bearing hairs like those of costa on

lower surface:

13. Pinnules lobed 4-2 towards costa; costules 4 mm

Of MOP apatt var sco ee aces 25. C. trichodesma

13. Pinnules lobed to within 1 mm from costa; costules

3-35 THE, BDAUL .. eehniteatickesdeuetinen 26. C. wallacei

12. Costules and veins lacking hairs on lower surface

Se ees eau ne aan ce eee it teereesere GEIR 20. C. trichophora

11. Long spreading hairs lacking on lower surface of rachises:

14. Largest pinnules with a free segment at base; pinnules

on stalks to 4 mm or more long ......... 27. C. polypoda

14. Largest pinnae lacking free basal segment, almost

sessile :

15. Bullate scales lacking on costae and costules; pin-

nules lobed to less than 1 mm from costa .........

Mths. ERE eden. aah eee teeta sad 28. C. agatheti

15. Bullate scales present on costae and costules;

pinnules less deeply lobed ...... 29. C. squamulata

15. Cyathea capitata Copel., Philip. Journ. Sci. 12C(1917) 49. Gard. Bull.

S. S. 7(1934) 201, 218. F.M. 142.

Trunk 1-3 m tall; stipe dark, to at least 40 cm long; basal scales brown,

firm, to 25 X 3-4 mm, edges bearing concolorous setae. Apex of frond broadly

deltoid and deeply lobed, grading into upper pinnae; pinnae to 40 pairs, jointed

to rachis, largest 15-19 x 2-3 cm, edges entire except near apex, base truncate

to cordate; veins in pinnate groups of c. 3 pairs, outer members of each group

anastomosing with those of adjacent groups. Sori usually in 2 rows on each side

of the costa of a pinna; indusium thin, completely covering young sorus, breaking

when old.

Distribution: Sarawak (Mt Murud), Sabah (Mt Kinabalu), in wet ground

near streams in forest at 4500-6000 ft.

Cyathea in Borneo 177

16. Cyathea angustipinna Holtt., Kew Bull. 16(1962) 52. F.M. 143.

Trunk to at least 50 cm tall; stipe c. 30 cm, basal scales pale, firm, to

20 x 14 mm, marginal setae dark. Pinnae c. 18 pairs, to 12 cm long, sterile to

1.6 cm wide, fertile 1.0-1.2 cm, base narrowly cuneate, stalks to 12 mm long,

edges entire except near apex; veins in groups of 3, basal one attached separately

to costa, free. Sori in 2 rather uneven rows on each side of costa, indusiate as in

C. capitata.

Distribution: known only from 2 collections on Mt Dulit, Sarawak, on

sandy bank of stream near waterfall, in forest, at 4000 ft.

17. Cyathea moluccanma R. Br. in Desv., Mem. Soc. Linn. Paris 6(1827)} 322.

F.M. 143. — C. kinabaluensis Copel., Philip. Journ. Sci. 12C(1917) 50. C. Chr.,

Gard. Bull. S. S. 7(1934) 218. — C. brunonis Hook., Gen. Fil. (1838) t.2. Holttum,

Rev. Fl. Mal. 2(1955) 117.

Trunk to 50 cm tall; stipe 20-30 cm, basal scales medium brown, 15-30 x

4—3 mm, edges with concolorous setae. Leafy part of frond to 150 cm or more

long; apical lamina a pinna like the rest; pinnae jointed to rachis, stalked or the

upper ones sessile, 12-28 xX 2-4 cm, base broadly cuneate, edges entire except

for short-acuminate crenate apex; veins as in C. angustipinna, but the middle

vein of each group forked once or twice. Sori in 1-3 rows on each side of costa,

commonly 4-6 on each vein-group, indusium as C. capitata or in some cases

only forming a disc which is hidden by sorus.

Distribution: Central Sumatra, Malaya, Lingga, Borneo (excluding south

and south-west), South & Central Celebes, Moluccas (Ceram, Amboina), in forest,

low country to 3000 ft.

Copeland in 1917 described three exindusiate species from Mt Kinabalu,

C. kinabaluensis, C. pseudobrunonis and C. fuscopaleata, distinguishing them

by size and colour of stipe-scales, but I cannot see clear distinctions between them,

and none are totally without indusia. See note under C. alternans.

18. Cyathea arthropoda Copel., Philip. Journ. Sci. 6C(1911) 134, t.13. F.M.

143.

Habit of C. moluccana, differing: all pinnae stalked, stalks of upper ones

5 mm, of lower ones 12-15 mm: shape of pinnae narrowly elliptical (sides not

parallel as in C. moluccana) with a caudate apex to 4 cm long; indusium a

narrow irregular ring hidden by sorus.

Distribution: Sarawak, Bongo Range. The shape of pinnae appear to dis-

tinguish this from C. moluccana, but the latter is variable and needs more study

in Borneo.

19. Cyathea altermans (Hook.) Presl, Abh. K. BoOhm. Ges. Wiss. V, 5(1848)

347. Gard. Bull. S. S. 7(1934) 219. F.M. 145 — Hemitelia alternans Hook., Ic.

Pl. 7(1844) t. 622.

Trunk usually less than 2 m tall; stipe to 60 cm, dark, basal scales to

30 X 2 mm. Pinnae jointed to rachis, lowest sometimes reduced, largest commonly

25 X 4-5 cm, in some cases to 40 X 9 cm, deeply lobed throughout (lobes entire,

rounded at apex) or with few to many of the lobes acute at apex and separately

joined by their contracted bases to axis of pinna, rarely 1-2 basal ones as quite

free pinnules; veins varying in number according to size of pinna-lobes. Sori

usually in one row on each side of costules of a pinna-lobe: indusium sometimes

178 Gardens’ Bulletin, Singapore — X XVII (1974)

completely covering sorus to maturity, more often forming an irregular disc

round base of sorus. Scales on lower surface of pinna-midrib and costules of lobes

usually sparse, narrow, setiferous; bullate scales sometimes present on costules,

sometimes also thick pale hairs.

Distribution: Sumatra, Malaya, Borneo, in forest, 1000-4000 ft.

The variable plants here included are always found growing in association

with C. moluccana, and are in several respects intermediate between C. moluc-

cana and C. squamulata (no. 29). The latter is fully bipinnate and quite exindu-

siate. If C. alternans does in fact represent a series of hybrids, the existence of

plants otherwise almost indistinguishable from C. moluccana could represent

extreme cases of introgressive hybridization. Experimental study of these plants

is desirable.

20. Cyathea trichophora Copel., Philip. Journ. Sci. 6C(1911) 363. F.M. 151.

— C. elliptica Copel., Philip. Journ. Sci. 12C(1917) 51. F.M. 146.

Trunk to 50 cm tall. Stipe 25-50 cm, + persistently scaly throughout, scales

to 20 X 3 mm, light brown, glossy; main rachis bearing narrow setiferous scales

and also hairs 2 mm long on lower surface. Lowest pinnae sometimes deflexed,

largest 30-45 cm long; largest pinnules 3-64 x 1.2-1.4 cm, lobed 4 way to costa;

costules 34-4 mm apart; veins 3-5 pairs. Sori medial, exindusiate. Hairs some-

times present on lower surface of pinna-rachis and costae, not on veins; bullate

scales on costules.

Distribution: Sarawak, Sabah, Philippines, in forest, lowlands to 4000 ft.

The type of C. trichophora was found in Luzon, that of C. elliptica on Mt

Kinabalu. Specimens on which Copeland based six other names are also here

included. The differences between them are slight. Small plants may have few

free pinnules.

21. Cyathea stipitipinnula Holtt., Kew Bull. 16(1962) 62. F.M. 147.

Stipe to more than 30 cm long, persistently scaly; scales to 25 X 3 mm, glossy

brown with paler edge bearing dark setae; minute scales also present; main rachis

glabrescent on lower surface. Pinnae to 45 cm long; pinnules to 6.5 x 1.2 cm,

lobed less than 4 way to costa, coriaceous, lowest pinnules with cordate bases

and stalks to 4 mm long; costules 34-4 mm apart; veins 3-4 pairs, thick. Sori

usually 3 to each pinnule-lobe, medial; indusium pale, firm, covering sorus to

maturity. Scales near bases of costae light brown, ovate-acute, flat, with crisped

marginal hairs or short setae, grading to light brown bullate scales on costules

and veins.

Distribution: Mt Kinabalu, Marei Parei ridge, in open places, 4000-5000 ft.

22. Cyathea assimilis Hook., Syn. Fil. (1865) 24. F.M. 150.

Stipes to 65 cm long, finely warty, persistently scaly near base; scales firm,

15-20 xX 1-2 mm. Lamina to almost 200 cm long; pinnae + jointed to rachis, lower

ones with stalks to 4 cm, largest 55 cm long; largest pinnules 8-9 x 24 cm, lowest

with stalks 4-8 mm, basal 1-2 segments of larger pinnules free, rest lobed to

1-2 mm from costa, lobes firm, crenate; costules S-7 mm apart, veins 8-10 pairs.

Sori medial; indusium pale, thin, covering sorus to maturity. Pinna-rachis glabrous

beneath; costa with narrow setiferous scales at base grading to bullate scales

distally and on costules; no hairs on lower surfaces of axes of frond.

—_ ———

Cyathea in Borneo 179

Distribution: Southern Sumatra, Sarawak, at 1000-7000 ft. Plants of ridge

forest on sandstone (Mt Dulit) have darker and more rigid fronds, sometimes

smaller than above described.

23. Cyathea discophora Holtt., Kew Bull. 16(1962) 54. F.M. 148.

Stipe persistently scaly near base; scales pale, to 25 X 2 mm, edges with short

dark setae; rachis glabrescent, finely and sparsely warty beneath. Pinnae to 50

cm long, pinnules rather widely spaced, sessile; largest pinnules 8 Xx 14 cm,

rather thin, lobed 4 towards costa except at base, lowest segment not free; costules

4-44 mm apart; veins 6-7 pairs. Sori medial; indusium a thin brown disc of

irregular shape covered by ripe sorus. Costae rather densely scaly, basal scales

pale, flat, elongate with stiff pale marginal hairs, grading to rather large pale

acuminate bullate scales distally and on costules; a few long hairs on costa near

apex of pinnule and on costules.

Distribution: only known from the type, at 8000 ft on Mt Kinabalu, in open

place in forest. This is intermediate between C. megalosora and C. squamulata.

24. Cyathea megalosora Copel., Philip. Journ. Sci. 12C(1917) 54. Gard. Bull.

S. S. 7(1934) 221. F.M. 148.

Trunk to at least 2 m tall. Stipe c. 30 cm long, densely scaly; scales thin,

pale, somewhat crisped, to 25 xX 14 mm, edges sparsely setose. Lowest pinna 20 cm,

largest 35 cm long; pinnules to 6 X 1.2 cm, almost sessile, lowest 1-2 segments

free, rest of pinnule lobed almost to costa; costules 4-5 mm apart, veins 5-7

pairs; lamina-segments very firm, crenate. Sori medial; indusium firm, covering

sorus to maturity. Pinna-rachis persistently densely scaly on lower surface, scales

long, pale, bases of smaller ones bullate; costae densely scaly, scales elongate,

not setiferous; many long hairs towards apex of costa; similar hairs on costules

and a few on veins; long pale hairs also on upper surface of costules and veins.

Distribution: Mt Kinabalu, in mossy forest on ridges at 7000-10000 ft; plants

at highest altitudes have smaller fronds, with pinnae to 104 xX 24 cm, free pinnules

c. 5 pairs, lobed only 4 way to costa.

25. Cyathea trichodesma (Scort.) Copel., Philip. Journ. Sci. 4C(1909) 55. F.M.

150. — Alsophila trichodesma Scort. in Bedd., Journ. Bot. 25(1887) 321. —

C. burbidgei sensu C. Chr., Gard. Bull. S. S. 7(1934) 222; sensu Holttum, Rev.

Fl. Mal. 2(1955) 124, non (Bak.) Copel.

Trunk to 44 m tall. Stipe densely scaly near base; scales medium to light

brown, firm, to 25 x 2 mm, closely setose. Pinnae to 60 cm long; pinnules in

Malaya 9-11 xX 14-2 cm, on Mt Kinabalu to 7 x 14 cm, nearly sessile, lobed to

2 mm from costa, no free basal segments; costules 44-5 mm apart; veins 6-8

pairs. Sori medial, often confluent at maturity, no indusia. Pinna-rachis, costae,

costules and veins bearing many pale erect hairs 1-2 mm long on lower surface;

scales on costae and costules sparse, pale, some bullate, most with dark setae;

in Malaya hairs present on upper surface of costules and veins.

Distribution: Malaya, Sarawak, Sabah, in lowland forest; to 5000 ft on Mt

Kinabalu. The Kinabalu specimens are perhaps smaller than those in Malaya

Owing to altitude.

180 Gardens’ Bulletin, Singapore — X XVII (1974)

26. Cyathea walliacei (Kuhn) Copel., Philip. Journ. Sci. 4C(1909) 48. F.M.

151. — Alsophila wallacei Kuhn, Linnaea 36 (1869) 153. — A. burbidgei Bak.,

Journ. Bot. 17(1879) 38; not Cyathea burbidgei sensu. C. Chr., Gard. Bull. S. S. 7

(1934) 222, which is C. trichodesma.

Stipe 30 cm or more long, pale and smooth above base; basal scales to 15 x 2

mm, pale brown; main rachis finely hairy on distal part of lower surface. Largest

pinnae 38 cm long; pinnules to 6.5 x 1.3 cm, sessile, lobed to within 1 mm from

costa; costules 3-34 mm apart; veins 4-6 pairs, mostly simple. Sori medial, no

indusia. Lower surface of pinna-rachis, costae, costules and veins bearing pale

hairs 1 mm long; pale bullate scales on costae and costules; long hairs present

on upper surface of costae and costules.

Distribution: Sarawak, Sabah, in lowland forest (including Bako National

Park, near Kuching). This differs from C. trichodesma in more deeply lobed

pinnules and closer costules. It has the aspect of a stunted member of sect.

Sphaeropteris but has the venation of sect. Schizocaena.

27. Cyathea polypoda Bak., Trans. Linn. Soc. II Bot. 4(1894) 250. F.M.

151. — C. kemberangana Copel., Philip. Journ. Sci. 12C(1917) 52. Gard. Bull.

S. S. 7(1934) 200, 219. — C. ampla sensu Holtt., Rev. Fl. Mal. 2(1955) 125, non

Copel.

Trunk to 3 m tall; small branches frequent on its lower part. Stipe to 80 cm,

green; basal scales medium brown, firm, to 30 c 2 mm, setae concolorous; rachis

glabrescent on lower surface. Pinnae to 60 cm long; lower ones long-stalked;

largest pinnules 84-11 xX 2.0-2.7 cm, thick and rigid when dry, all stalked, stalks

of lowest 9 mm; basal 1-2 segments of largest pinnules free, rest of pinnule

lobed to 1-2 mm from costa; costules 44-54 mm apart; veins 7-9 pairs. Sori

inframedial, no indusia. Small dark or brown setiferous scales near bases of

costae; bullate scales, often setiferous, on costules, all early caducous.

Distribution: Malaya, Borneo, Philippines (Panay, Mindanao); in open places

on ridge-crests and summits, 2000-7000 ft; specimens from the higher altitudes

very tough. Baker described the type of this species as indusiate; the “indusia”’

which he saw were bullate scales partly covering young sori.

28. Cyathea agatheti Holtt., Kew Bull. 16(1962) 51. F.M. 152.

Trunk hardly 5 cm tall. Stipe 35-75 cm long, dark towards base, rest smooth,

green; basal scales 10 xX 2-24 mm, light brown with dark setae on pale edge.

Lamina of frond 50-60 cm long; pinnae jointed to rachis, largest 18-25 cm long;

pinnules to 34 X 1 cm, lobed to within 1 mm from costa, lowest with stalks 1-mm

long; costules 3 mm apart; veins 4-5 pairs, simple. Sori medial, no indusia. Lower

surface of costae bearing scattered pale hairs, few hairs on costules; scales few, not

bullate; upper surface of costae, costules and veins bearing long hairs.

Distribution: only known from original collection, W. Kutai, Kalimantan, in

Agathis forest on water-logged white sand, at 2000 ft.

29. Cyathea squamulata (BI.) Copel., Philip. Journ. Sci. 4C(1909) 37. C. Chr. in |

Gard. Bull. S. S. 7(1934) 219 excl. syn. C. elliptica Copel. Holtt., Rev. Fl. Mal.

2(1955) 122, fig. 49. F.M. 152.

Cyathea in Borneo 181

Trunk to 2 m tall. Stipe 40-60 cm, persistently scaly throughout (main rachis

sometimes also); scales firm, medium brown with dark setae, to 30 xX 2-3 mm.

Lowest pinnae somewhat reduced, variable, largest 50 cm long; largest pinnules

8-10 xX 14-2 cm, rather thin with almost entire lobes, lobed 4-4 towards costa

(more deeply only in largest); costules 34-44 mm apart; veins 6-9 pairs, simple

in smaller pinnules, forked in larger. Sori a little inframedial, no indusia; pale

paraphyses longer than sporangia. Pinna-rachis glabrescent beneath, finely warty;

costae with flat brown setiferous scales at base, grading to bullate distally and on

costules; no hairs on lower surface of costules, a few on upper surface.

Distribution: Sumatra, Java, Malaya, Borneo, Sulu Archipelago, in lowland

forest; in Malaya rarely above 3500 ft; specimens from higher altitudes (to 7000

ft) on Mt Kinabalu are small, with densely scaly rachis.

Addendum: A new species from Mt Kinabalu

In 1966 Mr E. F. Allen collected a small plant of a tree fern at an altitude of

10,000 feet on Mt Kinabalu, and brought it to his home, near Ipswich in England,

where he successfully cultivated it. It flourishes in the open air in summer, but

in the winter needs the protection of a cool greenhouse. This plant has now pro-

duced a fertile frond, and proves to be distinct from any previously known species.

The key to the species of subg. Alsophila sect. Alsophila needs to be modified

at indentation 5 as follows, to accommodate the new species and to indicate its

distinctive characters.

Young sorus completely covered by indusium which breaks at maturity, leaving

the exposed sporangia surrounded by an irregular cup .................. C. oosora

Indusium hood-shaped, attached only on the side towards the costule, not

quite covering sorus at maturity:

Basal scales on stipe firm, pale, to 25 x 3 mm; stipe short, its pneumatodes

to 11 mm long; several pairs of lower pinnae gradually reduced, lowest 7

cm long; pinnules to 94 cm long, less than 2 cm wide; costules 34-4 mm

apart; small scales mostly lacking rigid marginal setae, bullate scales

Peeeianh rencias ait costiles int. 2iic2iees sda. aya AE od: C. loheri

Basal scales of stipe light brown, thin, to 10 xX 14 mm; stipe to 50 cm

long its pneumatodes 3 mm long, well spaced; lower pinnae little reduced;

fertile pinnules 7 X 2.2 cm, costules 44-5 mm apart; nearly all small

scales bearing several rigid setae; bullate scales few, often setiferous ......

See Sererene baw Prete te. eee ee OY SP et hee dee C. brachyphylla

Cyathea brachyphylla Holttum, sp. nov.

C. loheri affinis, differt: frondibus minoribus; stipite 50 cm longo, basi paleis

tenuibus usque 10 x 14 mm vestita, sursum paleis minutis plerisque copiose setiferis

praedito; pneumatodiis 3 mm longis, valde dissitis; pinnis inferioribus paulo reductis;

pinnis maximis 33 cm longis; pinnulis usque 7 X 2.2 cm; costulis 44-5 mm inter se

distantibus; paleis rhachidum costarumque fere omnibus setiferis, paleis bullatis

paucis distalibus etiam plerisque setiferis.

Type: cult. E. F. Allen, Ipswich, England, origin Mt Kinabalu, alt. 10,000 ft

(holotype K; isotype SING).

182 Gardens’ Bulletin, Singapore — X XVII (1974)

Mr Allen reports that he saw plants with trunks up to 8 feet (240 cm) tall.

They were growing in deep soil in a sheltered place, not on the crest of the ridge

where Cyathea havilandii is abundant at about the same altitude.

As indicated in the revised part of the key, the new species is related to

C. loheri in the form of its sori, but differs in its smaller size with fronds of a

different shape, and in the scales of all parts of the frond. The other two species of

Cyathea which have previously been found near 10,000 feet altitude (C. havilandii

and C. oosora) have sori of a different structure; careful examination with a hand

lens is necessary to distinguish the difference.

Morphological studies on some inland Rhizophoraceae

GEH SIEW YIN! ET HSUAN KENG?

I. Introduction

The family Rhizophoraceae (as Ordo Rhizophoreae) was first established by

R. Brown in 1814, and includes Rhizophora L., Bruguiera Lamk., and Carallia

Roxb. Other genera were subsequently added by Bentham and Hooker and others

and the Rhizophoraceae has since been expanded to comprise 16 genera and

approximately 120 species of trees and shrubs (Hou, 1958) which are pan-tropical

in distribution.

The best known representatives of the family are four mangrove genera, viz.

Rhizophora, Bruguiera, Ceriops, and Kandelia, which are characterized by their

peculiar adaptive features of viviparous fruits, in which the seeds undergo germina-

tion on the parent plant, and by their various special roots (pneumatophores and

knee roots), which enable the plants to thrive in a muddy, littoral habitat. These

mangrove genera have been extensively studied by many authors. The majority

of the Rhizophoraceous genera are, however, inland plants which exhibit none of

these special adaptive characters and are relatively less investigated. It is because ot

this that the classification of the family has been subjected to much revision and

rearrangement.

The present investigation is mainly confined to the species of the following

5 inland Rhizophoraceous genera, viz., Anisophyllea, Carallia, Combretocarpus,

Gynotroches and Pellacalyx, found in Singapore and Malaysia.

Members of the inland Rhizophoraceae are usually sun-loving, gregarious,

and generally found in secondary forests. Because they are normally small trees,

their timbers, except for construction, possess little or no commercial value, yet

from the ecological point of view they are among the important pioneer trees

which prepare a suitable environment for other superior trees such as dipterocarps

and others to grow.

The specimens of 8 species belonging to 5 different genera examined were

partly fresh material collected in Singapore, and partly from herbarium or wood

collections of the Botanic Gardens, Singapore, from Dr. T. C. Whitmore, formerly

of the Forest Research Institute, Kepong, and from Dr. J. A. R. Anderson, former

Conservator of Forests, Sarawak. The nomenclature of the material used follows

a recent taxonomic treatment by Hou (1958).

Fresh material including leaves, stems, flowers, fruits and seedlings were

preserved in formalin-acetic acid-alcohol (F.A.A.). Infiltration of the fluid was

facilitated by the use of the suction pump. Material taken from the herbarium

collections were first softened in a dilute solution of sodium hydroxide (5%) then

poroughly rinsed in water and stored in 70% alcohol. Wood specimens were cut into

1. Botanic Gardens, Singapore.

2. Department of Botany, University of Singapore.

183

184 Gardens’ Bulletin, Singapore — X XVII (1974)

small cubes and boiled for 1 ~ 2 hours to remove the air, transferred to 30%-40%

hydroflouric acid for periods of one month or longer, washed overnight in running

water and then stored in glycerin-alcohol.

Serial sections of twigs, petioles, leaves, flower buds and mature flowers were

obtained by the paraffin method. Dehydration and embedding were carried out

by the tertiary butyl alcohol series in accordance with the method described by

Johansen (1940). Sections of 8-15 yu were obtained by means of the rotary

microtome, and these were then stained with a safranin and fast-green combination.

Semi-permanent preparations of pollen grains were obtained by scraping

pollen grains from recently dehisced anthers into lactophenol, and sealing the

mounted slides with nail polish.

Cleared leaves and floral parts were prepared according to a modification of

Foster’s (1949) and Arnott’s (1959) methods.

Transverse, tangential, and radial longitudinal sections of wood material

were obtained with a sliding microtome. The sections were stained in safranin,

dehydrated through the xylene-alcohol or tertiary-butyl-alcohol series and mounted

in euparol. Fragments of wood were macerated in Jeffrey’s nitric-chromic acid

mixture.

Il. External Morphological Observations

External morphological observations on fruits, seeds and seedlings of various

genera and species were made during the course of this study.

Fruits of Anisophyllea disticha, Carallia brachiata, Gynotroches axillaris,

Pellacalyx axillaris and P. saccardianus were collected and the following observa-

tions recorded. Attempts to germinate the seeds from these fruits proved to be

unsuccessful, although several batches of seeds (8 for Anisophyllea disticha; 30-40

for the other species) in different media (soil, sand, filter paper) and under light

and dark conditions were experimented with. Consequently only the seedling stages

of two species, Anisophyllea disticha and Gynotroches axillaris, found in the field

are described below. Fruits of Combretocarpus rotundatus are described from

herbarium and pickled material obtained from Kuching, Sarawak and from the

Forest Research Institute, Kepong, Malaysia.

a. FRUIT

Genera of the Rhizophoraceae appear to differ considerably in fruit structure.

Baccate fruits are present in Carallia (Fig. 3 C), Gynotroches (Fig. 5 C) and

Pellacalyx (Fig. 6 C), while Anisophyllea (Fig. 2 C) produces drupaceous fruits and

Combretocarpus (Fig. 4 D) dry, indehiscent, winged fruits. Plants of the mangrove

genera, Rhizophora, Bruguiera, Ceriops and Kandelia, are well-known for their

peculiar one-seeded, vivaparous fruits.

The fruits of the Gynotrocheae vary in size from small and globular in

Carallia brachiata (2.5-3.5 mm across) and Gynotroches axillaris, to large and

turbinate in Pellacalyx axillaris and P. saccardianus (1-1.5 cm across). The fruits of

these plants are further differentiated by the glabrous character of the former two

species and the puberulous exterior of the Pellacalyx fruits.

The fruits of Carallia brachiata are many-seeded with the light-brown seeds

embedded in an axile placenta. The pericarp changes from green, through trans-

lucent pink to red as the fruit matures. Gynotroches axillaris produces similar

Morphological Studies of Some Rhizophoraceae 185

fruits which ripen to a shiny black on maturity. The fruits borne by Pellacalyx

axillaris and P. saccardianus are approximately four to five times the size of the

other two genera, and are very similar to one another in size; P. saccardianus can,

however, be distinguished by its four-shouldered berries which are situated on

longer stalks and are less densely puberulous than those of P. axillaris. Mature

fruits of Pellacalyx are yellowish-green in colour.

The fertilised ovary of Anisophyllea disticha develops into a succulent drupe

which ripens from pale pink to bright red. These fruits are often found in pairs

or singly, on the underside of the pendent branches. The succulent mesocarp

encloses a yellowish-brown stone which is acute at both ends and ridged longitu-

dinally; these ridges are obvious in dried specimens of the fruits. Combretocarpus

rotundatus produces a winged fruit in contrast to the other members of the family.

There are usually three, less often four, membranous extensions of the pericarp

which are delicately transversely veined. The single enclosed seed is spindle-shaped

and very slender.

The fruits of all the species examined possess persistent calyx lobes. These

lobes are situated above the pomaceous fruits which are formed by inferior or

half-inferior ovaries except in Gynotroches axillaris where it is found below the

berry. Very often, remains of stamens and style are also persistent on the fruit but

the petals of all species are caducous. The persistence of the calyx is also notable

in the fruits of the mangrove genera where they often become accrescent. There is

hardly any other common feature between the fruits of the tidal and inland species

than this.

b. SEED

Plants of the Gynotrocheae are characterized by comparatively small but

several to numerous seeds embedded in pulpy placentae of baccate fruits. Each

locule of the ovary in Carallia brachiata houses two ovules, while in the other

two genera, there are several to many ovules per locule. The ovules of Pellacalyx

species are arranged in fascicles of 8-25 attached to an axile placenta while those

of Gynotroches axillaris are found embedded in a mass of placental and endocarpic

tissue.

The seeds of these species are light yellowish-brown to dark brown in colour,

and the testa is often crustaceous or sometimes coriaceous. Endosperm is always

present surrounding the embryo which is axile, linear and practically extending

from one end of the seed to the other.

Both Combretocarpus rotundatus and Anisophyllea disticha have fairly large

single-seeded fruits. In C. rotundatus the pendulous seed lies in the centre of the

fruit attached to the apex of the locule and is enclosed by a thin dry pericarp which

is extended externally into 3-4 wings. The embryo is linear and cylindrical, and

sections of the fruit reveal the presence of two distinct cotyledons and a fairly long

hypocotyl; the plumule within the cotyledons is rudimentary. Surrounding the

embryo is a structurally differentiated endosperm which is composed of densely

cytoplasmic and more darkly stained cells. There is a clear demarcation between

the embryo and its surrounding tissue, contrary to Hou’s observation (1958) that

the embryo and endosperm form a solid whole and are not differentiable under

the microscope.

Freehand and microtome sections of the fruit and seed of Anisophyllea disticha

at different stages show that the embryo is yellowish-white, axile, linear and

embedded in a tissue which is endospermous in nature (Fig. 2 C 1 to C 35).

186 Gardens’ Bulletin, Singapore — X XVII (1974)

Hou (1958), possibly from dried material, has decribed the embryo and its

enveloping tissue as forming a solid unity. This was not found to be so in the

fresh specimens examined, as in all cases the embryo proper was quite naturally

separable from its surrounding tissues. In the young seed, the endosperm takes

up most of the space with the embryo confined to the apical portion, but as the

seed matures, the embryo gradually occupies the entire central region, at the

expense of the endosperm. Thus in longitudinal sections of a mature seed, it

reveals the embryo as a terete structure extending almost the entire length of the

seed. At the apical portion of the embryo two protuberances are observed which

can be interpreted as cotyledons*. The rest of the embryo is a solid cylindrical

structure with a sheath of small-celled vascular tissue separating the cortical from

the central region. Between the cotyledons is the plumule which elongates into a

young shoot and later bears the scale-like cataphylls of the seedling.

The seeds of the mangrove members of the Rhizophoraceae are distinguished

by their conspicuous hypocotyls which develop while the fruit is still attached to

the parent plant. In Rhizophora and Bruguiera the two cotyledons are connate into

a sheath and surround the plumule, serving to assimilate nutrients from the

endosperm for the developing embryo. This form of cotyledons is not confronted

in the inland genera and is absent in Anisophyllea disticha where the entire,

undifferentiated embryo is embedded in endospermous tissue.

c. SEEDLING

Although usually abundant fruits and seeds are produced by individual plants

belonging to the inland Rhizophoraceous genera, the number of seeds which

actually germinate appears to be very small indeed. Therefore, seedlings of these

species are not commonly found on forest floor. Fresh seeds of several species

(including Carallia, Gynotroches, Pellacalyx, Anisophyllea) collected in the field

mostly failed to germinate in the laboratory and consequently only seedlings of

the following two species are described.

Germination in Anisophyllea disticha seed is distinctly hypogeal (Fig. 7). The

seed lies prostrate on the ground at first, then is carried downwards together with

the radicle as the latter penetrates the soil, and soon produces lateral roots. Roots

are formed from both ends of the seed, with the plumular end producing a less

developed root system. The cotyledons and hypocotyl do not emerge outside the

seed, but the plumule elongates into a strong cylindrical stem portion bearing

small scale-like cataphylls before the first whorl of lateral branches is produced.

Gynotroches axillaris seed germinates into a seedling which does not produce

cataphylls. Cotyledons have not been found on any of the seedlings observed, thus

it is not possible to determine if germination was epigeal or hypogeal (Fig. 8).

However, since in the mangrove genera the first green leaves are not cotyledonary,

and the germination of Anisophyllea is hypogeal, it is perhaps reasonable to presume

that the germination in Gynotroches and other inland plants may also be hypogeal.

Seedling leaves of G. axillaris resemble very closely the mature leaves in their

general shape, colour, texture and especially in venation. Another prominent feature

* The seed structure of Barringtonia as observed by Payens (1967), in general, is very

similar to that of Anisophyllea disticha; in both cases, the embryos are not clearly differentiated

into plumule, cotyledons and hypocotyl. The two protuberances at the apical end of the

embryo of Barringtonia are considered by Payens to be scales, not cotyledons, on account

of their spiral arrangement and the presence of axillary meristems. Those at the apical end

of the embryo of Anisophyllea disticha, however, are strictly oppositely arranged, therefore

can be reasonably interpreted as cotyledons.

—_ 77 _-~—tt—=wte EE

Morphological Studies of Some Rhizophoraceae 187

is the conical interpetiolar stipules which cover the apical developing buds of the

plants; the stipules in the Rhizophoraceae overlap one another leaving one free

margin on each side, except in the case of Pellacalyx where the stipules are flat

as in most Rubiaceae. The presence of such overlapping stipules may hence be

used as a guide for identifying Rhizophoraceous seedlings, except in Anisophyllea

and Combretocarpus which are exstipulate.

Ili. Anatomical Studies

Serial sections were made of fresh specimens of the stem, wood, leaf, petiole

and flowers of Anisophyllea disticha, Carallia brachiata, Gynotroches axillaris,

Pellacalyx axillaris and P. saccardianus collected in Singapore, and of herbarium

collections of the flowers of Combretocarpus rotundatus and the wood of

Anisophyllea corneri, Carallia eugenioidea and C. rotundatus obtained from Dr.

J. A. R. Anderson, Sarawak and from Dr. T. C. Whitmore, Kepong, Malaysia.

A brief summary of the anatomical details of the leaf-blades, petioles, stems,

woods, flowers and pollen-grains of the species investigated is presented below.

a. LEAF-BLADE (Table 1; Fig. 15)

(1) Stomata. The stomata in the species of the Rhizophoraceae investigated

belong to the ranunculaceous or anomocytic type (Metcalfe & Chalk, 1950), ie.

the epidermal cells surrounding the two guard cells are indistinguishable from the

other epidermal cells; sometimes, however, the cells around the stomata are more

oblong than their neighbouring cells. Stomata are confined to the lower leaf

surface only.

(2) Epidermis. The upper epidermis varies from the uniseriate in Anisophyllea

disticha and Carallia brachiata, to 2-5 layered in Gynotroches axillaris, Pellacalyx

axillaris and P. saccardianus. The outermost layer of a multiple epidermis resembles

the ordinary uniseriate epidermis in having a cuticle (absent in P. axillaris), while

the inner layer or layers (or the hypoderm) are composed of larger cells which do

not contain chloroplasts, and commonly serve as a water storage tissue. Among the

hypodermal cells are frequently large mucilage producing cells which are especially

conspicuous in P. axillaris. The thickest hypoderm, of 3-4 layers, is found in

P. saccardianus.

The lower epidermis in all the species examined is single-layered. It is covered

by a thin cuticle except in Carallia brachiata and Pellacalyx axillaris where it is

nearly absent.

Cork-like warts occur as small black dots on the lower surface of Carallia

brachiata, while tufted trichomes are present especially on the abaxial surfaces of

Pellacalyx species. Crystalliferous inclusions in the form of druses are usually

present in the upper epidermal cells of C. brachiata.

In contrast to the thin cuticle present in the inland genera, the mangrove

Species are generally covered by a thick cuticle which may be composed of two

distinct layers. Multiple epidermis is also common among the mangrove genera

although species of Bruguiera may have a uniseriate epidermis. The number of

layers in the hypodermis has been directly correlated to the concentration of salt

in the water where the plants are growing, in the case of Rhizophora mangle Linn.

(Bowman, cited in Metcalfe & Chalk, 1950).

188 Gardens’ Bulletin, Singapore — X XVII (1974)

(3) Mesophyll. The mesophyll is differentiated into palisade and spongy tissues.

In A. disticha the leaf is isobilateral, and the single-layered palisade is distinguished

from the spongy parenchyma only by nature of its more longitudinally elongated

axis. C. brachiata also shows a one-layered palisade tissue which is distinct from

the spongy mesophyll and separated from it by 1-2 layers of parenchyma ceils

with tanniferous inclusions. The other species investigated have 2-4 palisade layers

which are closely packed in the first layer but become more loosely arranged, and

scarcely distinguishable from the spongy mesophyll in the case of P. saccardianus.

Palisade cells of P. axillaris are very much narrower and anticlinally elongated

than the other cells of the mesophyll.

The number of cells in the spongy mesophyll ranges from 6-17 rows in the

species examined. A. disticha and P. axillaris both have approximately 6 rows of

cells which are more compactly arranged in the former, but include extensive air-

spaces in the latter. In G. axillaris the spongy mesophyll cells are small in size

and loosely arranged. The lower 3-5 layers of cells are larger and more compact

in arrangement forming a pseudo-hypodermis, which as defined by Metcalfe and

Chalk (1950) is formed of cells not aligned with the epidermal cells and ontogene-

tically not derived from them. Mucilage cells are also observed in the pseudo-

hypodermis.

Cells of the spongy mesophyll in C. brachiata, G. axillaris and P. saccardianus

contain darkly stained, probably tanniferous substances. These are found in the

first 2-3 layers of the spongy mesophyll in C. brachiata, in the last 2-3 layers

beneath the lower epidermis in P. saccardianus, and scattered all over the mesophyll

in the case of G. axillaris. Druses are also found occasionally in the parenchyma

cells of all the species examined.

The proportion of spongy mesophyll in relation to the total leaf thickness, in

average, is found as follows: — A. disticha 72%; C. brachiata 75%; G. axillaris

69%: P. axillaris 63%; P. saccardianus 57%. Thus, a well-developed spongy

mesophyll is present in these species.

(4) Vascular Bundles. The mid-rib bundle is identical to and a smaller version

of the median petiolar bundle in all the species under observation. The vascular

bundles of the leaf blades are of the collateral type with xylem on the adaxial side

and phloem below it, facing the lower epidermis. The larger veins extend from the

upper epidermis or hypodermis to the spongy mesophyll, while the smaller veins

are present only in the spongy tissue.

The larger vascular bundles are completely encircled by two types of bundle

sheaths. The inner sheath is composed of thick-walled sclerenchymatous cells

and this is overlaid by a single layer of parenchymatous tissue. Sclerenchyma

frequently encloses the smaller veins of the leaf lamina which may be composed

only of xylem elements. Phloem cells are absent from vein endings which are

composed mainly of elongated tracheids surrounded by a parenchymatous bundle

sheath.

Bundle sheath extensions are not well-developed in the inland species examined,

and this is probably due to the increase of the thickness of the spongy parenchyma

as suggested by Wylie (1943).

(5) Sclereids. Sclerenchymatous idioblasts are notably absent in the species |

investigated in contrast to the well-developed H-shaped sclereids in the palisade >

and variously branched idioblasts in the spongy mesophyll of Rhizophora species. —

Solitary sclerenchymatous cells have also been recorded in the spongy mesophyll —

of Poga oleosa by Schimper (1898).

189

Morphological Studies of Some Rhizophoraceae

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190 Gardens’ Bulletin, Singapore — X XVII (1974)

b. PETIOLE (Fig. 16)

Serial sections made through the distal end of the petiole, immediately below

the lamina, show that the main vascular strand in this region is mostly U-shaped.

The two margins of the median bundle have a tendency to curve inward in

C. brachiata and in Pellacalyx species. The median strand of G. axillaris is a shallow

crescent-shaped structure with another small strand fitting into the hollow of the

crescent. There is no sclerenchymatous tissue surrounding the vascular strands, and

1-3 accessory bundles may be found on either side of the main trace.

The petiole of P. axillaris is characterized by the presence of large mucilaginous

cells in the peripheral region of the cortex. Darkly stained amorphous inclusions

and, occasionally, druses are present in the parenchymatous tissue of the cortical

region of the four other species.

c. STEM (Fig. 17)

The young branches are oblong or elliptical in transverse section. The outer

surfaces of the stems are glabrous in C. brachiata and G. axillaris, but possess

trichomes in the case of A. disticha, P. axillaris and P. saccardianus. The epidermal

cells are covered by a thin cuticle and the cortex is composed of closely arranged

ground tissue. Many of the cortical cells in the species investigated contain darkly

stained inclusions. A continuous ring of sclerenchyma forming the pericycle

surrounds the vascular cylinder in A. disticha, and is absent in the other species.

The vascular bundles are collateral, with phloem surrounding the xylem cylinder.

The pith is very wide in all cases and is composed of thin-walled cells which are

larger in the central region. In the older branches of Gynotroches and Pellacalyx

species, the central pith cells disintegrate leaving a characteristic hollow, which

is one of the field characters utilized in identifying these two genera. Cork

formation is sub-epidermal in origin.

The nodal structure in this family is trilacunar and many-traced, and agrees

with the observations by Sinnott (1914). The Rhizophoraceae has been included

customarily under the Myrtales which, however, is characterized by unilacunar

nodes.

d. WOOD ANATOMY (Tables 2 & 3; Figs. 18 & 19; Plate 2)

The wood structure of the five genera examined fall in harmoniously with

one another in most features. Except that Pellacalyx exhibits vessels with opposite

pitting to the ray cells and has the tendency for the pore clusters to be tangentially

arranged, and the occurrence of scalariform pitting in G. axillaris, there is no

sharp demarcation in the wood anatomy of the species investigated.

Growth rings are indistinct in members of the Rhizophoraceae and the wood

is commonly diffuse porous. The pores are solitary (especially in Anisophyllea),

or arranged in pairs, or in clusters (tangentially arranged in Pellacalyx). Pores are

usually scanty in Anisophyllea and Combretocarpus (3—6/mm?) but more abundant

in Gynotroches where they may number up to 25-30/mm?. The pores are mostly

spherical in outline and the walls are rather thin and evenly thickened. The largest

pores (up to 350 ») are found in Anisophyllea and Combretocarpus, while in the

other genera they are small to medium-sized. The length of the vessel members

range from medium-sized to very long in C. brachiata (to 1400 p).

Pits to ray cells are generally oval and alternate in arrangement, but in

G. axillaris they are exclusively scalariform and elongate, while in Pellacalyx they

are opposite and angular.

191

Morphological Studies of Some Rhizophoraceae

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TABLE 3

SUMMARY OF WOOD ANATOMY CHARACTERS OF THE TRIBES OF

RHIZOPHORACEAE

Character

Vessel member

Anisophylleeae

Gynotrocheae

Rhizophoreae

1. Arrangement Often solitary, some- | Solitary or more | Solitary or in multi-

times paired. often in pore multi- | ples of 2-14.

ples of 2-5.

2. Abundance 2-5 per mm2 7.25 per mm2 5-35 per mm2

3. Diameter and length (D.) (L.) (D.) S| i Be (L.)

lowest average 150 pp 380 pu 116 p 584 ph

mean of averages 292 p 483 pb 152 p 671 p

highest average 347 p 738 pp 209 pu 720

4. Pits to ray cells Alternate; oval Alternate oval or | Scalariform.

scalariform or oppo-

site angular.

5. End wall type Oblique-long caudate} Oblique steeply inclined taper-

ing ends.

Wood fibre

6. Wall thickness Thick Thick Thick

7. Length

lowest average 1,350 p 1,014 p

mean of averages 1,813 p 1,385 pp

highest average 2,550 pu 1,740 pw

Wood ray

8. Krib’s Type Heterogeneous I12 Heterogeneous II2 Heterogeneous Type

Ila or Homogeneous

(Rhizophora)

9. av. length of uniseriate | 13 cells 9 cells

rays

10. av. width of uniseriate | 6 cells 10(-26) cells 3-6 cells

rays

Wood parenchyma

11. Distribution Apotracheal or Paratracheal Scanty vasicentric

paratracheal confluent

Perforation plates are simple, or rarely, multiple in Anisophyllea, End walls

are oblique and long caudate in Anisophyllea or oblique with less pronounced

prolongations in others.

Wood fibres are thick-walled in all species examined, with a range in length

of extremely short (320 » in C. brachiata) to very long (2,800 » in Combretocarpus).

Wood rays are consistently heterogeneous of Krib’s Heterogeneous Type Ila.

Uniseriate rays may be present as 2-3 cells or 28 cells (in Gynotroches) but are

more often 4-10 cells high in Gynotroches. Multiseriate rays are commonly 100

or more cells high, to over 400 cells in Carallia. Cells of the rays may contain

amorphous depositions in species of all the genera, and are most abundant in

Carallia, but rarely found in Pellacalyx.

Morphological Studies of Some Rhizophoraceae 193

Wood parenchyma is often paratracheal confluent in Anisophyllea, Combre-

tocarpus, Carallia and Pellacalyx, and metatracheal bands are in addition, present

in the former three genera. G. axillaris shows a tendency towards aliform arrange-

ment of the axial parenchyma as well as the characteristic paratracheal confluent

distribution of Anisophyllea, Combretocarpus and Gynotroches.

Among the wood specimens of the various species examined, those of the

following three are investigated for the first time.

(1) Anisophyllea corneri D. Hou

(2) Carallia eugenioidea King

(3) Pellacalyx saccardianus Scort.

The main evolutionary trends of wood anatomy characters (pertinent to the

wood anatomical characters of the Rhizophoraceae) are summarised in Table 4.

TABLE 4

SUMMARY OF EVOLUTIONARY TRENDS OF WOOD ANATOMY CHARACTERS

Character

Wood porosity

Vessel elements

Primitive

diffuse-porous

Intermediate

semidiffuse-porous

Advanced

ring-porous

(a) Arrangement solitary paired clustered in pore

multiple

(b) Outline in t.s. angular oval or rounded

(d) Length .. | very long very short

(e) Perforation plate .. | Scalariform with reticulate simple

many bars

(f) End-wall very oblique, long | oblique, short horizontal

caudate

(g) Pits to ray cells Scalariform, elong- | opposite;transitional;| alternate; rounded.

ated oval

Wood fibres

(a) wall thickness very thin very thick

(5) length very long very short

Wood rays

(a) Krib’s Type Heterogeneous Heterogeneous Heterogeneous

Type I Type Ila Type Ill

(6) Length of uniseriate | very long very short

rays

Wood parenchyma

(a) Distribution diffuse-scanty paratracheal apotracheally banded

Following the putative phylogenetic trends (Carlquist 1961), the vessel members

of the Rhizophoraceae may be considered advanced in the possession of simple

perforation plates. Although the lengths and widths of the vessels in the genera

investigated do not differ greatly, it is evident that Carallia possesses the longest

vessel element length and the narrowest pore diameter, while Anisophyllea and

Combretocarpus show more advanced, broader and shorter vessel members. A

further advanced feature lies with the occurrence of alternately arranged pits to

contiguous ray cells, but this character is not common to all five genera;

Gynotroches shows a primitive scalariform pattern, while Pellacalyx possesses

Opposite pits, which condition is considered intermediate between those found in

194 Gardens’ Bulletin, Singapore — X XVI] (1974)

Gynotroches on the one hand, and Anisophyllea, Combretocarpus and Carallia on

the other. In the arrangement of the pores, the solitary condition, often found in

Anisophyllea is a less evolved condition which apparently progresses through that

found in Carallia and Gynotroches to the pore clusters of 2-5 found in Pellacalyx.

The longest wood: fibre lengths are found in Combretocarpus (2800 p), whereas

the shortest in Carallia (395 ). In this instance therefore, the wood of Carallia

shows a more advanced character in contrast to its other more primitive vessel

elements.

Wood rays of the genera studied are uniformly heterogeneous of Krib’s Type

Ila. The multiseriate rays are broad to very broad (28-30 cells) in Gynotroches

and generally over 100 cells high. The highest multiseriate rays are found in

Carallia (up to 500 cells). Uniseriate rays are shortest on the average in Gynotroches

and longest in Anisophyllea in contrast to the heights of multiseriate rays found

in the respective genera.

All members of the five genera show a tendency towards a paratracheal

confluent arrangement of wood parenchyma, which is considered an intermediate

condition in wood parenchyma phylogeny. Metatracheal bands also occur in

Anisophyllea, Combretocarpus and Carallia.

From the data obtained, it is difficult to consider any one genus as more

advanced than the other. Anisophyllea and Combretocarpus probably show a slight

advantage in the number of advanced features than the other genera, but on the

other hand, these features alone appear to be insufficient to consider that these

two genera are phylogenetically more evolved.

According to the observations of Marco (1935) and Metcalfe and Chalk

(1950), the members of the mangrove Rhizophoraceae, likewise, show a mixture

of advanced and primitive wood anatomy features, but in this group, the primitive ~

characters are more in preponderance, e.g. scalariform pitting on the lateral walls,

very oblique end walls, and scanty wood parenchyma. The genus Rhizophora

seems more advanced than the other three genera of the mangrove Rhizophoraceae

in the possession of homogeneous wood rays.

Considering the wood anatomical features of the family Rhizophoraceae as

a whole, the mangrove members which constitute the tribe Rhizophoreae may be

regarded as the most primitive, and the inland members, especially Anisophyllea

and Combretocarpus which constitute the tribe Anisophylleeae, the most advanced.

e. FLOWER AND POLLEN GRAINS

The inland Rhizophoraceae produce axillary flowers either in condensed

clusters (in Gynotroches (Fig. 5), Carallia (Fig. 3; Plate 1) & Pellacalyx (Fig. 6;

Plate 1)) or in fascicles or racemose inflorescences (in Anisophyllea (Figs. 1 & 2)

& Combretocarpus (Fig. 4)). In contrast, genera of the mangrove Rhizophoreae

exhibit paired or cymose arrangement of the flowers. The basic inflorescence

structures of the family, nevertheless, are probably cymose in nature.

The accessory floral parts are always valvate in aestivation and vary in

number from 3-4 (in Anisophyllea, Combretocarpus) to 4-15 (in Rhizophora,

Bruguiera). The sepals are always connate at the base into a calyx-tube with

triangulate or linear calyx-lobes, each of which receives a constant supply of a

single vascular trace. The petals are often clawed in Gynotroches, Carallia and

Pellacalyx, and apically fringed in various degrees. In Anisophyllea and Combreto-

carpus the petals are deeply and evenly 3-5 lobed, or entire, while in the mangrove

Rhizophoreae the petals are entire or bi-lobed and often associated with special

ms iat

Morphological Studies of Some Rhizophoraceae 195

appendages. The stamens are twice the number of the petals in the inland Rhizo-

phoraceae, and may be arranged in a single whorl around a crenulate or lobed

disc, or be attached to a calyx tube as in Pellacalyx species. The filaments are

unequal in length in Carallia brachiata and Pellacalyx species (the anti-sepalous

ones being shorter in C. brachiata and longer in P. axillaris and P. saccardianus).

Among the mangrove genera, the filaments of Rhizophora species are extremely

short, and those of Bruguiera are arranged in unequal pairs which are embraced

by the petals. The filaments of stamens in the other Malayan genera of Rhizo-

phoraceae are more or less equal in length. The anthers are generally small,

4-locular, dorsifixed and dehiscing longitudinally and introrse. Anthers of

Rhizophora deserve special mention as they are large and multiloculate, producing

clouds of pollen at anthesis.

The microspore mother cells in a developing anther are enveloped by 5 layers:

an inner tapetum, an outer epidermis and three intermediate layers. The layer

immediately underlying the epidermis produces ‘fibrous’ thickenings of the wall

which is so characteristic of the endothecial layer, and which presumably facilitates

pollen dispersal (Eames, 1961).

Following meiosis of the microsporocytes the tapetal and middle layers

disintegrate and are absorbed by the developing pollen grains so that at maturity,

only a spirally thickened endothecium and epidermis remain. In all the six species

the septum between the locules of each half-anther usually ruptures before the

pollen is released by a longitudinal line of dehiscence. The cells around the region

of dehiscence are smaller, isodiametric, and thin-walled, and the epidermal cells

are inconspicuous above this region.

According to Erdtman (1952) the pollen grains of Rhizophoraceae are 3(-4)-

colporoidate, colporate, oblate-spheroidal to subprolate, and with finely reticulate

or indistinct exine pattern. Table 5 shows the specific pollen type in the listed

species. The pollen grains of the species listed are more or less similar in

appearance. There is no pronounced difference between the pollen structure of the

species of the three genera, and pollen morphology in this case does not seem

to assist in the identification of the members of the family.

TABLE 5

POLLEN GRAINS OF THE RHIZOPHORACEAE

Species Size Shape | Exine pattern

ee ee. eee [ee ey

Anisophyllea disticha a 19.8px16.5p Subprolate | Indistinct

Combretocarpus rotundatus 24 ux 10.6 p | Subprolate | Indistinct

Carallia brachiata | 16.5pxiSp Prolate-spheroidal | Finely reticulate

Gynotroches axillaris | 10.54x86p Subprolate | Finely reticulate

Pellacalyx axillaris .. | 12px 105 p Subprolate | Indistinct

Pellacalyx saccardianus 13.24x 9.9 p | Subprolate Indistinct

Bruguiera | 2ipxi17p | Subprolate Finely reticulate

Ceriops candolleana ISpxi5.5p | Oblate-spheroidal Finely reticulate

Rhizophora mucronata |

25. x 21 p Subprolate Finely reticulate

196 Gardens’ Bulletin, Singapore — X XVII (1974)

The compound gynoecium is generally 2-12-loculate. The floral vasculature

of the plants show the presence of two series of traces, an outer series which

supplies the calyx, corolla and androecium, and an inner series which comprises

the gynoecial vasculature. Traces supplying the carpels diverge into the ovules as

well as ramify in the carpellary walls before supplying the style or styles of the

various species.

The ovary is usually inferior (Figs. 9, 11, 12, 14), rarely superior (e.g. in

Gynotroches (Fig. 13) ). Internally the gynoecium is divided by septa into as many

locules as the number of carpels. The ovules, varying from 1-2 to numerous

per locule, are pendulous and attached to an axile placenta.

The placentation in the Rhizophoraceae is generally described as axile, but

serial sections of flower buds show that the uppermost portions of the syncarpous

ovary of Combretocarpus (Fig. 12) and Gynotroches (Fig. 13) are unilocular,

indicating the parietal condition.

In Combretocarpus the carpels are fused by their ovarian regions but the

styles and stigmas are separate. Each style is grooved longitudinally on its ventral

(inner) surface, and as sections are taken down the lengths of the styles, there is

evidence of lateral concrescence of the adjacent margins of the styles. This condition

finds its equivalent in the corresponding junction of the margins of a whorl of

open conduplicate carpels. As a result, the lower part of the style and the upper

one-third of the inferior ovary are unilocular, while sections lower down show

the typical plurilocular syncarpous ovary with axile placentation.

In Gynotroches the style is simple and hollow in its lower half. This hollow

cavity leads down to the upper part of the superior ovary, and sections taken at

this point reveal a single locule deeply subdivided by projecting partitions of the

conjunct margins of each carpel, which do not meet in the centre. The lower

part of the ovary is by contrast, divided into 4-5 chambers as a result of the

union of the partitions. In the older flowers the axile condition is more pronounced

and the parietal arrangement becomes somewhat obscure.

Attempts to draw any conclusion about phylogenetic relationships of axile

and parietal placentation require great caution (Puri, 1952). The two conditions

may have arisen independently from marginal placentation or one condition may

have arisen from the other. The situation occurring in Combretocarpus and

Gynotroches, nevertheless, seems to suggest that there is possibly a general trend

from parietal to the commonly axile placentation (see Gundersen, 1939) in the

Rhizophoraceae.

Each ovule has two integuments. In Gynotroches, Carallia, and Pellacalyx,

the outer integument develops thick walls and becomes crustaceous while the

inner integument is membranous. In Anisophyllea disticha, the outer is also

composed of thick-walled cells which is lined by a thin-walled, 1-2-layered,

inner integument. In Combretocarpus rotundatus however, the outer integument

is composed of 3-4 layers of cells with less conspicuously thickened walls in

contrast to the single-layered inner integument.

In the mangrove and in some inland Rhizophoreae, a simple style is found

which is connected to a simple or lobed stigmatic crest. In Anisophyllea and

Combretocarpus however, as described before, 3-4 styles are present each of

which shows a ventral groove running from the apical stigmatic surface to the

ovary. The styles of Carallia brachiata, Gynotroches axillaris, and Pellacalyx spp.

have a stylar canal running from stigma or upper stylar region to the ovary.

Morphological Studies of Some Rhizophoraceae 197

IV. Taxonomic Considerations

The subdivision of the family Rhizophoraceae has been a subject of

controversy ever since it was established by R. Brown in 1814. Four major

systems of classification of the family have so far been proposed. They were by

Hooker (in Bentham and Hooker f., 1862-67), Schimper (1898), Ridley (1922),

and Melchior (1964).

Following Hooker’s system, the family includes 3 tribes, namely:

(1) the Rhizophoreae, members of the mangrove genera (Rhizophora,

Bruguiera, Kandelia, etc.), which exhibit inferior ovary, single style,

exalbuminous macropodous embryo, viviparous germination, and

opposite, entire, glabrous, stipulate leaves.

(2) the Legnotidae which includes Carallia, Gynotroches and Pellacalyx as

well as other members with inferior or half-inferior ovary, small embryo

embedded in fleshy albumen of the seed and opposite stipulate leaves.

(3) the Anisophylleeae, including Anisophyllea and Combretocarpus with

inferior ovary, 3-4 styles, exalbuminous macropodous embryo, and

alternate stipulate leaves.

Schimper’s proposed classification is altogether different; an outline of it

follows:

(1) Rhizophoroideae — includes genera with perigynous or epigynous

flowers, simple styles, baccate or rarely capsular fruits, opposite,

stipulate leaves.

(a) Gynotrocheae — comprising two sub-tribes.

(i) Gynotrochinae — consisting of Crossostylis, Gynotroches,

Ceriops, Kandelia and Rhizophora.

(ii) Carallinae — composed of Carallia, Pellacalyx and Bruguiera.

(b) Macarisieae — comprising Blepharistemma, Cassipourea, Dactylope-

talum, Macarisia and Weihea.

(2) Anisophylloideae — including genera with epigynous flowers, 3-4

styles, drupaceous or dry indehiscent fruits, alternate, exstipulate leaves.

Only two genera, Anisophyllea and Combretocarpus are included in

this sub-family.

Among the genera belonging to Hooker’s Legnotidae, Schimper found that

certain members, namely, Weihea, Cassipourea, Blepharistemma, Dactylopetalum

and Macarisia, are so distinctly related that he placed them in a tribe of their own,

Macariseae, to which he later assigned Anopyxis and Sterigmapetalum. The

remaining genera he placed with the mangrove species.

According to Hooker, the mangrove genera are placed in a group of their

own because of the several distinctive characters they exhibit, the majority of

which are no doubt of adaptive significance and origin. On this very point Schimper

(1898) voices his dissent, and considers it erroneous to separate a group of plants

On the strength of their response to environmental conditions. He favours a

‘natural’ classification which gives a more realistic indication of the relationships

of the plants within the family. The outcome of his proposal is the dispersion of

the mangrove genera into two different sub-tribes of Gynotrocheae, each of which

includes some of the inland genera.

198 Gardens’ Bulletin, Singapore — X XVII (1974)

Hooker’s proposal of a third tribe Anisophylleeae was accepted, but relegated

to a sub-family by Schimper.

Ridley agreed with Hooker’s arrangement of the Rhizophoraceous genera,

but raised all the three tribes to family rank. In his investigation he found that

the inland genera Carallia, Gynotroches and Pellacalyx are closely related and

warrant a group of their own. Other members of Hooker’s Legnotidae were not

investigated as his study was confined to the Malayan flora, and hence Schimper’s

suggestion of the Macarisieae as a distinct tribe was not discussed.

He also assigned Anisophyllea to a separate family Anisophylleaceae (as

Anisophylleae), and this was later taken up by Corner (1952).

In the latest (12th) edition of Engler’s Syllabus as revised by Melchior (in

Melchior 1964), the family Rhizophoraceae is divided into four co-ordinated tribes

as follows: Macarisieae, Gynotrocheae, Anisophylleeae and Rhizophoreae.

In the present investigation of hte Rhizophoraceous genera found in Johore,

Malaysia, and in Singapore, some morphological data have been assembled and

tabulated (see Table 6) in the three tribes proposed by Hooker (1898) but revised

by Melchior (1964). On the strength of the morphological features it appears that

these three tribes resemble one another very closely in many ways but yet have

their distinct differences. The mangrove tribe differs from the inland counterparts

not exclusively but manifestly in their ecological adaptive features.

TABLE 6

COMPARISON OF SOME MORPHOLOGICAL CHARACTERS OF THE 3 TRIBES

OF RHIZOPHORACEAE

Gynotrocheae [Legnotideae]

(Garallin® Gpnorocnes Anisophylleeae (Anisophyllea,

Rhizophoreae (Rhizophora,

Combretocarpus)

Bruguiera, Ceriops)

Pellacalyx)

1. Trees of littoral habitats. 1. Trees of inland forests. 1. Trees of inland forests.

2. Leaves opposite, entire, glab- | 2. Leaves opposite, entire or | 2. Leaves alternate, entire,

rous. serrate, glabrous or densely glabrous to pubescent.

puberulous.

3. Stomata anomocytic. 3. Stomata anomocytic. 3. Stomata anomocytic.

4. Stipules present. 4. Stipules present. 4. Stipules absent.

5. Phellogen subepidermal in | 5. Phellogen subepidermal in | 5. Phellogen subepidermal in

origin. origin. origin.

6. Flowers medium-sized to large | 6. Flowers small to medium- | 6. Flowers minute (0.15-0.5

(0.5—4 cm.) sized (0.25—1.5 cm.) cm.)

7. Calyx lobes persistent; 4-15 in | 7. Calyx lobes persistent; 4-8 | 7. Calyx lobes persistent; 3-4

number. in number. in number.

8. Petals entire or lobed, with or | 8. Petals clawed, fringed at the} 8. Petals laciniate or entire.

without appendages. apex in various degrees.

9. Stamens twice the number of | 9. Stamens twice the number | 9. Stamens twice the number

petals or numerous, equal or of petals, usually unequal in of petals, equal or unequal

unequal in length. length. in length.

— a

Morphological Studies of Some Rhizophoraceae 199

TABLE 6—continued

COMPARISON OF SOME MORPHOLOGICAL CHARACTERS OF THE 3 TRIBES

OF RHIZOPHORACEAE

| Gynotrocheae [Legnotideae]

(Carallia, Gynotroches

Pellacalyx)

Rhizophoreae (Rhizophora,

Anisophylleeae (Anisophyllea,

Bruguiera, Ceriops)

Combretocarpus)

10. Anthers basifixed, multi-locu- | 10. Anthers basifixed, 4-locu- | 10. Anthers basifixed 4-locu-

late or 4-locular. lar. lar.

11. Filaments shorter or longer | 11. Filaments several times | 11. Filaments several times

than anthers. longer than anthers. longer than anthers.

12. Disc present or absent. 12. Disc present. 12. Disc present.

13. Pollen grains 24 »- 26yp in | 13. Pollengrains 12-15 min! 13. Pollen grains 20 p — 24 pin

longest axis. longest axis. longest axis.

14. Carpels 1-4; ovary half-in- | 14. Carpels 3-12; ovary super-| 14. Carpels 3-5; ovary inferior,

- ferior or inferior, syncarpous. ior or inferior, syncarpous. syncarpous.

15. Ovules 2-6 per locule. 15. Ovules 2-several per | 15. Ovules 1-2 per locule.

locule, fasciculate or not.

16. Fruit a specialized structure | 16. Fruit a berry, usually glo- | 16. Fruit a drupe or dry and

with seed which germinates bose or pyriform. indehiscent with winged or

while still attached to parent woody pericarp.

plant.

17. Pericarp hard and tough. 17. Pericarp succulent. 17. Pericarp succulent or dry.

18. Seeds one per fruit. 18. Seeds numerous, embed- | 18. Seeds one per fruit.

ded in pulpy placentae in

fruit.

19. Endosperm absent. 19. Endosperm present. 19. Endosperm present.

20. Embryo macropodous; coty- | 20. Embryo minute, embedded} 20. Embryo macropodous.

ledons connate or free. in fleshy endosperm; coty- cotyledons rudimentary

ledons free. or not.

The following discussions are centrated on the three tribes as proposed by

Hooker but revised by Melchior.

(A) On the tribe Rhizophoreae

The Rhizophoraceae may be conveniently divided into mangrove and inland

groups on ecological grounds. Members of the former group belong to the tribe

Rhizophoreae, those of the latter, to the tribes Gynotrocheae (or Legnotidae) and

Anisophylleeae. In spite of Schimper’s (1898) strong objection to the adoption of

this scheme, the mangrove genera (namely, Rhizophora, Bruguiera, Ceriops and

Kandelia) in many aspects, do form a closely related unit. The development of

such characters as xerophytic leaves, prop roots, pneumatophores, in mangrove

plants are undoubtedly brought about as a result of ecological adaption. However,

there are many other features such as persistent and accrescent sepals, ex-

endospermous seed and wood anatomical characters which set this tribe apart.

200 Gardens’ Bulletin, Singapore-— X XVII (1974)

The unique method of viviparous germination of seeds in the Rhizophoreae

seems to be another adaptive feature. However, as van Steenis (in Hou, 1958)

pointed out, it is not universal or even common in other equally successful mangrove

plants such as Sonneratia (Sonneratiaceae) or Barringtonia (Barringtoniaceae), thus

the general assumption that this character is a special adaption to the environment

may be disputable.

(B) On the tribe Gynotrocheae

In the tribe Gynotrocheae (= Legnotidae, in part) only 3 genera, namely,

Carallia, Gynotroches and Pellacalyx were studied. They appear to fit into a

natural group. However, according to Marco (1935), this tribe is heterogeneous,

and therefore should be further divided into 3 groups: (1) Group I including

Carallia, Combretocarpus, Anisophyllea, Gynotroches and Crossostylis, (2) Group II

including Macarisia and four other genera which are not present in this part of

the world, (3) Group III including Pellacalyx and Poga. Unfortunately most of

this material are not available for the present study.

It is noted that Marco’s Groups I and II correspond closely to Schimper’s

tribes Gynotrocheae and Macariseae. Marco (1935) also considered Pellacalyx

to be wood anatomically unclassifiable with the others. Although this genus differs

in its tangential arrangement of 2—5 vessels in pore clusters, and in the possession

of opposite, angular pits to ray cells, yet with regard to the other wood anatomical

characters, it seems to fit in homogeneously with the rest of the Gynotrocheae.

Therefore it would appear to be unwarranted to separate Pellacalyx out solely on

these points of difference.

Anisophyllea and Combretocarpus, which constitute the Anisophylleeae, are

conspicuous by the alternate arrangement of their leaves as contrasted to the

paired condition found in the other tribes, and by their Jack of the characteristic

interpetiolar stipules of the family. The flowers of these two genera are very tiny by

comparison, their 3-4 styles are free, yet the fruits produced are relatively large

to very large (for example, 8-12 x 5-7 cm in Anisophyllea grandis Burk.).

Anatomically, Anisophyllea shows a tendency towards formation of isolateral

leaves without a hypoderm, and has less palisade tissues than the other inland

genera. Transverse sections of the young stem show the presence of a sheath of

sclerenchymatous cells surrounding the vascular cylinder, which is not found in

the other genera examined. These features are probably among the deciding

factors for the separation of the two genera, Anisophyllea and Combretocarpus,

into a tribe or even into a family as advocated by Ridley (1922) and Corner (1952).

The embryos in the seeds of the Anisophylleeae resemble those of the

Rhizophoreae in being macropodous. Other common features between these two

tribes include the deeply bi-lobed petals and the comparatively larger size of the

pollen grains. The wood anatomical characters of the Anisophylleeae on the other

hand, in general, agree very well with those of the Gynotrocheae, but appear to

be more advanced. For example, they possess shorter. broader vessel elements

than Carallia, Gynotroches and Pellacaiyx (of Gynotrocheae), and the pores are

larger, often solitary and fewer in number. Therefore, on morphological grounds

it would hardly seem justifiable to establish a separate family to accommodate

these two genera.

Morphological Studies of Some Rhizophoraceae 201

To summarize, after due consideration of the data available, it appears to

be most appropriate that the Malayan members of the family Rhizophoraceae

should be accommodated in three tribes: namely Rhizophoreae, Gynotrocheae and

Anisophylleeae, as suggested by Hooker (1865) and revised by Melchior (1964).

Acknowledgements

The senior author wishes to express her gratitude to Professor A. N. Rao,

Department of Botany, University of Singapore for his continuous interest, and

to Messrs. R. Tay, D. Teow and J. Wee for their technical assistances. Special

thanks are due to Dr. T. C. Whitmore, formerly of the Forest Research Institute,

Kepong, and to Dr. J. A. R. Anderson, former Conservator of Forests, Sarawak,

for their assistance in obtaining wood specimens.

| Bibliography

1. Arnott, H. J.

1959. Leaf clearings. Turtox News, 37: 192-194.

2. Bentham, G. & J. D. Hooker

1862-67. Genera Plantarum. Vol. 1. London.

3. Carlquist, S.

1961. Comparative Plant Anatomy. Holt, Rinehart and Winston, N.Y.

2 -Comner, -E.: J. H.

1952. Wayside trees of Malaya. 2 vols. 2nd edition. Govt. Printing Office,

Singapore.

> eames, A. J.

1961. Morphology of the Angiosperms. New York: McGraw-Hill.

6. Erdtman, G.

1952. Pollen morphology and plant taxonomy. Angiosperms. Stockholm:

Almgvist and Wiksell.

7. Foster, A. S.

1949. Practical plant anatomy. 2nd ed.; van Nostrand. New York.

8. Gundersen, A.

1939. Flower buds and phylogeny of dicotyledons. Bull. Torrey Bot.

Club 66: 287-295.

9. Hou, Ding

| 1958. Rhizophoraceae in Flora Malesiana Ser. I. 5: 429-493.

10. Johansen, D. A.

1940. Plant microtechnique. McGraw-Hill. New York.

11. Marco, H. F.

1935. Systematic anatomy of the woods of the Rhizophoraceae. Trop.

Woods. 44: 1-21.

202

13.

14.

1S;

16.

17.

18.

19:

20.

Gardens’ Bulletin, Singapore — XXVII (1974)

Melchior, H.

1964. A. Engler’s Syllabus der Pflanzenfamilien. Band II Gebrueder

Borntraeger, Berlin.

Metcalfe, C. R. and R. Chalk.

1950. Anatomy of the dicotyledons; leaves stem and wood in relation

to taxonomy, with notes on economic uses. 2 vols. Clarendon Press.

Oxford.

Payens, J. P. D. W.

1967. A monograph of the genus Barringtonia (Lecythidaceae). Blumea

15: 157-263.

Pot tye

1952. Placentation in Angiosperms. Bot. Rev. /8: 603-651.

Ridley, H. N.

1922. Flora of the Malay Peninsula. Vol. 1. L. Reeve and Co. London.

Schimper, A. F. W.

1898. In Engler & Prantl, Die Nat. Pflanzenfamilien 3, 7. 42-50. Leipzig.

Sinnott, E. W.

1914. Investigations on the phylogeny of the Angiosperms I. The anatomy

of the node as an aid in the identification of angiosperms. Amer. J. Bot.

I: 303-322.

and I. W. Bailey.

1914. Investigations on the phylogeny of the Angiosperms III. Nodal

anatomy and morphology of stipules. Amer. J. Bot. 7: 441-453.

Wylie, R. B.

1943. The role of the epidermis in foliar organisation and its relation to

minor venation. Amer. J. Bot. 30: 273-280.

Plate 1. Flowering branches. A, B, Carallia brachiata (Lour.) Merr. C, D, Pellacalyx

axillaris Korth. (Scale: A, B and D, | cm. divisions; C, 1 mm. divisions),

Plate 2. Wood anatomy. Carallia brachiata: a, TS. b, TLS. Combretocarpus rotundatus:

¢, TS. d, TLS. Gynotroches axillaris: e, TS. £, TLS.

Morphological Studies of Some Rhizophoraceae 203

Figure 1.

Anisophyllea disticha Baill.: A, pendent lateral branch bearing a pair of fruits.

B, flowering branch. C, male inflorescence. D, female inflorescence. E, female flower. F,

male flower.

204 Gardens’ Bulletin, Singapore — X XVII (1974)

Figure 2. Anisophyllea disticha Baill.: A, male flower. Al, half-flower (male), A2, stamen.

AS. Sepal. A4, petal! B, female flower. B1, half-flower (female). B2, style and stigma. C, _

mature fruit, C1-C5, longisections of various ‘stages in fruit development.

Morphological Studies of Some Rhizophoraceae

Figure 3. Carallia brachiata (Lour.) Merr.: A, flowering branch. B, flower with stamens

removed. Bl, half-flower. B2, petal. B3, stamen. B4, ovary with di

attached filaments. C, fruit, ‘ . : y with surrounding disc and

205

206 Gardens’ Bulletin, Singapore —- X XVII (1974)

Figure 4. Combretocarpus rotundatus (Miq.) Danser: A, flowering branch. B, flower bud.

Bi, sepal and antisepalous stamen. B2, flower bud with petals and one sepal removed to

show stamens, lobed disc and styles. C, mature flower, Cl, style and stigma with surrounding,

deeply lobed disc. C2, stamen. D, mature fruit. D1, younger fruit.

Morphological Studies of Some Rhizophoraceae 207

Figure 5.

Gynotroches axillaris B1.: A, flowering and fruiting branch, B, flower. Bl, half-

ower. B2, sepal. B3, petal with apical appendages. B4, stamen. C, fruit with basal persistent

calyx. Cl, cluster of fruits. C2, seed.

208 Gardens’ Bulletin, Singapore — X XVII (1974)

Figure 6. Pellacalyx saccardianus Scort.: A, flowering branch; B, flower, Bi, half-flower,

B2, petal, B3, style and lobed stigma, B4, stamen, C, fruit, Cl, longitudinal section of fruit.

P. axillaris Korth.: D, flower, D1, half-flower.

Morphological Studies of Some Rhizophoraceae 209

Figure 7. Anisophyllea disticha Baill.: a-c, progressive stages in seedling development.

Stages b and ¢ show the remains of the fruit which is embedded in the soil, Roots arise

from both ends of the fruit.

210

SA (

Figure 8. Gynotroches axillaris Bl.: a-d, stages in seedling development.

Gardens’ Bulletin, Singapore — X XVII (1974)

Morphological Studies of Some Rhizophoraceae 211

Figure 9. Anisophyllea disticha Baill.: A, B & C, different longitudinal sections of female

flower. a-j, transections at different levels as indicated in C.

212 Gardens’ Bulletin, Singapore — X XVII (1974)

Figure 10. Anisophyllea disticha Baill.: A, longisection of male flower. a-f, transverse

sections at levels indicated in A.

Morphological Studies of Some Rhizophoraceae

Oo 980A. «<o

213

Figure 11. Carallia brachiata (Lour.) Merr.: A, longitudinal section of flower, a-g, tran-

sections at levels corresponding to those shown in A

214 Gardens’ Bulletin, Singapore — X XVII (1974)

Figure 12. Combretocarpus rotundatus (Miq.) Danser: A, longisection of flower, a-h, trans-

verse sections at levels shown in

i A

Morphological Studies of Some Rhizophoraceae 215

Figure 13. Gynotroches axillaris BI.:

sections at different levels.

A, longitudinal section of flower, a-i, transverse

216 Gardens’ Bulletin, Singapore — X XVII (1974)

Figure 14. Pellacalyx saccardianus Scort.: A, longitudinal section of flower, a-g, transections

of flower at levels indicated in

Morphological Studies of Some Rhizophoraceae 217

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Figure 15. Transverse section of leaves. A, Anisophyllea disticha Baill. B, Carallia brachiata

(Lour.) Merr. C, Gynotroches axillaris Bl. D, Pellacalyx axillaris Korth. E, P. saccardianus

Scort. Mesophyll cells stippled to represent chlorenchymatous condition. Amorphous cell

| inclusions, heavy black, Mucilage-producing cells are hatched.

Gardens’ Bulletin, Singapore — X XVII (1974)

218

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Morphological Studies of Some Rhizophoraceae 219

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220 Gardens’ Bulletin, Singapore — XXVII (1974)

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Figure 19. Uniseriate and multiseriate wood rays as seen in tangential] longitudinal section.

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Danser. D, Carallia brachiata (Lour.) Merr. E, C. eugenioidea King. F, Gynotroches

axillaris B1. G, Pellacalyx axillaris Korth. H, P. saccardianus Scort.

Report of the Botanic Gardens for the year 1973

1. ADMINISTRATION

The Botanic Gardens and the Parks & Trees Branch, PWD, were integrated

to form the Parks & Recreation Division, PWD as of 1.7.73. Mr A. G. Alphonso,

acting Director of the Botanic Gardens was appointed acting Deputy Commissioner

of the Parks & Recreation Division with effect from 2.7.73. A number of posts

were redesignated as shown below.

Redesignated Post

Botanist Curator

Horticultural Assistant Assistant Curator

| Junior Horticultural Assistant Horticultural Assistant

Mr Quek Wai Yan, Assistant Curator, who was granted 3 years’ no-pay

leave to pursue a Diploma in Horticulture course at the Royal Botanic Gardens,

Kew, resumed duty on 21.6.73.

2. GARDENS’ MAINTENANCE

The Gardens’ lawns were well-maintained throughout the year. Several areas

were landscaped with flowering and foliage plants and many old plants were

replaced by more vigorous specimens.

Additional road lights and spot lights were installed in the earlier part of the

year to further improve the lighting facilities, however all spot-lights were

switched off from November, in an effort to conserve electricity during the current

energy crisis.

Over 110 offenders of the Environmental Public Health Act 1968 were sum-

moned during the year.

3. HERBARIUM

The Herbarium facilities were fully made use of throughout the year by

visiting botanists. About 202 specimens were identified for the public, and 1,878

specimens were sent out on loan to various institutions. 1,232 duplicates were

sent out in exchange for 4,310 received from other herbaria.

During the year, 7,650 specimens were mounted, 1,028 old specimens were

repaired, and 10,170 sheets were indexed. About 5,646 specimens were incorpo-

rated in the herbarium.

Several collecting trips were made in the year, details of which are given

in Appendix I.

4. ORCHIDS

Twenty-five new orchid hybrids were described in 1973. Two of these,

Dendrobium Dzemal Biiedic (D. Tumphal x D. ostrinoglossum) and Dendrobium

Rahah (D. Noor Aishah x D. Peggy Shaw) were named in honour of the Prime

221

222 Gardens’ Bulletin, Singapore — X XVII (1974)

Minister of Yugoslavia and Toh Puan Rahah, wife of the Prime Minister of

Malaysia, respectively, during their visit to the Gardens.

Three orchid hybrids were awarded prizes at the Orchid Show organised by

the Orchid Society of South East Asia. They are:

Aranda Majula Ist Prize

(Arachnis Maggie Oei x Vanda insignis)

Paphiopedilum Shireen lst Prize

(Paphiopedilum glaucophyllum

x Paph, philippinense)

Vanda Kinloch Smith 2nd Prize

(Vanda Josephine x Vanda coerulea)

During the year 106 hybrid pods were harvested from the Gardens’ nursery.

13 pods were received from local growers and 33 pods were received from the

Royal Botanic Gardens, Sydney, for germination. 99 of these 152 pods produced

viable seeds. A total of 1,141 flasks of orchid seedlings were transplanted into

pots and 2,794 seedlings were removed from the seedling houses and grown into

bigger pots.

260 plants were received from various institutions and private individuals

in exchange for 10 seed pods, 4 flasks of seedlings and 189 orchid cuttings and.

plants.

A total of 31,875 orchid sprays were supplied to various organisations,

government departments and statutory bodies.

5. NURSERIES

a. Pot Plant Nursery

For the period January to September 1973, 191 requests were received for

the loan of 7,635 potted plants for decoration purposes. Loan of potted plants

were handled by the Logistics and Maintenance section from October 1973.

Production figures of plants at the Pot Plant Nursery are shown in

Appendix I.

b. Plant Introduction Nursery

A total of 662 species of plants were introduced through collections, gifts,

exchanges, and purchases from various institutions, nurseries and private indivi-

duals. Several of the more promising species were sent to the Primary Production

Department and to the Logistics and Maintenance section for future experimenta-

tion and propagation. Altogether, 289 plants of 70 species and varieties were

distributed. Seeds of 11 species of plants were also supplied to the Logistics and

Maintenance section. The plants included new varieties of roses and pot Chrysan-

themums as well as seedlings of a fruit tree (Artocarpus odoratissima) well-known

in the Philippines and Sabah, and which should become popular in Singapore.

6. DECORATIONS AND FLORAL ARRANGEMENTS

The Gardens assisted in decorations with orchids, flowering and foliage

plants for the following: —

1. National Day Parade Float of ““The Garden City”’.

3. National Day State Banquet.

Botanic Gardens, 1973 Report 223

3. The 50th Birthday Dinner for the Prime Minister organised by the

National Trade Union Congress.

4. Meeting of the 4th ASEAN Permanent Committee on Commerce and

Industry.

5. Orchid Show organised by the Orchid Society of South East Asia.

7. TREE PLANTING DAy

Tree Planting Day is an annual national event which serves to remind the

public of the importance of growing trees.

This year’s Tree Planting Day was held on Sunday, 4th November 1973.

Tree-planting activities were organised in various constituencies. A total of 6,850

ornamental trees, 5,291 fruit trees and 8,450 shrubs and climbers were planted

this year.

8. SCHOOL OF ORNAMENTAL HORTICULTURE

There is a total enrolment of 26 students in the School. Twelve of them are

in their second year and 14 are in their first academic year.

A first year examination was conducted for the senior students in August.

The School prospectus was revised by a School Board Sub-Committee at the

end of the year.

The School Board comprises of : —

Mr Lim Leong Geok — Chairman

Mr A. G. Alphonso

Mr Kee Chin Hin

Mr Wong Yew Kwan

Mr K. C. Chung

Miss Kwok Mei Yong

Mr Hardial Singh —- Secretary

9. EDUCATIONAL ASSISTANCE AND TRAINING SCHEMES

Mr Djunaedi G. Widjaja of Indonesia, ASEAN Scholar in Botanic Studies

underwent a two-month practical training course in the Gardens.

Four students from the University of Singapore and Nanyang University

were given six weeks’ practical experience in the Gardens as part of the Industrial

and Business Orientation programme organised by the Science Council of Singa-

pore.

Miss Artis Djusar of Indonesia was awarded a Colombo Plan Scholarship

to pursue a six-month course in orchid culture in the Gardens.

Two students participating in the Youth Week in Commerce and Industry

programme were attached to the Gardens for two days.

10. LIBRARY AND PUBLICATIONS

The Library, which maintains a valuable collection of botanical and horti-

cultural publications is referred to by staff members, botanists and research

workers preparing taxonomic revisions and floristic studies of this region. In-

terested members of the public may obtain permission from the Deputy Com-

missioner of Parks and Recreation to make use of the Library during normal

office hours.

224 Gardens’ Bulletin, Singapore — X XVII (1974)

At the close of the vear, the accessioned book collection totalled 9,878. There

were 384 current titles of serial publications obtained from subscriptions and

exchanges. Gifts received during the year numbered 8 periodical titles, 76 reprints

and 18 books.

Volume 26, part II of the Gardens’ Bulletin was published on 15 September

1973. There were 178 exchanges for the Gardens’ Bulletin and the Botanic

Gardens’ Annual Report 1972.

11. SALE OF PUBLICATIONS AND PLANTS

Revenue collected from the sale of botanical and horticultural publications

amounted to $2,039.20.

1,401 orchid seedlings and 1,257 orchid plants and cuttings were sold for a

total sum of $7,874.00.

12. VISITORS

HE Mr Dzemal Bijedic, Prime Minister of Yugoslavia and Madame Razija

Bijedic visited the Gardens on 18.3.73. An orchid, Dendrobium Dzemal Bijedic

was named in honour of the Prime Minister.

Mr R. J. Tizard, Minister of State for Health Services, New Zealand,

presented and planted a Pohutukawa tree in the Gardens. He was accompanied

by Mrs Tizard.

Toh Puan Rahah, wife of the Prime Minister of Malaysia was welcomed at

the Gardens on 15.11.73. She chose an orchid, Dendrobium Rahah, to be named

after her.

Other eminent visitors included: —

Mr George Avery, Director Emeritus, Brooklyn Botanic Gardens and Mrs

Avery.

Mr J. J. Aves, Director of Parks, Pretoria, South Africa.

Mr Ian D. Galloway, Director of Parks, Wellington, New Zealand.

Professor H. Kamemoto, University of Hawaii, Honolulu, U.S.A.

Professor Charles H. Lamoureux, University of Hawaii, Honolulu, U.S.A.

13. NATURE RESERVES

The present Nature Reserves constitute the Bukit Timah Reserve, the

Catchment Area, and Labrador Reserve. Kranji Reserve was deleted late in the

year.

The Bukit Timah Nature Reserve comes under the supervision of the

Botanic Gardens. The Labour force concentrated on maintenance and repair

work during the year. Over 400 steps along the footpaths were repaired and a fence

was installed along a precipitous boundary aligning one granite quarry. Danger

signs were posted to alert the public of dynamite blasting at specified times by

the quarries. A new set of benches and 18 litter bins were installed. The Reserve

is closed to public vehicles at the following times :—

7 a.m. — 8 p.m. on Sundays and Public Holidays.

4 pm. — 8 p.m. on weekdays.

2 p.m. — 8 p.m. on Saturdays.

Botanic Gardens, 1973 Report 225

Earlier in the year, paths and bridges in Kranji Reserve were repaired when

they were damaged during high tides.

A new Board of Trustees for the Nature Reserves was set up under the

Nature Reserves (Amendment) Act of 1973.

The Board members are: —

Mr Quek Kiah Huat (Chairman),

Mr Lim Leong Geok.

Mr A. G. Alphonso.

Dr Hsuan Keng.

Mr Chan Chor Cheung.

Mr Sie Chu Hua.

Mr Wong Yew Kwan.

Mr Kee Chin Hin.

Mr Christopher Giles Roche.

Dr Tay Kah Seng.

Dr Chang Kiaw Lan was nominated Honorary Secretary by the Board.

14. METEOROLOGICAL RECORDS

Temperature, rainfall and humidity records for the years 1969-1973 are

given in Appendix III. The figures were recorded at the Botanic Gardens and

at the Bukit Timah Nature Reserve.

226 Gardens’ Bulletin, Singapore — X XVII (1974)

PRODUCTION FIGURES OF POTTED PLANTS AT

THE POT PLANT NURSERY FOR 1973

Month

January

February

March

April

May

June

July

August

September

October

November

December

Total

Botanic

Gardens

Nursery

1,090

895

965

785

885

1,060

1,075

2,230

2,836

1,400

1,250

520

Ulu Pandan

Nursery

1,295

850

1,015

1,032

508

1,590

1,166

1,170

1,913

880

740

887

APPENDIX I

Total

2,385

1,745

1,980

1,817

1,393

2,650

2,241

3,400

4,749

2,280

1,990

1,409

28,037

227

Botanic Gardens, 1973 Report

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Gardens’ Bulletin, Singapore — X XVII (1974)

242

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Botanic Gardens, 1973 Report

January

February

March

April

May

June

July

August

September

October

November

December

January

February

March

April

| May

June

July

August

September

October

November

December

1969

METEOROLOGICAL REPORT

KRANJI WAR GRAVES CEMETERY

Recorded daily at 8.00 a.m. from

1-1-69 to 31-12-69

Month

Total

1970

Total

Rainfall

inches

135

935

6.95

7.65

9.47

6.33

4.37

4.25

6.45

8.10

5.40

26.35

102.02

METEOROLOGICAL REPORT

KRANJI WAR GRAVES CEMETERY

Recorded daily at 8.00 a.m. from

Month

1-1-70 to 31-12-70

Total

Rainfall

inches

5.58

4.93

11.20

6.65

11.48

10.78

10.40

4.28

4.25

8.45

13.47

11.83

104.70

243

APPENDIX IIIp

with rain

107

No. of

days

with rain

Highest fall

in 24 hours

inches

Highest fall

in 24 hours

inches

SSL moou

SRSaRaESES

244 Gardens’ Bulletin, Singapore — XXVII (1974)

APPENDIX IIIq

1971

METEOROLOGICAL REPORT

KRANJI WAR GRAVES CEMETERY

Recorded daily at 8.00 a.m. from

1-1-71 to 31-12-71

No. of

Total Highest fall

Month . days :

Rainfall with cain in 24 hours

inches inches

January Ke Ae. se mas 3.55 a 0.75

February - ve # ss 5.49 6 1.95

March i, & Ly me 3.73 6 1.65

April i. ie Es oe 0.48 3 0.18

May oh oe - _ 2.47 d 1.50

June # is iS S. 6.65 10 1.78

July Bs m es vis 7.48 7 3.00

August as - as 53 10.10 18 1.94

September 2 a% se, om 5.06 9 1.15

October x J Ri: 2 8.25 i7 1.69

November a La a os 8.65 16 1.43

December 4s i, os os 7.26 14 3.65

TotalyT.. 69.17 120 20.67

1972

METEOROLOGICAL REPORT

KRANJI WAR GRAVES CEMETERY

Recorded daily at 8.00 a.m. from

1-1-72 to 31-12-72

No. of ;

Total ; Highest fall

Month : days ;

Rainfall wath sain in 24 hours

inches inches

January bi. sf b dfs 1.40 4 0.81

February be i Ce rs 8.93 12 2.30

March M, it By of 2.33 8 0.66

April i. ig ms Me 9.73 16 2.08

May ad ee £ a 4.50 10 1.29

June x 4 es cs ue 4.19 8 1.07

July be rs Be oh 0.94 3 0.68

August i a a me S22 13 0.80

September i Ee ee 9:73 13 2.18

October A A, e ah 0.83 i? 1.16

November ft Pe at i 8.19 15 42

December As a * es 8.65 19 2.01

Totab?T.. 70.64 133 | 16.76

Botanic Gardens, 1973 Report 245

APPENDIX Ilr

1973

METEOROLOGICAL REPORT

KRANJI WAR GRAVES CEMETERY

Recorded daily at 8.00 a.m. from

1-1-73 to 31-12-73

No. of ;

Total Highest fall

Month . days :

Rainfall En | 24 hours

inches inches

January 7 a z i 7.91 13 1.91

February ae iF za ee 8.02 16 1.56

March .... ae - = ss 8.98 9 1.352

April ~ os as oe LET? 16 2.33

May S =f Ms - 8.16 15 2.30

June ie a ek a 5.20 1] Lk

July 5 * a: rate 5.87 9 2.60

August = ast ME Pe 10.01 17 2.66

September . ae me oe 9.26 17 2.30

October - os . sa 9.81 ti 2.90

November io ms te a 8.43 15 2.50

December a aa i s 6.23 14 2.40

Total, .. 99.65 163 26.29

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Annotated list of seed plants of Singapore (III)*

HSUAN KENG

Department of Botany

University of Singapore

II. Angiosperms-Dicotyledons (continued)

19. CAPPARIDACEAE

Key to the genera

Leaves simple; scandent shrub; fruit a many-seeded berry Capparis

Leaves palmately divided or compound.

B. Herbs; fruit cylindric, dry Cleome

B. Erect trees; fruit globose, fleshy Crataeva

Capparis micracantha DC.

Shrub, scandent; consisting of two forms: the wild form, ssp. korthalsiana

(Miq.) Jacobs, with 60-100 stamens per flower was once collected from Changi

(Ridley 4418) and Bukit Timah, called C. finlaysoniana Wall. in Ridley’s

Flora; the cultivated form, ssp. micracantha, with 20-35 stamens per flower,

is occasionally planted in gardens. Vern. Jambol merah.

Cleome aculeata Linn.

cl.

el.

Cl.

Herb, often with stipular thorns; flowers white to cream; stamens 6; Ovary on

a short stalk; weed, of tropical American origin.

gynandra Linn.

Tall herb, often cultivated in gardens; flowers white; stamens and ovary both

elevated by a long stalk (androgynophore); also called Gynandropsis

pentaphylla DC. Native of Tropical America.

rutidosperma DC.

Herb; flowers violet-blue to pink; stamens 6; seeds with an open cleft;

weed, of African origin.

speciosa Rafin.

Tall herb, often growing in gardens as an ornament; flowers pink; stamens

and ovary both elevated by a long stalk; native of trop. America. PRIE,E «

viscosa Linn.

Herb; flowers bright yellow; stamens 10-20; ovary sessile; weed. pantropical.

Crataeva religiosa Forst. f.

* continued from Gdns’ Bull. Sing. 27:83.

Small tree, occasionally planted in gardens; leaves 3-foliolate; flowers creamy

yellow; native of Malaya and Tropical Asia. Vern. Cadat. fA °

247

248 Gardens’ Bulletin, Singapore — XXVIII (1974)

20. CRUCIFERAE

Key to the genera

A. Flowers white or purplish.

B. Erect herbs; petals 1.5-2 cm long Raphanus

B. Creeping or floating herbs; petals less than 7 mm long Nasturtium

A. Flowers brightly yellow.

C. Garden vegetables; petals over 0.5 cm long; disc-glands 4 Brassica

C. Weeds; petals less than 0.5 cm long; disc-glands 6 Rorippa

Brassica* alboglabra Bailey

Kai Lan or Chinese kale; native of S. China; growing locally from the

Bras. chinensis Jusl.

Pak Choy or Chinese white cabbage; prob. native of S. China; one of the

commonest vegetables in markets; growing locally and also imported; many

varieties, including; Pak Choy (var. chinensis 3 ), Choy Sam (var. para-

chinensis 32,{) ), etc.

Bras. juncea Czern. & Coss.

Kai Choy or mustard cabbages; native of subtropical Asia; mostly imported;

several varieties, including the crisp leaved (var. crispa) and the broad-leaved

(var. rugosa) and others. 7 3é «

Bras. oleracea L.

The cabbages; native of S. Europe; almost all imported; several varieties,

including cabbages (var. capitata®[[3é » @t)3Z), cauliflower (var. botrytis

ZERRSZ ), kohl-rabi (var. gongylodes 7¢ BFR ) etc.

Bras. pekinensis Rupr.

The celery cabbage or the pale and soft-leaved cabbage; native of China;

mostly imported; common varieties including the compact-headed Shangtung

cabbage (var. cylindrica #{#$ 432), the loose headed celery cabbage (var. laxa

eA ), ete.

Nasturtium officinale R. Br.

Water cress, a native of Europe; aquatic herb, creeping or floating, growing

Raphanus sativus L.

Radish, prob. a native of temperate Asia or the near East; annual or perennial

herb, the fleshy tap root is a common vegetable, mostly imported. #27] -

Rorippa indica Hiern

Annual weed, occasionally found in waste land; flowers very small, bright

yellow; formerly called Nasturtium indicum DC.

*For the nomenclature of the Brassica species, G.A.C. Herklot’s Vegetables in South-East

Asia (1972) is followed.

Annotated list of seed plants of Singapore (111) 249

21. MORINGACEAE

Moringa pterygosperma Gaertn.

Horse radish tree, native of India; small tree; leaves 3 or 4 times pinnately

compound; flowers pale yellow, in large panicles; fruit a long cylindric

capsule often cultivated near villages for its edible leaves and fruits. Vern.

Lembugai, #87 -

22. CRASSULACEAE

Kalanchoe laciniata (L.) DC.

Succulent herb; flowers yellow or orange; cultivated, native of continental

Asia.

Kalan. pinnata (Lamk.) Pers.

Flowers purple; detached leaves producing young plants from leaf margins

or midrib: also called Bryophyllum calycinum Salisb.; native of Africa.

23. SAXIFRAGACEAE

Key to the genera

A. Herbs; leaves rounded Saxifraga

A. Shrubs or trees; leaves ovate or oblong.

B. Flowers 4-merous, functional, in terminal racemes Polyosma

B. Flowers 5-merous, all sterile, in dense terminal corymbs Hydrangea

Hydrangea macrophylla (Thunb.) Ser. ex DC.

Shrub or subshrub; flowers all sterile, in large corymbs, pink or blue; garden

Polyosma conocarpa Ridl.

Small tree; petals linear, white; specimens not available.

Poly. fragrans Benn.

Small tree; once collected by Wallich (No. 8472), Not Seen.

Poly. ridleyi King.

Tree; flowers white; recorded from Seletar.

Saxifraga stolonifera Meerb.

Small herb, with long thin runners; leaves rounded, lobed, hairy; occasionally

cultivated in pots, never flowering; native to China, often called Saxif.

24. PITTOSPORACEAE

Pittosporum ferrugineum Ait

Shrub or small tree; leaves crowded at twig tips, in false whorls; flowers

yellowish white; capsule orange, 2-valved; Kranji, (Mat 5972) Changi, Seletar,

often along sea-shores. Vern. Giramong.

250 Gardens’ Bulletin, Singapore — X XVII (1974)

25. DICHAPETALACEAE

Dichapetalum sordidum (Ridl.) Leenh.

Shrub or small tree; infl. axillary; petals oblong, yellow, bifid at apex; Bajau,

(Mat 6752). Called Chailletia sordida Ridl. in Ridley’s Flora.

26. ROSACEAE

Key to the genera

A. Herbs or shrubs, creeping, scandent or climbing.

B. Tiny herb with creeping stolons; leaves 3-foliate Duchesnea

B. Shrubby with prickly stems.

C. Leaves pinnate; carpels on a concave receptacle Rosa

C. Leaves simple, entire or shallowly lobed; carpels on a convex

receptacle Rubus

A. Trees or shrubs, erect.

D. Style terminal; carpel 1, 2-ovulate Prunus

D. Style basal.

E. Ovary 1-loculate.

F. Stamens 5-10 Licaria

F. Stamens 2 Parastemon

E. Ovary 2-loculate Parinari

Duchesnea indica Focke

Tiny herb with slender stolons; aggregate fruits red, spheroid, a miniature of

the strawberry; also called Fragaria indica Andr. A weed, occasionally found

in shade and wet places.

Licaria splendens (Korth.) Prance

Small tree; leaves alternate; flowers small, white; fruit a small red drupe;

common in secondary forests in Water Catchment area (Cantley 2592). Called

Coccomelia nitida Ridl. and Angelesia splendens Korth. in Malayan literature.

Parastemon urophyllus A. DC.

Large tree; flowers very small, in racemes; fruit oblong, yellowish or pink;

in open places often near the sea; formerly found in Jurong (Ridley 6066),

Changi, Chua Chu Kang.

Parinari corymbosa (BI.) Mia.

Large tree; Changi (Ridley 4792) and Fort Canning. Called Parinarium

griffithianum Benth. in Ridley’s Flora, and Maranthes corymbosa Bl. in

Whitmore’s Tree Flora.*

Pari. excelsa (Jack) Kosterm.

Large tree; flowers white, in racemes or panicles; fruit rounded, slightly

compressed; formerly found in Chua Chu Kang. Called Parinarium asperulum

Mig. in Ridley’s Flora, and Atuna excelsa Kosterm. in Whitmore’s Tree

Flora.

*Tree Flora of Malaya 2 (1973)

Annotated list of seed plants of Singapore (111) 251

Pari. oblongifolia Hook. f.

Large tree; leaves glaucous beneath; fruit ellipsoid (6 x 3 cm); in dense

forests, Bukit Timah and Mandai (Kiah s.n. in 1940); one of the valuable

timber trees in Malaya. Vern. Balau.

Pari. sumatranum (Jack) Benth.

Large tree, buttressed; flowers white; fruit ellipsoid, hard, brown with white

dots; in forests, Chua Chu Kang (Ridley 3901), Seletar. Called Parinarium

costatum Bl. in Ridley’s Flora.

Prunus arborea (Bl.) Kalkman

Large or small tree; in forests, Kranji, Tanglin, Cluny Road (Ridley 4452):

called Pygeum parvifiorum Hook. f. and Pyg. persimile Kurz in Ridley’s Flora.

Prun. grisea (C. Muell.) Kalkm. var. tomentosa Kalkm.

Shrub; Seletar (Sinclair 40272). Called Pygeum lanceolata Hoff. in Ridley’s

Flora.

_ Prun. polystachya (Hook. f.) Kalkman

: Tree; leaf-blades with 2 conspicuous glands at the base; fruit rounded, 2-lobed:

common in forests, Gardens jungle, Catchment forests, etc.

Rosa chinensis Jacq.

Native of China, called R. indica L. in Burkill’s Dictionary; including: var.

semperflorens or the crimson China rose, var. minima the dwarf fairy rose, etc.

i Several other garden species and hybrids are introduced and cultivated. A #7E-

Rubus glomeratus BI.

Climbing shrub; leaves ovate deltoid, obscurely 3-lobed; petals white; fruit

orange, of few drupes: Jurong (Ridley s.n. in 1880). Vern. Akar balan adap.

Rubus moluccanus L.

Shrub, armed with small hooked prickles; leaves ovate or rounded, cordate.

3-8 shallowly lobed; fruit red, of many small drupes; Bukit Timah (Sinclair

37800), Tanglin, Jurong. Called R. angulosus Focke in Ridley’s Flora. Vern.

Tempu rengat.

27. CONNARACEAE

Key to the genera

A. Carpels 4 to 5 in a flower, usually more than 1 carpel developing into

fruit (except Rourea); calyx usually enlarged in fruiting stage.

(except Cnestis).

B. Leaves 3-foliolate; fruit warty Agelaea

B. Leaves pinnately compound; fruit glabrous Cnestis, Rourea

A. Carpeil 1 per flower; calyx not enlarged.

C. Leaves 1-foliolate; flowers small, in axillary clusters Ellipanthus

C. Leaves mostly pinnate; flowers in large terminal panicles Connarus

252 Gardens’ Bulletin, Singapore — X XVII (1974)

Agelaea borneensis Merr.

Climber; leaves 3-foliolate, leaflets clliptic to oblong; flowers small, pinkish

red; fruit papillose; common in forests; Tanglin, Sembawang, Bukit Timah

(Ridley 5585), Changi; called Agel. vestita in Ridley’s Flora. Vern. Akar

kachang kachang.

Agel. macrophylla (Zoll.) Leenh.

Lateral leaflets + symmetric: Changi (Hullett 426); called Agel. hullettii in

Ridley’s Flora.

Agel. trinervis Merr.

Lateral leaflets very oblique; fruit warty; Changi (Ridley s.n. in 1893); called

Agel. wallichii in Ridley’s Flora. Vern. Akar kachang jantan.

Cnestis platantha Griff.

Climber, sometimes a shrub; leaves odd-pinnate, leaflets 21-31; flowers white;

fruit 1 or 2 per flower, pear-shaped; common in dry woods and open places;

Bukit Timah (Ridley s.n. in 1894), Tanglin, Changi.

Connarus ferrugineus Jack

Climber, sometimes a shrub; leaves odd-pinnate, leaflets 7-11; branches

densely ferruginous-tomentose: fruit ellipsoid; Bukit Timah (Md. Shah 744),

Bukit Mandai.

Conn. grandis Jack

Leaflets 3-5, lanceolate to oblong-ovate; fruit obovoid; Changi, Nee Soon

(Sinclair 40321).

Conn. monocarpus L. ssp. malayensis Leenh.

Leaflets 5-9; fruit obliquely spindle-shaped; formerly found at Bukit Mandai,

Changi and Chua Chu Kang (Ridley s.n. in 1894); called Conn. oliogophyllus

in Ridley’s Flora.

Conn. semidecandrus Jack

Leaflets 3-7, elliptic to lanceolate; fruit pear-shaped, compressed; Bukit

Mandai, Cluny Road, Seletar (Mohd. Noor 26).

Ellipanthus tomentosus Kurz.

Large tree; leaves 1-foliolate; flowers small, white; Bukit Timah (Corner

34639), Gardens jungle; called Ellip. griffithii in Ridley’s Flora.

Rourea fulgens Planch.

Large climber or shrub; leaves odd-pinnate; leaflets 13-51, very oblique at

base; young leaves pink; flowers white or pink; fruit ovoid to ellipsoid,

curved; formerly collected at Kranji (Goodenough 2027), and at Gardens

jungle.

Annotated list of seed plants oj Singapore (III) 253

Rour. minor (Gaertn.) Leenh.

Leaflets 1-19; Bukit Timah, Changi (Ridley 3981); called Rour. acuminata in

Ridley’s Flora.

Rour. mimosoides Planch.

Leaflets 5-51, the base slightly oblique; fruit narrowly ellipsoid. curved;

Changi (Goodenough 2020) and also very common in reservoir woods; Vern.

Serilat.

Rour. rugosa Planch.

Leaflets 15-41; fruit oblong-ovoid, curved; Changi, Seletar (Hullett 594); Vern.

Semilat puteh.

28. LEGUMINOSAE

Key to the subfamilies.

A. Flowers regular; calyx and corolla mostly valvate in bud I. Mimosoideae

A. Flowers irregular; perianth segments conspicuously imbricate

in bud

B. Corolla caesalpinaceous, namely petals 5, free, with the

uppermost (vosterior) petal innermost II. Caesalpinoideae

B. Corolla papilionaceous, namely petals 5, with the uppermost

petal outermost, and the two lower (anterior) petals often

basally connate III. Papilionoideae

(I} Mimosoideae — Conspectus of tribes*

A. Calyx-lobes valvate.

B. Stamens more than 10.

1. Ingeae (Filaments united into a tube): Serianthes, Samanea, Pithecellobium,

Albizia, Calliandra.

2. Acacieae (Stamens free or nearly so): Acacia.

—————————— oo rLhrhhrCOCUC rT TTT”

B. Stamens as many or twice as many as petals.

3. Mimoseae (Anthers glandless): Leucaena, Mimosa.

4. Adenanthereae (Anthers crowned by a gland; seeds albuminous): Neptunia,

Adenanthera.

5. Piptadenieae (Anthers usually glandulate; seeds exalbuminous): Entada,

2 ea

A. Calyx-lobes imbricate.

6. Parkieae: Parkia.

———

(II) Caesalpinoideae — Conspectus of tribes

A. Leaves pinnate, or sometimes simple or 1-foliolate.

B. Sepals free to the base.

1. Cynometreae (Anthers dorsifixed, dehiscing by slits; leaves simple pinnate or

ples bracteoles small or large, not enclosing the flower buds): Cynometra,

Sindora.

2. Ambherstieae (Anthers and leaves as above; bracteoles well developed, enclosing

the flower buds, valvate, persistent): Saraca, Trachylobium, Hymenaea, Tama-

rindus, Intsia, Amherstia, Brownea.

3. Cassieae (Anthers firm in texture, usu. dehiscing by pores; leaves usu. simple

pinnate): Dialium, Koompassia, Cassia.

*Adapted and modified from J. C. Willis’ Dictionary (7th ed. 1972).

254 Gardens’ Bulletin, Singapore — XXVII (1974)

B. Sepals joined below.

4. Cercideae (Leaves usu. simple, bilobed; stamens 10 or fewer): Bauhinia.

A. Leaves bipinnate.

5. Caesalpinieae (Flowers in racemes or panicles): Caesalpinia, Peltophorum, Delonix.

(il) Papilionoideae — Conspectus of tribes

A. Stamens free or almost so.

1. Sophorieae (Trees, shrubs or rarely woody herbs or lianes; leaves pinnate or

1-foliolate with a joint between petiole and lamina): Ormosia, Sophora.

A. Stamens mon- or di- adelphous (i.e. filaments joined into one or two groups).

B. Herbs or shrubs (or trees or lianes in tribes 3 & 7); pods dehiscent unless short

and 1-2 seeded, or inflated.

C. Pods not transversely jointed.

2. Genisteae (Usually shrub; leaves simple or digitately 3- or more foliolate, leaflets

entire; stamens usually monadelphous, anthers often of 2 sizes): Crotalaria.

3. Astragaleae (Galegeae) (Leaves pinnately S-many or rarely 3- or 1-foliolate;

leaflets usually entire; rachis not ending in a tendril; stamens usually diadelphous):

Indigofera, Psoralea, Tephrosia, Milletia, Giliricidia, Sesbania.

4. Fabeae (Vicieae) (Herbs, leaves even-pinnate, without stipels, rachis ending in a

point or tendril; stamens 10, diadelphous): Vicia, Pisum.

5. Abreae (Shrubs or twiners woody at the base; leaves even-pinnate, rachis ending

in a point, usually stipellate; stamens 9, united): Abrus.

6. Phaseoleae (Often twining; leaves pinnately 3-foliolate, usually stipellate, rarely

1 or 5-7 foliolate; leaflets entire or lobed; stamens di- or monadelphous):

Clitoria, Centrosema, Glycine, Erythrina, Mucuna, Canavalia, Cajanus, Maughania,

Phaseolus, Vigna, Pachyrrhizus, Dolichos, Psophocarpous.

C. Pods transversely jointed.

7. Coronilleae (Hedysareae) (Shrubs or twiners; leaves pinnately 3- to many-foliolate,

stipellate or not): Aeschynomene, Arachis, Zornia, Desmodium, Alysicarpus;

Uraria.

B. Trees, shrubs or lianes; pods indehiscent.

8. Dalbergieae (Leaves pinnately 5-many-foliolate, rarely 1—3-foliolate, stipellate or

not): Dalbergia, Pterocarpus, Pongamia, Derris, Andira.

Abrus precatorius L.

Twining subshrubs; leaves even-pinnate, leaflets 8-17 pairs; seeds hard,

scarlet with a black spot; in hedges and on sea-shores (Changi. Ridley s.n. in

1891). Probably of African origin, naturalized. Vern. Akar saga, #43

Acacia auriculiformis A, Cunn. ex Benth.

Tree; phyllodes (seemingly leaves) dull green; flowers small in yellow clusters;

pods curling up into a ring when ripe, dark brown. Native of New Guinea

and Australia (or Thursday Island in the Torrey Strait); extremely common

all over the island.

Acac. farnesiana Willd.

Shrub or small tree, thorny; leaves bipinnate; flower heads bright yellow,

fragrant. Probably of American origin. Vern. Lasana, 8K °

Acac. cincinnata F. Muell.

Shrub or small tree; phyllodes silvery-coloured (‘Silver Wattle’); native of

tropical Queensland. Sometimes called Acac. mangium Willd. (from Moluccas)

which may prove to be conspecific with this species.

Annotated list of seed plants of Singapore (IIT) 255

Acac. pseudo-intsia Miq.

Big climber; leaves bipinnate; pinnae 4-10 pairs; leaflets 10-30 pairs on each

side stalk; in forests, Bukit Mandai, Changi (Hullett 458), Tuas, Chua Chu

Kang. Vern. Akar kapok.

Adenanthera bicolor Moon

Tree; pods curved in a ring; seeds one third black and two thirds scarlet;

in lowland forests, Bukit Mandai (Ridley 3636a), Pasir Panjang, etc.

Ad. pavonina L.

Large tree: leaves bipinnate; leaflets 9-15 on each side stalk; pod curved:

Aeschynomene indica L.

Herb; leaves pinnate, with 20-30 pairs of leaflets; flowers yellow; pods jointed,

breaking into 8-10 prickly parts; in open wet places; Galang (Ridley 10355).

Albizzia falcataria (L.) Fosberg

Tall tree, very fast-growing; native of Moluccas, planted and wild in Singapore,

common in waste places. Also called A. falcata Back. or A. moluccana Miq.

Alb. retusa Benth.

Tree; leaves bipinnate; pinnae |I-4 pairs: leaflets 3-10 pairs on each side

stalk: flower heads pink; rare, once collected at Pulau Ubin (Ridley 4752).

Called A. littoralis T. & B. in Ridley’s Flora.

Alb. pedicellata Baker

Tall tree; leaves bipinnate; leaflets 12-16 pairs on each side stalk; pods flat

(30-45 cm jong) and thin; in forests, Kranji (Ridley 6297).

Alysicarpus vaginalis DC.

Ascending herb; uni-foliolate, variable; flowers white or dark red; pods

cylindric, jointed, of 4-9 1-seeded pieces: in open places, Changi (Ridley

2080a), Telok Kurau.

Ambherstia nobilis Wall.

Small tree; leaves pinnate; leaflets 4-7 pairs, opposite; flowers in hanging

racemes from branches, Native of Burma, a well-known garden ornamental.

Andira inermis (Wright) H.B.K.

Tree; leaves odd-pinnate, with 4-8 pairs of leaflets; flowers dark purple:

pods pear-shaped, indehiscent. Native of Surinam in trop. America.

Arachis hypogaea L.

Ascending herb, rooting at the base; leaves even-pinnate, with 2 pairs of

leaflets; flowers bright yellow; after pollination, the ovary on a lengthening

stalk is forced into the ground, where the pod develops “‘ground-nut”. Native

256 Gardens’ Bulletin, Singapore — XXVIII (1974)

Bauhinia acuminata L.

- Shrub; leaves simple, 2-lobed, the lobes pointed; flowers white; stamens 10.

Native of S.E. Asia, cultivated.

Bauh. flammifera Ridl.

Large climber; flowers in large panicles, yellow, orange then red. Called

Phanera flammifera (Ridl.) de Wit. Native of Malaya.

Bauh. griffithiana Prain

Large climber; petals white; stamens 6; in thickets. Also called Phanera

griffithiana (Prain) de Wit.

Bauh. monandra Kurz

Tree; petals yellow then pale red; perfect stamen 1. Native of America,

cultivated.

Bauh. purpurea L.

Bush or small tree; flowers pink: stamens 3 with pink filaments, Continental

Asia, cultivated.

Bauh. semibifida Roxb.

Large climber; flowers white then greenish yellow; stamens 3, white; in

secondary jungle, Bukit Mandai, Bukit Timah (Burkill 323), Gardens jungle.

Also called Phanera semibifida (Roxb.) Benth.

Bauh. tomentosa L.

Shrub or tree; flowers solitary or in pairs; petals pale yellow; stamens 10.

Trop. Africa and continental Asia, cultivated.

Bauh. variegata BI.

Bush or small tree; leaves simple, 2-lobed; flowers in short racemes, pink to

violet; perfect stamens 5. Native of continental Asia, cultivated. A natural

hybrid between this species and Bauh. purpurea is called Bauh. blackeana

Dunn which also has 5 stamens but remains sterile, commonly cultivated in

gardens.

Brownea ariza Benth. |

Shrub or small tree, branches drooping; new leaves pink, developing in tassels;

flowers red, in a crowded head. Native of Colombia, occasionally cultivated

in gardens.

Caesalpinia bonduc Roxb. emend. Dandy & Exell

Prickly shrub; leaves bipinnate; pinnae 6-10 pairs; leaflets 12-24; stipules

pinnate; petals yellow; near the sea.

Caes. crista L.

Prickly black-stemmed climber; leaves bipinnate; flowers bright yellow; on

sandy beaches and mangroves, Kranji, Jurong. Called C. nuga in Ridley’s

Flora.

Caes. globulorum Bakh. f. & Van Royen

Leaves and leaflets larger than those of Caes. honduc; stipules subulate; near

the sea.

Annotated list of seed plants of Singapore (IIT) 257

Caes. pulcherrima (L.) Swartz

Shrub; flowers yellow or red; known as Peacock flower, native of S. America.

Also called Poinciana puicherrima, <3 JAUZE -

Caes. sappan L.

Prickly shrub or tree; flowers bright yellow. Native of Continental Asia,

occasionally cultivated. Vern. Sappan tree, fi HK »

Cajanus cajan (L.) Huth.

Shrub, white hairy; leaves 3-foliolate. Vern. Kachang kayu, pigeon pea, AE. «

Calliandra brevipes Benth.

Erect shrub; leaves bipinnate; pinnae in one pair; leaflets 20-40 pairs; free

part of the filaments pinkish above and white below. Native of Tropical

America.

Call. haematocephala Benth.

Like the above species, but leaflets 6-10 pairs, and free part of the filaments

entirely blood red. Native of Trop. America.

Canavalia catharica Thou.

Twiner; leaves 3-foliolate; flowers rose-pink to dark purple; pod linear

oblong (7-12.5 x 4-6 cm), strongly 3-keeled; beans poisonous; common on

sandy shore. Also called C. turgida, C. obtusifolia.

Cana. ensiformis DC.

Jack bean; native of the New World tropics; pod 20-30. x 2-3 cm. Vern.

Kachang parang puteh.

Cana. gladiata DC.

Sword bean; native of the old world tropics; pod 30-60 x 3-4.5 cm, edible.

Vern. Kachang parang.

_ Cana. maritima Piper

Twiner; flowers pink, sweet scented; pod 6-15 x 1.5-3; beans edible; common

| on seashores: Changi. Also called C. turgida Grah. ex Gray.

Cassia alata L.

Shrub; leaves simple pinnate; leaflets 8-20 pairs; flowers orange, in racemes,

at first covered with large dark orange bracts; pod black, with 2 broad wings

along its length: native of tropical America. Vern. Gelenggang. In addition

to those listed below, there are a number of others species planted. All the

Cassia species in Singapore are introduced; 2 of them, C. nodusa and

C. siamea are native to the Malayan mountains, the rest are from various

parts of the World. ac |

Cas. fistula L.

Tree; leaflets 3-8 pairs; hanging racemes with yellow flowers (hence ‘“‘Golden

shower’); pod woody, cylindric. Native of India and Ceylon.

Cas. fruticosa Mill.

Shrub; leaflets 2-pairs; flowers pale yellow, in loose terminal clusters. Native

of Tropical America.

258 Gardens’ Bulletin, Singapore — X XVI] (1974)

Cas. hirsuta L.

Hairy herb; leaflets 3-6 pairs; flowers 1-3, axillary, yellow; pod linear,

rounded. Tropical America.

Cus. mimosoides L.

Subshrub; flowers 1-3, axillary. A weed, native of S.E. Asia.

Cas. nodusa Buch.-Ham. ex Roxb.

Tree; leaflets 5-12 pairs; flowers in pink clusters (“pink Cassia”), often behind

leaves. Native of the Malayan mountains.

Cas. obtusifolia L.

Herb, resembling C. tora but not foetid; flowers yellow. Weed, American

origin.

Cas. occidentalis L.

Herb or subshrub; flowers orange-yellow. Weed. native of S. America.

Cas. siamea Lam.

Tree; flowers bright yellow. Ornamental and timber tree; native of Malaya

and Thailand.

Cas. tora L.

Herb or subshrub; leaflets 3 pairs; flowers pale yellow; pod narrow and

curved, 4-angled.

Centrosema plumieri Benth.

Twining subshrub; leaves 3-foliolate; flowers white with 2 large reddish violet

blotches. ““Butterfly pea’’, native of America.

Clitoria laurifolia Poir.

Erect herb; leaves 3-foliolate; flowers white to pale blue. Native of Brazil.

Clit. ternatea L.

Slender twiner; leaves 5-7 foliolate; flowers blue (often in double form). Native

home unknown (either S. America or the Malay Islands).

Crotalaria bialata Schrank

Subshrub; leaves 1-foliolate; stems winged; flowers pale yellow; in sandy

places or on waste ground, Jurong, Changi (Ridley 4671). Vern. Kachang

hantu darat. Called C. alata Buch.-Ham, ex D. Don in Ridley’s Flora.

Crot. mucronata Desv.

Shrubby; leaves 3-foliolate; flowers yellow with reddish or purplish stripes;

in open places, Changi, Geylang. Called Crot. saltiana Andr. in Ridley’s Flora.

Crot. quinquefolia L.

Herb; leaves 5-(3—7—) foliolate; flowers bright yellow; in open waste ground,

rare, Kranji (Ridley 575).

Crot. retusa L.

Shrubby; leaves 1-foliolate; stems not winged; flowers yellow; on sea-shores,

Changi, (Ridley 2087).

Annotated list of seed plants of Singapore (II1) 259

Cynometra cauliflora L.

Tree; leaves of 1 pair of leaflets; flowers white, small, in clusters on trunk:

pod kidney-shaped, can be eaten raw or cooked. Prob. a native of E. Malesia.

vern. Nam nam, Num num.

Cyn. ramiflora L.

Leaflets 1-2 pairs; pod thick, ovoid, wrinkled, not edible; in tidal rivers and

mangroves, Jurong, Kranji (Sinclair SFN 40957).

Dalbergia candenatensis Prain

Climber; leaflets usually 5; panicles axillary; flowers white; pods thick, flat,

1- or 2-seeded rounded; in littoral scrubs, Kranji, P. Ubin (Ridley 4678),

P. Damar. Called D. torta Grah. in Ridley’s Flora.

Dalb. hullettii Prain

Shrub; flowers white, produced when the leaves are fallen; rare, in open

swampy ground, Seletar (Ridley s.n. in 1896).

Daib. junghuhnii Benth.

Woody climber; flowers small, greenish white; in secondary woods, Changi,

Sembawang, P. Ubin, Tanglin (Hullett 161). Called D. scortechinii Benth. in

Ridley’s Flora.

Dalb. parviflora Roxb.

Large climber, spiny. Bukit Timah (Ngadiman 358).

Dalb. rostrata Grah.

Woody climber; flowers white; Gardens jungle, Changi (Ridley 6090), Bukit

Mandai.

Dalb. velutina Benth.

Woody climber; Tempinis, Changi, Seletar (Sinclair 39617).

_ Delonix regia Rafin.

A large deciduous tree with umbrella-shaped crown; leaves bipinnate; flowers

scarlet (hence “flame of the forest’); pods woody, very large, with 20-40

seeds inside, splitting open on the tree. Native of Madagascar, widely planted.

Vern. Gul mohm, JAJA

Derris amoena Benth.

Large climber; leaves pinnate, leaflets about 7; flowers pink, in panicles;

pods flattened, 2-seeded; once found in Siglap (Ridley 10380).

Der. elliptica Benth.

Native of continental Asia, often planted for the roots which are used for

killing insects on vegetables and for catching fish. Vern. Tuba, 72/4 % =

260 Gardens’ Bulletin, Singapore — X XVI] (1974)

Der. heptaphylla (L.) Merr.

Woody climber; flowers greenish; tidal river, Yeo Chu Kang, P. Ubin

(Hullett 6194). Called D. sinuata Thw. in Ridley’s Flora.

Der. heterophylla (Willd.) Backer

Scandent shrub; leaflets 3-5; flowers pink; pods thin, flat, 1-seeded; common

on sea coast and by tidal rivers, Changi, P. Ubin, Kranji (Ridley s.n. in 1893),

etc. Called D. uliginosa Benth. in Ridley’s Flora. Vern. Akar ketuil.

Der. scandens Benth.

Scandent bush; flowers white; Pulau Merambong (Ridley 9492).

Der. thyrsiflora Benth.

Scandent bush; flowers in dense erect panicles, white; pods long and thin,

reddish; common in open places, Bukit Timah, Tanglin, Changi (Ridley 2092).

Desmodium heterocarpon (L.) DC.

Shrub; leaves 3-foliolate; flowers white or deep crimson; pods of several

1-seeded joints; in open places, Changi (Ridley s.n. in 1890), Tanglin, Seletar.

Called D. polycarpum DC. in Ridley’s Flora. A variety (var. ovalifolium

Prain) with creeping habit, round silky leaflets and pale lavender flowers was

recorded at Ang Mo Kio. Vern. Kalumbar.

Desm. heterophyllum DC.

Creeping herb; flowers pink; common in the grassland all over the island;

pod 3-4 mm wide. Singapore (Hullett 418).

Desm. triflorum DC.

Like the above species, pods narrower (2-2.5 mm wide). Roadsides and

grassland; Tanglin, Changi (Ridley 2079), Bukit Timah.

Desm. umbellatum DC.

Large shrub; flowers white; on sea coasts around the island; Pulau Senang

(Sidek 87).

Dialium laurinum Baker

Large tree; leaves pinnate, leaflets 5-7; pods ovoid, velvetely dark brown,

each containing one seed surrounded by sweet edible pulp; the hard, heavy

wood is well-known; Kranji, MacRitchie Reservoir (Sinclair 8921). Vern.

Kraniji.

Dial. maingayi Baker

Like the above, with 7-9 leaflets; Catchment Area, Bukit Timah (Ngadiman

34621).

Annotated list of seed plants of Singapore (I11) 261

Dial. wallichii Prain

Like the above, with 9-11 leaflets; Garden’s jungle, Bukit Timah (Sinclair

39568).

Dolichos lablab L.

Also called Lablab niger Medik.; prob. native of India, cultivated for its

edible pods and beans (“hyacinth bean”). Vern. Kachang kara, Karkaras, BG -

Entada spiralis Rid.

Woody climber; leaves bipinnate, pinnae 4, leaflets usually 4 pairs in a

pinna; flowers small, in dense racemes; pods curled with 5-11 very large seeds

(3 x 5 cm); in secondary forests, formerly at Tanglin (Ridley 2102), Chua

_Chu Kang, now survives in Catchment Area.

Erythrina fusca Lour.

Tree; leaves 3-foliolate, terminal leaflets much longer than broad, tapering

to base; flowers dark purple. A widely distributed species from India to the

Pacific Islands, cultivated.

Ery. orientalis Murr.

Tree; terminal leaflets broader than long, truncate at base; flowers dark

scarlet. Also called Ery. indica Lam., from India to the Pacific Islands, but

cultivated here. Vern. Indian coral tree, #iJ Ha -

Ery. parcelli Hort.

Leaflets with a broad yellow midrib and several yellow stripes along the

side-veins, the base truncate; flowers bright orange red; cultivated in gardens

Or as a roadside tree. Native of Trop. Asia.

Ery. subumbrans Merr.

Like Ery. orientalis, but terminal leaflets nearly as long as broad; and flowers

comparatively smaller (3-5 cm long); native of Malesia. cultivated.

Gliricida sepium Walp.

Small tree; leaves pinnate; leaflets 13-17; flowers pinkish native of Mexico

(hence ‘‘Mexican lilac’’); cultivated in gardens or along roadside.

Hymenaea courbaril L.

Tall, fast-growing tree with a spreading crown; leaves consisting of a pair

of oblique leaflets; flowers white; pods oblong with 2 or few seeds inside,

stinking. The resin (collected by cutting the bark) is called ‘‘copal’ of the

trade. “Locust tree’, native of the West Indies.

Indigofera tinctoria L.

Shrubby, much-branched; leaves pinnate; leaflets 5-13; flowers greenish

yellow. Indigo, a deep blue dye, is obtained by allowing plants to soak and

ferment in water. Native of India and Ceylon. Vern. Nila, tarum, #2 % -

Indig. hirsuta L.

Herb, stems, leaf rachis, calyx and pods covered with long brown hairs;

leaflets 5-9; a weed, found on sandy seashores.

262 Gardens’ Bulletin, Singapore — X XVII (1974)

Intsia bijuga (Colebr.) O. Ktze.

Small tree; leaflets in 1 (or less. commonly in 2) pair; pods woody, large,

oblong, slightly flat; common along sea coast, Bajau (Ridley 4675), Changi,

Kranji. Called Afzelia bijuga A. Gray and A. retusa Kurz in Ridley’s Flora.

Vern. Merbau ayer.

Koompassia malaccensis Maing. ex Benth.

Gigantic tree with bug buttresses; leaflets 5-9; pods 1-seeded, flat, oblong.

Formerly very common alli over the island. now restricted to Bukit Timah

and Catchment Area (Ridley 6403). Vern. Kempas.

Kunstleria ridleyi Prain

Large climber; leaflets 5; flowers small, dark purple, in large panicles; pods

golden pubescent; in woods, Gardens jungle, Mandai (Corner 37735), Seletar.

Endemic to Singapore.

Leucaena leucocephala (Lamk.) De Wit

Shrub; leaves bipinnate; flowers white, crowded in globular heads; pods thin

flat, 2-valved; seeds many. “‘Lead tree’, native of trop. America; occurs near

villages. Called L. glauca Benth. in literature. Vern. Petai Jawa.

Mezoneuron sumatranum W. & A. ex Benth.

Prickly climber; leaves bipinnate; flowers flame-coloured; pods bright red;

in edge of woods and open places, not common; Bukit Timah, Kranji, Jurong.

Vern. Akar Darah Blut (Eel’s blood).

Millettia atropurpurea Benth.

Tree; crown large dome-shaped; leaflets 9-11; flowers large, dark purple;

pods thick leathery, 1—2-seeded. Native of the Malayan mountains, planted;

fine specimens in MacRitchie Reservoir. Vern. Tulang daing.

Mill. eriantha Benth.

Woody climber, covered with golden hairs; leaflets 2—3 pairs; flowers coppery

red; pods woody, 1-seeded; found from Gardens’ jungle, Bukit Mandal,

Tanjong Gul (Sinclair 10750). Called Adinobotrys erianthus Dunn in Ridley’s

Flora. Vern. Akar pua.

Mill. maingayi Baker

Big climber with hanging panicles of scented pink flowers; leaflets 11-17; pods

l-seeded; recorded from Reservoir woods, Chua Chu Kang and Tanglin

(Hullet 145). Called Padebruggea maingayi (Baker) Dunn in Ridley’s Flora.

Mimosa invisa Mart. ex Colla

Subshrub or herb, scandent at base, erect above; pinnae 5-9 pairs; flower-heads

reddish purple; in open places; native of trop. America.

Mim. pudica L.

The sensitive plant; subshrub, spreading; stems prickly; leaves bipinnate,

pinnae in two pairs, arranged at the top of a long stalk; flower-heads pink;

pods jointed, the edges covered with bristles; native of trop. America,

naturalized in waste places. Vern. Rumput si-malu, &7EE o

Annotated list of seed plants of Singapore (III) 263

Mim. sepiaria Benth.

Shrub; pinnae 5-9 pairs; flower-heads white or pale yellow; native of trop.

America.

Maughania strobilifera St. Hil. ex O. Kuntze

Shrub, leaves 1-foliolate; on seashores, uncommon; recorded from Changi

(Ridley 2076) and Blakang Mati. Formerly called Flemingia strobilifera Br.

Mucuna bennetti F. Muell.

Climber; leaves trifoliolate; flowers large, scarlet, in racemes. Introduced from

New Guinea. Seeds obtained only through hand-pollination.

Muc. gigantea (Willd.) DC.

Climber; leaves large, trifoliolate; flowers greenish white; recorded from

Blakang Mati and Telok Paku (Sinclair 9972).

Neptunia natans (Linn. f.) Druce

A water-sensitive plant; floating; leaves bipinnate; leaflets numerous; flowers

bright yellow; perhaps a native of S.E. Asia; in ditches, formerly very

abundant, now very rarely cultivated as a vegetable. Also called N. oleracea

Lour.

Ormosia bancana (Mig.) Merr.

Tree; leaflets 7-13, small, deep green; flowers white; pods round; seeds 1-3,

red; Catchment Area, Changi, Pasir Panjang (Ridley 8096). Formerly called

O. parvifolia Baker.

Ormos. macrodisca Bak.

A big tree; flowers pink; pods flat and round, with a large scarlet and black

seed; a timber tree, very rare, once recorded in Dalvey Road (Ridley 2103).

Ormos. sumatrana (Miq.) Prain

Tree; flowers white; rare, recorded from Tanglin and Seletar (Ridley 5574).

Pachyrhizus erosus (L.) Urb

Twining herb, spreading, with a tuberous tap root; leaves 3-foliolate. Native

of America; leaves, beans and seeds poisonous but the tuberous roots (“yam

bean’’) are edible. Also called P. tuberosus Spreng. Vern. Bengkuang, FG

Parkia speciosa Hassk.

Tall tree, buttressed; leaves bipinnate; leaflets 20-35 pairs on side stalks;

flowers very small, crowded on pear-shaped heads; pods large, flattened and

twisted, with onion-smell, edible; scattered in forests, Bukit Timah, Catchment

Area. Vern. Petai.

Peltophorum pterocarpum Backer ex Heyne

Tree, with dome-shaped crown; leaves bipinnate; flowers showy, yellow

(hence “‘yellow flame’’); pods flat, thin, with a wing around; formerly a native

tree at Changi coast (Ridley 4676), now extinct, but widely planted as orna-

mental or as roadside trees. Also called Pelt, ferrugineum Benth. Vern. Balai,

ETEK ©

264 Gardens’ Bulletin, Singapore — X XVII (1974)

Phaseolus lunatus Linn.

Lima bean, native of S. America, occasionally cultivated for its edible beans.

Phas. vulgaris Linn.

French bean or kidney bean, native of trop. America, commonly cultivated

species of Phaseolus (or Vigna) of which the dried beans are selling in stores

including: Phas. aureus Roxb., the green gram, #47 » native of India, and

Phas. angularis F. W. Wright, the adzuki bean, #5 » native of the Orient).

Pithecellobium clypearia Benth.

Small tree; leaves bipinnate; pinnae 2-9 pairs; leaflets 4-14 pairs, rhombic,

asymmateric; flowers in large panicles; pods flattish, lobed, twisted and loosely

curled; common in secondary forests, Seletar, (Goodenough 2097), Chua Chu

Kang. Vern. Petai belalang.

Pith. contortum Mart.

Small tree, like P. clypearia but twigs rounded (not angled as in the latter);

common, Tanglin, Changi, Seletar, Bukit Mandai (Ridley 4755).

Pith. dulce Benth.

Tree, with a bushy crown; pods pale greenish, seeds black covered by thick

white pulp. A native of trop. America, erroneously called “Madras Thorn”.

Pith. ellipticum Hassk.

Tree; pods flat, 3 cm wide, twisted into a ring; Seletar, P. Merambong

(Corner 29963), P. Ubin. Vern. Jering hutan.

Pith. globosum (Bl.) Kosterm.

Small tree. Tanjong Bunga (Ridley 6408). Called P. affine Baker in Ridley’s

Flora.

Pith. jiringa (Jack) Prain

Tree, pods large, 5-6 cm wide, strongly swollen at each seed, curled and

twisted, garlic smelled, can be cooked and consumed as a vegetable, wild in

secondary forests, commonly cultivated for the edible pods. Called P. lobatum,

Gardens jungle (Ridley s.n. in 1897). Vern. Jering.

Pith. microcarpum Benth.

Small tree, flowers white; pods curly, bright orange; in secondary woods,

Tanglin, Changi (Ridley 188) Chua Chu Kang, Bukit Timah.

Pith. splendens Corner

Large tree; pods flat, swollen at the seeds, slightly curved; rare, Bedok

(Ridley 8446), Seletar. Called P. confertum Benth in Ridley’s Flora.

Pongamia pinnata (L.) Pierre

Small tree; leaflets 5-7; flowers pink; formerly common on the coast, Changi,

Siglap, Jurong, P. Ubin (Ridley) s.n. 1891), now occasionally found in

Annotated list of seed plants of Singapore (III) 265

Psophocarpus tetragonolobus DC.

Perennial twinner, with a tuberous root; leaves 3-foliolate; flowers pale blue-

violet; pods cylindric with 4 wings (“four angled bean”). Prob. native to

Psoralea corylifolia Linn.

Annual herb; leaves simple. Native of India, grown on several occasions in

Botanic Gardens; the seeds are used medicinally. #4 fi§

Pterocarpus indicus Willd.

Large tree, leaves pinnate, flowers yellow, fragrant; pods 1-—2-seed, wood

valuable. Native of the continental Asia and Malaya; extensively planted as

roadside trees, easily propagated by large cuttings. Vern. Angsana, Sena,

FU Eats

Samanea saman (Jacq.) Merr.

Large tree, trunk usually branched; crown broadly dome-shaped; leaves

bipinnate. “‘Rain tree”, a native of Tropical America, often planted as a shade

tree in large gardens or on roadsides. Also called Enterolobium saman Prain.

Saraca indica Linn.

Small tree; leaves pinnate; leaflets 4-6 pairs; flowers orange-yellow, in large

clusters on trunk or branches; pods large, flat, purple. Native of Malaya and

the continental Asia, Cultivated. Vern. Gapis. Yalan.

Sar. thaipingensis Cantley ex Prain

Like above species, but leaves larger and leaflets with a conspicuous stalk.

Native of Malaya, sometimes cultivated in gardens.

Serianthes dilmyi Fosberg

Tree; leaves bipinnate; pods woody, thick; once collected from Bukit Timah

(Wallich 5285). Recently collected from Pulau Pawai (Sinclair 38902). Called

Seri. grandiflora Benth. in Ridley’s Flora.

Sesbania grandiflora Pers.

Shrub or small tree, with showy white or blue flowers. Native country

unknown, cultivated as an ornamental plant for its edible leaves. Vern.

Turi, KiE HS

Sindora wallichii Grah. ex Benth.

Large tree with a massive crown; leaves with 3-4 pairs of leaflets; pods oval

to oblong, flat, 1-seeded, covered with close, stout hard spines; Gardens’

jungle, Changi (Bakar s.n. in 1893). Vern. Sepetir daun tebal.

Spatholobus ferrugineus Benth.

Large climber, sometimes to the tops of the tallest trees; leaves 3-foliolate;

flowers dark purple; pods flat, thin, with 1 seed near the tip, indehiscent;

Gardens’ jungle, Tanglin, Changi (Ridley 3609a) Bukit Timah. Vern. Ajar

Sejangat.

266 Gardens’ Bulletin, Singapore — X XVII (1974)

Spath. maingayi Prain

Flowers white or pinkish white; Gardens’ jungle, Bukit Timah Road (Ridley

6397).

Spath. ridleyi Prain

Flowers white or yellow; Botanic Gardens (Ridley 6401), Chua Chu Kang.

Strongylodon macrobotrys A. Gray

“Jade vine’, from the Philippines, sometimes cultivated for its large, hanging,

blue-green flowers.

Tamarindus indica Linn.

Large tree, with a dense, rounded crown; leaves pinnate; leaflets 10-20 pairs;

flowers pale yellowish; pods brown. The pulp of the ripe pods is edible, also

employed in the native medicine; timber valuable. Native of tropical E. Africa

and W. Asia. Vern. Asam jawa, Tamarind, 72ti=- o

Tephrosia noctiflora Bojer ex Baker

Bushy herb; leaflets 13-19; flowers white or pinkish; on waste grounds. Native

of Africa.

Uraria crinita Desv. ex DC.

Shrubby; leaflets 5-7; flowers lavender, in dense racemes, to 30 cm long; in

open waste places. Native of Tropical Asia.

Vigna marima Merr.

Small twiner; leaves 3-foliolate; flowers yellow (“yellow vetch’’); on sandy

seashores, Changi (Ridley s.n. in 1890), Blakang Mati. Called V. retusa Walp.

in Ridley’s Flora.

Vigna unquiculata (L.) Walp.

Twining herbs; leaves 3-foliolate; prob. native of India; two forms are

cultivated: ssp. sinensis Fruw. (pods up to 30 cm long, vern. Kachang bol,

¥L@) and ssp. sesquipedalis Fruw. (pods 30-90 cm long, vern. Kachang perut

ayam, $£§[’7@ ); pods of both forms are served as vegetables.

Zornia diphylla Pers.

Creeping herb; leaves of a pair of narrow leaflets; flowers yellow; pods spiny;

in Open sandy places, A pantropical weed.

PUBLICATIONS OF THE BOTANIC GARDENS

SINGAPORE

1. The Agricultural Bulletin of the Malay Peninsula (Series I).

Only Nos. 3, 5, 7, 8 and 9 available, at 20 cents each.

2. The Agricultural Bulletin of the Straits and F.M.S, (Series II).

Vols. 1-10, 1901-1911, monthly issues.

Many parts available.

Price: $5 per volume, 50 cents per part.

3. The Gardens’ Bulletin, Straits Settlements (Series III).

Vols. 1-11, 1912-1947. i

Vol. 1 (1-5) January-May 1912 issued under title of Agricultural Bu

of the Straits & F.M.S. 3

Prices on application.

4. The Gardens’ Bulletin, Singapore (Series IV).

Vols. 12-27, 1949-1971. Vol. 28 Pt. I in print.

Price: Vol. 13 Pt. I (New impression) $12 per copy, $20 per vol.

Vols. 12 & 14 $13 per vol.

Vol. 15 $20 per vol.

Vols. 16-25 $25 per vol. Individual parts vary in price.

Vol. 26 $36 per vol. (Pt. I $18; Pt. If $18).

Vol. 27 Pt. I $18, Vol. 28 Pt. I (in press).

Frequency: Parts are published as materials become available.

5. Materials for a Flora of the Malay Peninsula, Monocotyledons.

Parts 1, 2 and 3 remain available.

Price: $10 per set, $5 per part.

6. Annual Reports.

1909-19772.

7. (a) Malayan Orchid Hybrids by M. R. Henderson and G. H. Addison. $15.

(5) Malayan Orchid Hybrids, Supplement I by G. H. Addison. $21. Z

8. A Revised Fiora of Malaya.

(a) Vol. 1, Orchids, by R. E. Holttum. $30 (3rd ed. 1972 Impr.).

(b) Vol. 2, Ferns, by R. E. Holttum. $20 (2nd ed., 1968).

9. Boletus in Malaysia by E. J. H. Corner. $50 (1972).

Items 1-6 obtainable from the Commissioner, Parks & Recreation Division, PW

Botanic Gardens, Cluny Road, Singapore 10. -

Items 7-9 obtainable from Singapore National Printers (Pte) Ltd, Upper Serar ng

Road, Singapore 13. ag

Prices quoted are in Singapore Dollars ay

Overseas postage is extra >:

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