CONTENTS

Vol. XXVI

PART I — 30th December, 1972

KRAMER K. U.: The Lindsaeoid Ferns of the Old World VI — Continental

Asia, Japan and Taiwan

Forest Research Institute, rneenes Malaya: Notes on the : Systematy of Malayan

Phanerogams XI- XVII :

WHITMORE T. C.: Studies in id irantabe ti vs M. lowii sd

DRANSFIELD J.: The Genus Johannesteijsmannia H. E. Moore Jr. ... ve

Hitt R. D.: Soil Moisture under Forest, Bukit Timah Nature Reserve,

Singapore aie

RATNASABAPATHY M.: — — Sanedd jeval (Kedah Peak), Sita yela

REVIEW: Illustrated Guide to Tropical Plants (P. S. Ashton)

REVIEW: Pollen Flora of Taiwan (H. Keng)

STONE B. C.: Arthrodactylis and Pandanus: A brief sitcmuat on the /Chavkcteres

Generum Plantarum’

Lim SiEw-NGo: Cytogenetics aid Pe ucaaiiiy ‘a the pe $ Globba é: in Malaya

— IV Distribution in relation to polyploidy

HsuAaN KENG: Two new Theaceous plants from Malaysia and a - proposal to

reduce Tutcheria to a synonym of Pyrenaria , 5 os

TIXIER P.: Mount Maquiling Bryoflora

SHIGEO KURATA and MASAMI TOYOSHIMA: Philippine ee of ee

CorRNER E. J. H.: Studies in the basidium — spore es and the Boletus

spore ee a oo

PART II — 15th September, 1973

WHITMORE T. C.: Frequency and Habitat of Tree ing in the Rain Forest

of Ulu Kelantan

HouitruM R. E.: A New uci speeds Mt. Kitabale

MariE A. MarTIN: Notes on the ve of the desert "Mciunieatnt

Cambodia ;

HSUAN KENG and E. ‘ Heuer: The Xylopia malayana fruit: ‘uipadindbince

if its dehiscence us}

SHIGEO KuRATA: Nepenthes from ) Witigaiaes me ale 4 =

HsuaAN KENG: Annotated list of seed plants of Singapore (I)

DRANSFIELD J.: Korthalsia hispida Becc. in Malaya

Rao A. N. and LEE War CHINH Pollen bomen and Cetinination in Sortie

Orchid Hybrids

Nayar M. P.: A Review of the tir Credghtoh Stapf. Bitbdistienatnduilas

KUEH TIONG-KHENG: New Plant Disease Records for Sarawak for 1970 and

1971

KOCHUMMEN K. M. iad As Maw Wiaresi-sne Notes on the Systematy of Malayan

Phanerogams XVIII-XXII

ANNE JOHNSON: A ey of the occurence of epiterranean soil algae in Sings-

pore Island ;

GALIL J.: Pollination in Bisteteos Figs:

Ceratosolon hewitti ue

INDEX

“Pollination of Woiia: fistulosa by

Pages

1-48

49-61

63-83

85-93

95-110

111

112

113-114

115-126

127-135

137-153

155-158

159-194

195-210

211

213-222

223-225

227-232

233-237

239-244

245-257

259-261

263-268

269-287

289-302

303-311

313-325

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INDEX, VOL. XXVI

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

boldface. Plates are expressed by page numbers with letters.

Acanthocladium longipilum 140 Alternanthera 263

piliferum 140 longissima 267

Achnanthes 99 ECE 207 CUS

Acroporium sp 218 Altingia siamensis 214

diminutum 140 Amanita 159 162 164

hermaphroditum 141 Amomum spp 218

secundum 141] kravanh 218 221

Actinotaenium 103 Amoora 210

Adenanthera 210 Anabaena 96

pavonina 219 Anacolosa maingayi 285

Adiantum orbiculatum 21 Ananas 263

tenuifolium 5

Anaxagorea 224

Adina 210 Andrachne calcarea 50

Aerobryidium filamentosum 139 fruticosa 50

Aerobryopsis longissima 139 Anerincleistus 259 261

var. prostratula 139 philippinensis 259 261

Afzelia xylocarpa 222 Anisoptera 207

Agaricus 159 sp 214 219 220

Agathis 235 Annona 263

alba 236 Anthocephalus sp 219 220

dammara 210 236 chinensis 204

loranthifolia 236 Antiaris toxicaria 206

Agrostistachys borneensis 52 Antirrhinum 255

latifolia 52

A h 222

leptostachya 52 a ome a 2 1as igh

longifolia 52 Str. Org kg

var leptostachya 52 Aquilaria 209

var latifolia 52 krassna 217 221

var malayana 52 Arachnis 248 263

sessilifolia 52 Maggie Oecei 245 247 248 249 254 256

Albizzia sp 218 Aranda 248 249

Alchornea villosa 53 Hilda Galistan 245 247 248 249 254

tilifolia 53 256

Alloburkillia album 55 Lucy Laycock (long spray) 245 247

Aloe 133 248 249 254 256

(short spray) 245 247 248 249

Alphonsea 224 254 256

Alpinia spp 218 220 Wendy Scott var Greenfield 245 247

Alstonia 210 248 249 254 256

313

314 Gardens’ Bulletin, Singapore — X XVI (1972-73)

Aranthera 248

Lilliput 245 247 248 249 254 256

Araucaria 235 236

bidwillii 236

cunninghamii 236

excelsa 236

kunsteinii 236

Araucariaceae 236

Archilejeunea mariana 141

Areca triandra 218

Aromadendron elegans 205

Artocarpus integer 206

kemando 210

lanceifolius 207

rigidus 204

scortechinii 204

Arundinaria falcata 222

Ascochyta 265

Asplenium 217

nidus 218

Athrodactylis 113 114

spinosa 114 113

Austrobuxus nitidus 51

var montanus 51

Baccaurea 51

griffithii 208

motleyana 51

oxycarpa 217 219

pubescens 51

sapida 219

Bambusa 220

Barbella cf clemensiae 139

Barringtonia 207

payensiana 284

basidiospores

adaxial patch 162

angle 163

elongate 161

globose 165

interfering tetrads 162

length 165

measurement in Boletus 179 180

positioning 163

reversed 163

spacing 161

lateral in tretrad 164

sterigmatic length 165

sterigmatic patch hypothesis 169

tangentially compressed 162

basidium

apex 164 165

geometry 163 167

length 165

unit 166

with 6-8 spores 169

Batrachospermum 97 106 106A 106B

Bazzania sp 218

Begonia 263

Bixa 263

Blumeodendron borneense 53

concolor 53

elateriospermum 53

vernicosum 53

tokbrai 53

Boletellus 174 175 178 189

Boletus 162 182 189 191

spore 166 174

albo-ater 175 178 179 182 183

altissimus 182

aureomycelinus 178 180

emodensis 178 179 188 189

funerarius 178

longicollis 179 182 189

nanus 175 178 179 182 184

obscurecoccineus 189

phaeocephalus 178 179 180 181

ravenelii 162 178 180

retisporus 178 179 182 185

singaporensis 182

umbilicatus 178 179

Bombax ceiba 221 222

valetonii 205

Botryodiplodia theobromae 267

Bouea 210

Bowenia 233

Brassica 263

Bromelia sylvestris 113

Bryophyllum 255

Buchanania reticulata 222

Buddleja 133

Bulbochaete 102

Bulbophyllum 218

Burkillia alba 55

Burkilliodendron album 55

Calamus 81

Callitris 235

mackayana 236

= =

Index

Calocedrus 235

formosana 236

Calophyllum 205

sp 219

austrocoriaceum 269 270

coriaceum 270

inophylloide 270

var singapurense 270

intramarginale 270

rupicolum 271

var elatum 270

tahanense 270

Calymperes serratum 139

Camellia 130 132 135

lanceolata 127

shinkoensis 135

spectabilis 134

virgata 135

Campnosperma 204

Canangium 264

odorata 210

Cantharellus 163

Capsicum 264

Carallia brachiata 207

Careya sphaerica 222

Carica 264

Carludovica 63

Cassytha filiformis 218

Castanopsis 208

cambodiana 219

pierrei 217

Cattleya guatmalense 255

Centrolepis cambodiana 220

Centrosema 85

Cephaleuros virescens 263 267

Cephalotaxus 233

Ceratocystis paradoxa 264

Ceratolejeunea maritima 142

Ceratozamia 233

Cercospora 264

capsici 264

chrysanthemicola 264

fukushiana 265

sorghii 267

Cerebella andropogonis 265

Chaetomitrium sp 218

perarmatum 140

Characiopsis 101

315

Characium 101

Cheilolejeunea ceylanica 142 218

verdonii 218

Chiloscyphus communis 141

Choanephora circinans 265

cucurbitarum 263

Choriophyllum montanum 51

Chroococcus 96

Chrysanthemum 264

Cicinobella 267

Cinnamomum sp 214 217

aff javanicum 217 219

Cintractia exicola 264

Citrus 264

Cladosporium oxysporium 264

Clastobryum cuculligerum 140

Clavaria 164 167

Closterium 103 104 106 106C

Clypeolella ricini 267

Cochliobolus geniculatus 266 267

lunatus 266

Cocos 264

Codiaeum 264

Coelostegia 210

Coffea 264

Colletotrichum capsici 263 266 267

graminicola 265

Cololejeunea 152

complex 145

sp 218

armata 142 145 149 151

flavicans 142

floccosa 142

gynophthalma 142

haskarliana 142

var thermarum 142 148

var luzonensis 147 148

hispidissima 142 145 146

maquilinensis 142

aff nymanii 142

panchoana 142

pseudoschmidtii 142 145

schmidtii 145 146

selaginellicola 142 149 150

venusta 145

vulcania 142

aff yulensis 142

316 Gardens’ Bulletin, Singapore — X XVI (1972-73)

Colona merguensis 59 Dacrydium 235

Columbia curtisii 59 elatum 71 235

diptera 59 Dactylococcopsis 96

Colura acutifolia 142 Daemonorops 81

Combretum quadrangulare 222 aff ne, 218

Coniothyrium fuckelii 267 Dalbergia aff. cochinchinensis 222

; aff lanceolaria 222

Coprinus 159 165. nigrescens 221 222

Coptosapelta 135 Davallia biflora 4

Corticium rolfsii 263 265 266 gracilis 8

salmonicolor 264 266 luzonica 8

solani 267 PE ae 5%

; schizophylla

Corypha lecomtei 222 tnisiotia’ s

Cosmarium 103 104 105 106 106D var lata 4

Craterellus 163 trichomanoides Bedd 29

Cratoxylum 209 221 Debarya 102

arborescens 210 Dendrobium 218

formosum 207 Desmogonium 98

Creaghiella philippinensis 259 261 Dialium 208

purpurea 259 260 261 cochinchinensis 222

setosa 259 260 261 Dicksonia linearis 8

Crudia 269 repens 45

gracilis 285 Dicranodontium uncinatum 137

subsimplicifolia 285 eI

viridiflora 285 Dicranoloma braunii 218

Cryptomeria 235 Dicranopteris sp 220

japonica 236 Dillenia 208

Ctenolophon parvifolius 206 oa 222

Cucumis melo 255 Dinobryon 100 106

Cunninghamella echinulata 264 Dinococcus 98 106B 106C

Cupressaceae 236 Dioon spinulosium 234

Cupressus 235 Diospyros 207

macrocarpa 237 erhetioides 222

Curvularia eragrostidis 263 265 266 Dischidia aff imbricata 218

pallescens 265 ee ae

senegalensis 267 Diplodia zeicola 267

Cyathea latebrosa 218 Dipteris conjugata 71

podophylla 218 Dipterocarpus 205

appendiculatus 205

Cycadaceae 234 apes 210

Cycas 234 baudii 208

circinalis 234 chartaceus 210

revoluta 234 cornutus 199 207

rumphii 234 costatus 219 220

siamensis 234 crinitus 199 206

Cylindrocystis 103 104 gracilis 210

grandiflorus 207

Cymbella 99 intricatus 222

Cynometra malaccensis 199 208 kunstleri 207

Cyrtopodium punctatum 255 oblongifolius 207

Index 317

Dipterocarpus (cont.) Eugenia 209

obtusifolius 221 222 Euglena 101

sublamellatus 207 Eunotia 98

tuberculatus 222 .

turbinatus 219 meee ‘ee 9

verrucosus 207 pay*

nummularia 218 Euphorbia 133

Drechslera musae-sapientum 265 Fagraea 207

Drepanolejeunea bakeri 143 aetna 214

dactylophora 143 Feroniella lucida 222

thwaitesiana 218 Ficus 209 264

Drosera 220 sp 214 219 221

Drynaria 217 218 Fimbristylis 264

Dryobalanops 205 Fissidens aff nobilis 218

aromatica 205 sylvaticus 137

oblongifolia 199 205 Flemingia 265

Durio 208 Fomes lignosus 264 266

Dyera costulata 208 Frullania apiculata 143

Dysoxylum cauliflorum 217 integristipula 143

aff hoaensis 217 squarrosa 143

procerum 217 ternatensis 143

Ectropothecium buitenzorgii 141 218 — Frustulia 99

falciforme 141 rhomboides 106

ichnotocladum 141 Pasauim 266.267

Elaeis 264 equiseti 264 265

Elaeophorbia 133 heterosporum 264 266

monileforme 264

Elateriospermum tapos 199 207 oxysporum 267

Encephalartos 234 semitectum 267

villosus 234 solani 264 263

Endospermum diadenum 53 Garcinia 210

malaccense 53 54 199 204 atroviridis 272

moluccanum 54 bancana 276

peltatum 54 var curtisii 276

Endotrichella compressa 139 burkilli 273

dimorphophylla 144 cantleyana 273 |

maquilinensis 139 144 ni age ey 214

: caloneura

Engelhardtia 204 cataractalis 273 274

Entoloma 159 cornea 275

Eria 218 costata 272

Eriobotrya japonica 255 cowa 273

: ; curtisti 276

Eriopus microblastus 140 cuspidata 271 272

parviretus 140 densiflora King 277

Ervatamia curtisii 50 diversifolia 272

jasminiflora 50 dulcis 273 277

pauciflora 50 dumosa 273 274

Erythrophloeum cambodianum 221 eugeniaefolia 272 277

Fuastrum 105 106A mente

uast um forbesii 272

Eugeissona 65 gaudichaudii var minor 274

tristis 71 glomerata 277

318

Garcinia (cont.)

griffithii 272

hanburyi 217

holttumii 272

hendersoniana 273 275

hombroniana 272

alliance 276

kunstleri 275

lanceolata 278

nervosa 273 277

maingayi 272

malaccensis 272

mangostana 272

merguensis 272 278

minutiflora 272

monantha 272

montana 272

murdochii 272

murtonit 272 275

nigrolineata 273 275

opaca 272 276 277

var dumosa 277

parvifolia 273 |

penangiana 269 272 275 276

group 275 276

prainiana 273 277

pyrifera 273 277

rostrata 272 277 278

and allies 277

scortechinii 273 274

tenuifolia 274

uniflora 273

urophylla 273 274

vilersiana 217

wrayl 277 278

xanthochymus 273

group 277

Gelonium bifarium 54

glomerulatum 52

multiflorum 54

tenuifolium 54

Gironniera 210

Gleichenia aff norrisii 218 220

Gliocladium 265

Globba

distribution 116 119 120 121

infraspecific polyploidy 116

albiflora 116 126

var aurea 116

aurantiaca 117

bracteata 123

bulbifera 123

cernua 117 118 123 124

Gardens’ Bulletin, Singapore — XXVI (1972-73)

ssp crocea 118 123 124

ssp prophyria 118 124

curtisii 117 124

fasciata 116

fragilis 116 124

holttumiit 117 125

ssp aurea 123 125

x intermedia 125

leucantha 118 123

var bicolor 118

var flavidula 118

var peninsularis 118 126

var violacea 118

marantina 123 124

patens 117 123

x cernua 125

var costulata 125

pendula 117 123

var elegans 117 126

ssp montana 117 126

strobilifera 123

unifolia 116

var sessiliflora 124

variabilis 118 123 125

ssp pusilla 118 123 125

Glochidion brunneum 50

coronatum 51

goniocarpum 50

microbotrys 51

pedunculatum 50

penangense 51

trilobum 51

villicaule 51

wallichianum 51

Gloeocantharellus 191

Gloeocapsa 96

Gloeotheca 96

Glomerella cingulata 264 265

266 267

Gluta 206

Glycine 265

Glycosmis calcicola 55

var kelantanica 57

parkinsonii 57

var ovatofoliolis 55 57

Glyptocarpa 133

Gnetaceae 237

Gnetum 235

campestre 237

gnemon 237

gnemonoides 237

kingianum 237

latifolium var funiculare 237

Index

Gnetum (cont.)

macrostachyum 237

microcarpum 237

sylvestris 237

wrayl 237

Gonystylus 210

Gordonia concentricicatrix 210

hirta 134

Guizotia 265

Gunong Jerai map 107

Hantzschia 99

Hapalosiphon 97

Heimiella 174 175

retisporus (see under Boletellus)

Helicia 286

Heliciopsis cockburnii 286

montana 287

whitmorei 287

Hendersonia zingiberi 267

Herberta cf minima 141

Heritiera 204

javanica 219

Hibiscus floccosus 203 204

Himantocladium cyclophyllum 140

Homaliodendron crassinervium 218

flabellatum 140

Hopea 203 204 214

odorata 214

pierre 219 220

Hyalotheca 105 106 106D

Hygrophorus 159 165

Hymenophyllum ramosissimum 5

Hypnodendron vitiense 139

Impatiens 265

Imperata 265

cylindrica 221 222

Intsia palembanica 199 208

Irvingia malayana 207 219 221

Isoloma divergens 44

Isopterygium albescens 141

Ixonanthes icosandra 210

Johannesteijsmannia

description of genus 64

distribution Malay Peninsula 67

fruit 64A

geography 81

history 63

key to species 66

nat. history 65

uses 65

319

altifrons 63 64B 66 67 68 69 70 70A

70B 71 73.75 81\ 82

native names 65

lanceolata 66 68 74B 78 79 80 82

magnifica 64A 65 66 68 75 76 78 80

perakensis 64 66 72 73 74 74A 75 82

Juniperus 235

chinensis 237

Kaida 113

Kayea 282

assamica 278 279

caudata 279

curtisii 281

elegans 279

kunstleri 281 282

parviflora 282

rivolorum 281 282

Keura 113

odorifera 113

Koompasia 206

excelsa 199 208

malaccensis 206

Korthalsia 81 239 240 242

echinometra 239 241

flagellaris 239

hispida 239 240 242 243

macrocarpa 239 240 241 242

rigida 239 241

robusta 239 240 242

scaphigera 239 241

scortechinii 239

squarrosa 239 242

teysmannia 241

Lactarius 163

Lagerstroemia 221 222

Lagynion 100

Lansium domesticium 210

Leersia 265

Lejeunea flava 143

Lepidozia cf. subintegra 141

Lepiota 159

Lepocinclis 101

Leptocarpus disjunctus 220

Leptocolea hispidissima 142

Leptolejeunea foliicola 143

schiffneri 143

subacuta 218

320 Gardens’ Bulletin, Singapore — X XVI (1972-73)

Leptopus australis 50 ssp. ensifolia 32

calcareus 50 flabellulata 21

hirta 50 var gigantea 27

Leptosphaeria 267 gigantea 27

typhae 267 glandulifera 2 10 45

Leptosphaerulina trifolii 264 ibe eipee

pa a griffithiana 32

Leucobryum bowringii 139 218 hainanensis 22 23 25

Leucoloma molle 137 218 heterophylla 10 13 31 33

Liberbaileya gracilis 82 himalaica 15 43

Libocedrus formosana 236 uiegts 2 ee

; javanensis 2 11 12 25

Licuala 66 81 kawabatae 2 10 16 33

Lindsaea kirkii 38

doubtful species 47 kusukusensis 44

putative hybrids 33 laguginosa 48

sect. AULACOLINDSAEA 9 lancea auct. div. 38 39 47

sect. AULACORHACHIS 39 lancea (L.) Bedd. 25 39 40

sect. ISOLOMA 44 var caudata 39

sect. OSMOLINDSAEA 40 liangkwangensis 27

sect. PSPAMMOLINDSAEA 43 lobata 14 36 37

sect. PSEUDOLANCEA 46 var epirotes 2 14 34 37

sect. SCHIZOLOMA 15 var hainaniana 2 35

sect. STENOLINDSAEA 44 lobbiana 44

subgen. ODONTOLOMA 9 10 45 longipes 21 24

subgen. SCHIZOLOMA 10 16 longipetiolata 47

annamensis 2 10 lucida 2 15 44

austro-sinica 2 11 24 25 ssp lucida 44

boniit 22 23 macraeana auct. div. 45 46

bouillodii 12 28 47 macraeana ching 46

cambodgensis 2 10 28 30 33 malabarica 2 13 15 33

caudata 2 9 39 malayensis 2 13 33

changii 44 merrillii 9 46

chienii 12 18 19 33 ssp yaeyamensis 46

var deltoidea 27 minima 44

chinensis 27 napaea 2 13 14 33

chingit 2 10 19 47 neocultrata 40

commixta 22 nitida auct. 35

concinna 44 oblanceolata 9 46

var kusukusensis 44 obtusa 2 14 34

conformis 47 odorata 2 3 40 42 43

cultrata auct. 37 var darjeelingensis 41

cultrata (Willd.) Sw. 13 14 35 37 var japonica 7 42

var assamica 43 orbiculata auct. 19 29

var attenuata 40 44 orbiculata (Lam.) Kuhn 2 11 12 15

var japonica 42

var minor 40 43

var pallens 40

20 21

key to varieties 21

davalloides Bl. 36 var chienti 19

davallioides auct. 34 var commixta 21 25 30 33

decomposita auct. 37 var deltoidea 27

decomposita Willd. 35 var gigantea 27

dissectiformis 2 7 10 16 18 var orbiculata 21 23

divergens 15 44 parallelogramma 2 14

doryphora 2 15 25 38 39 40 parasitica 9 39 40 47

eberhardtii 16 parvipinnula 30

ensifolia 13 pectinata Bl 45

Index 321

Lindsaea (cont.) Mangifera 209 265

pectinata auct. 46 Manihot 265

recedens 47 Marasmiellus scandens 265

recurvata 35 Marasmius 159 191 192

repanda 11 23 28 campanella 266

repens 2 10 45 nigripes 190 191 192

f. minor 45 ; ;

Sit iwaney 45 Maxburretia rupicola 82

var pectinata 45 Medinilla coerulescens 217 218

scandens 24 47 a Melanochyla 206

var terrestris etartyger

schizophylla 2 10 23 29 33 Melanolepis diadena 53

simulans 21 47 Melanorhoea 206

taiwaniana 47 laccifera 222

tenera auct div 28 Melastoma 221

tenera Dryander 47 Melia azedarach 219

var chienii 19 ’ : :

var conmixta 22 Meloidogyne javanica 266

var gigantea 27 Memecylon 210 221

trapeziformis var caudata 39 :

venusta 2 13 34 35 37 38 ae peeteenle

walkerae 15 43 esua

acuminatissima 278

aff assamica 278 279

assamica group 278 279

yaeyamensis 46

yunnanensis 47

Lindsaeoid ferns caudata 2719 280

key to genera 3 curtisii 281

Lithocarpus 200 elegans 279 280 282

cerifera 219 ferrea 199 205

Livistona tahanensis 71 ferruginea 278 279

Lobelia 133 kochummeniana 280

kunstleri 280 281 282

Lomatophyllum 133 var curtisii 281 282

Longetia lepidota 282

malayana 51 var parviflora 282 283

montana 51 nervosa 282

nivenii 283

Beene applanata 143 nuda 269 282 283 284

subfusca 143 parviflora 283

Lophopetalum 206 purseglovei 284

Maba sp 219 rivulorum 281

Macaranga sect pseudorottlera 62 wrayi 283 284

auriculata 62 Metroxylon sagu 65

gigantea 210 Metzgeria furcata 141

lowil 62 Michelia 265

Macromitrium cuspidatum 139 Microlejeunea cucullata 143

pungens 139 Mikania 85

semipellucidum 139

Macrozamia 234

denisonii var hopei 234

Mitragyna brunonis 222

Mniodendron fusco-mucronatum 139

Mallomonas 100 106 Mougeotia 102

Mallotus leucodermis 210 Musa 265

muticus 210 Mycena illuminans 174 176 177

vernicosus 53 Mycosphaerella 267

322 Gardens’ Bulletin, Singapore — XXVI (1972-73)

Myristica 265

Navicula 99

Neesia 210

Neonema 101

Nepenthes alata 155 157 158 227

ampullaria 227

bongso 227

burkei 155

campanulata 227 228 228A 228B 232

carunculata 228

dubia 229

globamphora 155 156 156A

gracilis 229

inermis 228

merrilliana 157

mirabilis 157 229

x alata 157

pectinata 229

petiolata 158

x alata 158

rafflesiana 229

rhombicaulis 227 229 230

singalana 231

tobaica 232

truncata 158

xX petiolata 158

Nephelium 266

lappaceum 203 208

Netrium 103 106C

Nicodemia 133

Nypa 63

fruticans 65

Ochanostachys amentacea 199 207

Odontosoria chinensis 4

var divaricata 5

eberhardtii 16

tsoongil 4

uncinella 40

Oedogonium 98 102

Ophiobolus 267

Ormosia 210

Oryza 266

Pachyrrhizus 266

Pagiantha 49

Panaeolus 162

Pandanus 113 114 219

odoratissimus L 114

tectorius 114 214

Panicum 266

Parapyrenaria 133

Parashorea lucida 199 207

Parinari anamensis 222

Parkia 200

streptocarpa 217 219

Paspalum 266

Paxillus 191

— Phylloporus aff 191

Pellacalyx 206

Peltophorum aff pterocarpa 222

Pennisetum 266

Pentace 207

excelsa 59

grandiflora 59

Pentacme siamensis 222

Pentaspadon 209

Peperomia 266

Periconiella musae 265

Pestalotia leprogena 265

Pestalotiopsis 265 266

disseminata 266

palmarum 266

theae 263

Phalaenopsis 255

Phoenix dactylifera 66

Phormidium 96

Phragmites sp 220

Phylloboletellus 191

chloephorous 191

Phylloporus 189 191

borneensis 189

cingulatus 178 180 189

coccineus 189

Phyllosticta glycinea 265

manihot 265

zingiberi 267

Pinaceae 236

Pinanga duperreana 218 219

Pinnatella cf ambigua 140

nana 140

Pinnularia 99

Pinus 235

Caribaea 236

insularis 236

massoniana 236

merkusii 221 236

Piper 266

Pithecellobium splendens 209

Pithomyces chartarum 266

sacchari 266

Index

Plagiochila sp 218

Platycerium 217

coronarium 218

Pleurotaenium 104

Pleurotus (Hohenbucehelia) testudo 266

Pluteus 162

Podocarpaceae 235

Podocarpus 235

blumei 235

imbricatus 210 217 219 235

motleyi 235

neriifolius 71 235

polystachyus 235

wallichianus 235

Pogostemon 266

Pometia pinnata 199

var alnifolia 208

var pinnata 207

Porphyrellus 174 175 189

sect. GRACILES 175 187 189

Pothos spp 218

Pottsia 63

Pratia 133

Psathyra 162

Pteridium esculentum 218

Pterocarpus sp 221

Pterocymbium javanicum 205

Pterospermum 208

Pterygellus 163

Pterygota horsfieldii 203 204

Ptychocoleus cumingianus 143

Pycnolejeunea bidentula 143

eximia 143

fitzgeraldii 143

Pyrenaria 127 129 130 132 133 134 135

acuminata 132 132A

camellioides 133

championi 134

garrettiana 133

greeniae 134

hainanensis 133

hirta 134

kunstleri 132A

microcarpa 134

multisepala 134

ovalifolia 134

pahangensis 129 130B 131

serrata 130

shinkoensis 135

symplocifolia 135

tawauensis 128 129 130A

—tutcheria complex 127

virgata 135

Pyricularia oryzae 265 266

zingiberi 267

Quercus 200

chrysocalyx 217 219 220

Racemobambos hirsuta 211

tessellata 211

Radula acuminata 218

anceps 141

borneensis 141

Rainforest tree groups

averse to granite 209

323

basal area & tree numbers 200 202

206

frequency of families 201

map 197

preferences

elevation 207

granite 204 205

sedimentary 205 206

ubiquitous 208

Randia scortechinii 210

tomentosa 222

Raphidolejunea subacuta 218

Raphidostichum eberhardtii 140

luzonense 140

piliferum 140

Rengas 206

Rhizogonium spiniforme 139

Ricinus 267

Rosa 267

Rubus 220

Russula 159 162 163 191

Saccharum arundinaceum 220

Sageraea elliptica 17 219

Salacca 63

Sanchezia 267

Sandoricum koetjape 210

Santiria 209

Sapium baccatum 208 219

Sarcotheca 207

Scaphium 207

Scaphocalyx spathacea 54

parviflora 54

Schima crenata 217 219

Schistochila aligera 141

324

Schizoloma chienii 19

ensifolium 32

griffithianum 32

heterophyllum 31

intertextum 31

javanense 25

lobatum var malabaricum 33

malabaricum 33

recurvatum 37

tenerum 28 47

walkerae 43

Schoutenia furfuracea 59

Schroederia 101

Sciadopitys 233

Scorodocarpus borneensis 208

Scytonema 97 103

Selaginella 141

sp 221

siamensis 218

Semecarpus 206

Septobasidium 264

Septoria 266

chrysanthemella 264

Shorea 206

assamica 204

atrinervosa 210

balanocarpoides 207

bracteolata 199 205

curtisii 198 199 200 205 206

dasyphylla 206

faguetiana 206

glauca 210

guiso 199 206

kunstleri 204

laevis 210

leprosula 198 199 207

macroptera 206

maxweliana 210

multiflora 199 205 210

obtusa 222

ovalis 199 206

ovata 206

parviflora 198 199 205

pauciflora 208

platyclados 199 204

resina-nigra 210

scrobiculata 210

singkawang 205

sumatrana 206

talura 222

Sindora 200 208

cochinchinensis 22] 222

Sinopyrenaria 133

Gardens’ Bulletin, Singapore — X XVI (1972-73)

Sirogonium 103

Smilax spp 218

Sorghum 267

Sphaerostilbe repens 265

Sphaerulina 264

Sphenomeris 3

key to the species 3

biflora 3 4

chinensis 2 3

var divaricata 5

var littoralis 4

chusana 5

var littoralis 4

var tenuifolia 5

gracilis 7

Spirogyra 103

Spondylosium 105

Stangeria 233

Staurastrum 105

Stauronema 264

Stenolejeunea apiculata 143

Stenoloma biflorum 4

chinense 5

chusanum 5

var littorale 4

eberhardtii 7 16

gracile 7

littorale 4

Stichococcus 102

Stigonema 97 103

Stipitococcus 100

Streblus elongatus 210

Strobilanthes 217 218

Strobilomyces 174 175 178 189

velutipes 175 182 186

Strombosia 285

maingayi 285 286

multiflora 285 286

rotundifolia 208 285 286

Suregada angustifolia 54

multiflora 54

Surirella 100

Swintonia 205 206

Symphysodon subneckeroides 139

Synaphlebium recurvatum 35

Synechococcus 96

Syrrhopodon tristichus 139

Index

Syzygium 217 219

aff cochinchinensis 217

zeylanicum 214

Tabernaemontana 49

Tapeinidium 8

key to species 8

bisseratum auct. 8

gracile 8

lineare 8

luzonicum 8

pinnatum 2 8

Taxodiaceae 236

Terminalia 205 222

alata 222

nigrovenulosa 221 222

Tetmemorus 104 106D

Tetrameles nudiflora 219

Tetraploa aristata 267

Teysmannia 63

altifrons 63

Thea shinkoensis 135

virgata 135

Thelephora 162

Thuidium glaucinum 140

meyenianum 218

Thuja 235

orientalis 237

Thujopsis 233

Tolypothrix 92

Trachelmonas 101

Trachypus bicolor 218

Trentepohlia 102 103

Tribonema 101

Trichocolea pluma 141 218

Trichomances chinense 4

Trigonostemon salicifolius 52

verticillatus 52

var salicifolius 52

Trismegistia calderensis 140

Tristania 207 219

burmanica 214 221

Triticum 255

Tutcheria 127 130 132 133 134 135

championi 130 134

greeniae 134

hirta 134

microcarpa 134

multisepala 134

ovalifolia 134

shinkoensis 135

spectabilis 130 134

symplocifolia 135

villosa 134

virgata 135

Ulothrix 102

Uncaria 267

Uredo scabies 267

Vanda 248 249 254 255 256 267

Josephine 245 247 248 249 252 253

Miss Joaquim 245 249 251 253 254

254A 255 256

Poepoe 245 247 248 249 250 251 253

254 254A 255 256

Vanilla 267

Vatica 205

odorata 219 220

Vittaria parasitica 47

resecta 47

Volvariella esculenta 266

Wallachia 63

Warburgiella cupressinoides 140

Weinmannia blumei 71

Xanthophyllum 206

glaucum 214

Xerospermum 203 210

Xylia kerri 221 222

Xylopia 224

ferruginea 210

malayana 223 224 224A

var obscura 49

Yunnania 130

Zamia 234

floridana 234

integrifolia 234

media var gutierrezii 234

var tenuis 234

tenuis 234

Zea 267

Zingiber 267

Zizyphus cenoplia 222

Zygnema 102

Zygnemopsis 102

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COED PEOPLE OPE FE GFE a

MAR 7 1973

P\ et i K. U.: The Lindsaeoid Ferns of the Old World VI te Continental Asia,

- Japan and Taiwan .

! Forest Research Institute, Kepong, Loh Notes on the i of ss ee oa

1 Phanerogams XLXVII - - . -

y DRANSFIELD J.: The Genus J ohannesteijsmannia H. E. Moore Jr. S

x Hn R. D.: Soil Moisture under Forest, Bukit Timah Nature Reserve, Singapore

(J ion RATNASABAPATHY M.: Algae from Gunong Jerai (Kedah Peak), Malaysia -

p, A) b ee: Illustrated Guide to Tropical Plants (P.S. Ashton) - : :

_ REVIEW: Pollen Flora of Taiwan (H. Keng) -, s ° :

AY hs ~ STONE B. C.: Arthrodactylis and Pandanus: A brief comment on the ‘Characteres

4 x fis Generum Plantarum’~ - - = i : é z :

Lim ‘SIEW-NGo: Cytogenetics and Taxonomy of the pelted Globba L. in Malaya

_ — IV Distribution in relation to polyploidy’ - -

OX : scan KENG: Two new Theaceous plants from Malaysia a Proposal to reduce

A _ Tutcheria to a synonym of Pyrenaria - . a 5

- TIxter P.: Mount Maquiling Bryoflora’- - - - - -

‘ a SHIGEO KuraTA and MaSsaMi ToYOSHIMA: Philippine Species of Nepenthes :

sscxmilamn E. J. H.: Studies in the basidium — spore spacing and the Boletus spore

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THE LIBRARY

GARDENS’ BULLETIN

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Vol. XXVI, Part I | 30th December, 1972

CONTENTS

PAGE

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fiz

113-114

115-126

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YR DLE EYER YO OIE OYE YEEIOIEEIEIIEOWPIOIPEO OE

Kn?

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oe ERE ea

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THE

GARDENS’ BULLETIN

SINGAPORE

Vol. XXVI, Part I 30th December, 1972

The Lindsaeoid Ferns of the Old World VI

Continental Asia, Japan and Taiwan

By K. U. KRAMER

Institute for Systematic Botany,

University of Utrecht, Netherlands

I. Introduction

The present revision deals with the Lindsaeoid ferns of Asia from India,

Nepal, and Ceylon in the West to China, Japan with the Bonin, Volcano, and

Ryukyu Islands, and Taiwan in the East; South it covers the parts of Further

India not dealt with in Flora Malesiana, i.e., Thailand up to the border of Malaya.

Prior to the present author’s work the knowledge of the taxonomy of the

Lindsaeoid ferns of this region was very uneven. Modern and more or less

comprehensive treatments were available for Japan (Tagawa 1937, 1938, supple-

mented by Iwatsuki 1961; Tagawa 1959; Ohwi 1965), China (Ching 1959),

Hainan (Chun, Chang & Chen 1964), Burma (Dickason 1946), and Indo-China

(Christensen & Tardieu-Blot 1936, Tardieu-Blot & Christensen 1939). No recent

or no complete treatments were available for India and the neighbouring countries,

Thailand, and Taiwan.

As shown by the synonymy and citations, the author’s views on affinity,

classification, and synonymy of the species agree well with those of Tagawa and

of Iwatsuki but diverge more or less widely from those of Ohwi, Ching, Tardieu-

Blot & Christensen, and Dickason. With regard to Ching’s work the author was

in a difficult position. It proved impossible to examine his holotypes, or photo-

graphs of them, and only in about half of the cases there were isotypes or

paratypes available which permitted to establish the identity or affinity of the

taxa, or published illustrations which to a certain extent served the same purpose.

Judging from the cases where a definite conclusion about Ching’s species could

be reached, most of them are only extreme forms of other, previously described

ones.

2 Gardens’ Bulletin, Singapore — XXVI (1972)

Due to the relative scarcity of Chinese material in herbaria outside China,

except for a few well explored localities like Hong Kong, the citations for China

are consequently very incomplete. For this reason, and because of the fluctuating

borders between certain Chinese provinces and the difficulties encountered in

attempts to locate all, often equally unstable names of places on maps, Chinese

specimens are as a rule only cited by province, although it is realized that this

does not give more than a very rough impression of the distribution of the taxa.

For India it seems also likely that several species are more widely distributed

than is shown by the specimens in the herbaria consulted by the author. It is

hoped that additional data will some time in the future be furnished by local

workers.

II. Phytogeographic Notes

The assortment of Lindsaeoid ferns in the area under consideration is very

unevenly distributed. The main centre of the group being in Malesia, the

continental areas of several more or less widespread Malesian species may be

regarded as extensions of their Malesian ones, although there is no proof that

at least in some cases the converse is not true. In the case of the taxa of section

Synaphlebium with its strong representation in Malesia and a much weaker one

on the continent this may be true, although some endemics do occur in the

“continental” part (L. venusta, L. malabarica, L. lobata var. hainaniana and

var. epirotes). Section Schizoloma, on the other hand, is so well represented in

Continental Asia, especially in Further India and southern Japan, and most

species are so much more variable there, that it seems to be the Malesian rather

than the continental part of the area that is an extension, or a secondary centre

at best. Of the 17 Asiatic species of this section 9 are confined to the continental

part, 3 to Malesia (but one extending to Australia), and 5 are common to both

areas.

There seems to be two regions in which chiefly Malesian species find their

continental limit. One is in the Malay Peninsula (the exact limits are not known

due to very incomplete exploration) where L. integra, L. malayensis, L. napaea,

L. parallelogramma, and L. doryphora stop. Another is in the central and eastern

Himalayas from Nepal to Assam, where the north-westernmost stations of

L. javanensis, L. orbiculata, and L. repens are; the two last-named species extend

to Ceylon, L. orbiculata even to S. India. There are very few examples of wide-

spread Malesian species reaching further North on the East Asiatic islands but

not on the continent: Tapeinidium pinnatum (S. Thailand, Philippines; Taiwan

and Ryukyu) and Lindsaea obtusa (Malaya, Philippines; Taiwan). Three species

extend far into China and Japan, viz. Sphenomeris chinensis, Lindsaea odorata,

and L. lucida; it would be difficult to decide whether these are originally

continental or insular species.

The only examples of notable disjunctions are L. glandulifera (Ceylon;

East Java and Lesser Sunda Islands), L. cambodgensis (Ryukyu; S. Cambodia),

and Tapeinidium pinnatum (S. India; Malesia, westernmost station in S. Thailand).

L. kawabatae (Yakushima) and L. dissectiformis (Annam and Hainan) form a

species pair.

Endemics occur in some concentration in Indo-China, sometimes extending

into South China and/or Hainan (L. austro-sinica, L. dissectiformis, L. annamensis,

L. chingii) and in Ceylon, sometimes extending into South India. (L. caudata,

L. venusta, L. schizophylla). Most, if not all, other species described as endemics

from South China by Ching are only doubtfully distinct.

The Lindsaeoid Ferns of the Old World VI 3

The species that occur furthest from tropical Asia and form the outposts

of the group as a whole are L. odorata (Cheju Do and Honshu; Nepal) and

Sphenomeris chinensis (south coast of Korea; mountains of Uttar Pradesh in

NW. india). It is not surprising that these are widespread species, either of open

habitats or euryoecious.

if. Taxonomy

KEY TO THE GENERA

(1) Sori on 1-3 (4) vein-ends; indusium laterally entirely or largely adnate to

the lamina; ultimate divisions never dimidiate; veins free; spores monolete.

(2) Ultimate free divisions not of a linear- or cuneate-divaricate type, subentire

to pinnatifid; sori on the lateral margin of the divisions or in their lobes;

pluricellular filiform paraphyses usually (always ?) present ..................

he ee A IE hah Ua OD. L2GRDS.. A834 0.18 2. Tapeinidium

(2) Ultimate free or nearly free divisions of a linear- or cuneate-divaricate

type, with the sorus (sori) on their apical margin; paraphyses 2- or 3-celled,

cS ee ae eee he et ee 1. Sphenomeris

(1) Sori on many vein-ends, or, if on 4 or fewer, the sides of the indusium free,

or the pinnules dimidiate, or the veins anastomosing, or these characters

Combined: spores with few exceptions trilete ....................... 3. Lindsaea

N.B. In order to avoid redundance, those genera and their subordinate taxa dealt

with in the author’s treatment of the group in Flora Malesiana (1971) are not

described in the following.

1. Sphenomeris

Maxon J. Wash. Ac. Sc. 3 (1913) 144, nom. cons.; Holttum Rev. Fl. Mal. 2

(1954) 340; Tagawa Col. Ill. Jap. Pterid. (1959) 54, 256; Ohwi FI. Jap. (1965) 40.

Stenoloma Fée Gen. Fil. (1852) 330, p.p. min.; Beddome Ferns Br. India

(1892) 70; Tardieu-Blot & Christensen Fl. Gen. I.-C. 7 (1939) 130, p.p.; Ching FI.

Reip. Pop. Sin. 2 (1959) 275, p.p. mai.; Chun, Chang & Chen FI. Hain. 1 (1964)

S0,..p.p.

For further synonymy and description see Fl. Mal. (gen. 1). Eleven species

in the tropical and northern subtropical parts of both hemispheres, but wanting

in Australia.

KEY TO THE SPECIES*

Rhizome scales up to 5—6-seriate at the gradually broadened base; sori uni- or

binerval; larger free ultimate divisions ca. 2 mm wide; lamina subcoriaceous or

coriaceous, usually bipinnate & pinnatifid or tripinnate & crenate ...............

a rR cas we BaD 6 igh dodE «ie + sans opens 1. Sph. biflora

Rhizome scales 1—3-seriate (or to 4-seriate at the abruptly broadened base); sori

1—3-(4-) nerval; larger free ultimate divisions 1 mm or more wide, or, if narrower,

spathulately broadened at the sorus; lamina herbaceous to subcoriaceous, in

full-grown plants bipinnate & bipinnatifid, tripinnate & pinnatifid, or even more

mnRR MEO, SOO, OMe WORE sated, STS Lv. aU, vas. - cyan SuAmeccners <4 2. Sph. chinensis

* For Sph. gracilis (Tagawa) Kurata and Sph. minutula Kurata see the notes at the end

of the genus.

4 Gardens’ Bulletin, Singapore — X XVI (1972)

1. Sphenomeris biflora (Kaulfuss) Tagawa J. Jap. Bot. 33 (1958) 203; Col. Il.

Jap. Pterid. (1959) 54, 256, fig. 89; Kramer Blumea 15 (1968) 573, 17 (1970) 162.

— Davallia biflora Kaulfuss Enum. (1824) 221. — Stenoloma biflorum (Kaulfuss)

Ching Sinensia 3 (1933) 338; Tagawa J. Jap. Bot. 22 (1948) 160; Ching Fl. Reip.

Pop. Sin. 2 (1959) 277, pl. 24 fig. 7-9. Type: Chamisso s.n., Manila, Luzon (B).

— Davallia tenuifolia Swartz var. lata Hooker ex Moore Ind. Fil. 2 (1861) 301,

based on: var. 8 of Hooker Sp. Fil. 1 (1846) 186, nom. subnud. Lectotype:

“Imp. Acad. Petersb.’’ 44, Bonin Is (K.) — Odontosoria tsoongii Ching Bull.

Fan. Mem. Inst. 1 (1930) 149. Type: several coll. cited, none seen; the collection

Tsoong 1423 from the Hailin Is, Kwangtung, China, prob. most eligible as type.

— Stenoloma littorale Tagawa Acta Phytotax. Geob. 6 (1937) 25. — Stenoloma

chusanum (L.) Ching var. littorale (Tagawa) Ito Bot. Mag. Tokyo 52 (1938) 6.

— Sphenomeris chusana (L.) Copel. var. littoralis (Tagawa) Ito ex Mizushima

Misc. Rep. Inst. Natur. Res. 38 (1955) 115 (not seen; quoted by Tagawa 1959,

loc. cit.). — Sphenomeris chinensis (L.) Maxon var. littoralis (err. ‘littorale’)

(Tagawa) Ohwi Fl. Jap. (1965) 40, comb. invalid sine cit. compl. basion. (but

possibly validly published elsewhere). Type: Tasiro s.n., Oshima, Japan (KYO,

not seen). — Sphenomeris (or Stenoloma) chinensis or chusana of various authors,

in part.

For further synonymy and description see Fl. Mal. (gen. 1 sp. 2).

Distribution. Southern Japan, S.E. coast of China, Taiwan, Marianas, northern

Philippines.

Ecology. In more or less exposed places, often by the sea, not in forests, at lower

and middle elevation.

JAPAN. Honshu: Yamada 1525 (GH); Higuchi 362 (E, L, U); Uno 7941 (GH).

— Oshima: Faurie 4607 (B); Kurihara, Kurihara & Ohba 3607 (U). — Hachijo

Shima: Suzuki 391007 (GH). — Shikoku: Tagawa 6846 (Pic-Ser), 6859 (E,

Pic-Ser, U, US); Ito 79 (SING); Iwatsuki 600 (US). — Kyushu: Ichikawa 200599

(S-PA, W); Wright sn. (GH, K, US). — Koshiki I.: Hatusima 16537 (US). —

Ryukyu. Yakushima: Koidzumi s.n. (MICH, US); Hatusima 14871 (US). —

Amami-Oshima: Iwatsuki 5046 (E, K, L, U); Hosoyamada s.n. (US). — Okinawa:

Elliott & Nakanyina 562 (US); Walker, Sonohara, Tawada & Amano 6097 (MICH).

— Bonin Is.: Mertens 68 (L); Acad. Imp. Petersb. 44 (K, lectotype of Davallia

tenuifolia var. lata). — Volcano Is. Iwo Jima: Porter 8 (BISH, MICH, US):

Henderson s.n. (GH).

TAIWAN. Tanaka & Shimada 13523 (E. GH, SING, S-PA, W); Oldham s.n.

(GH, S-PA, W); Faurie 273 (S-PA), 620 (W). Orchid I., Huang & Kao 7526 (U).

CHINA. ‘Macai Peninsula’ (prov.?): Hance 12294 (GH). — Fukien: Kuluntu I.

near Amoy, Sampson s.n. (BM, W). — Taitan I. near Amoy, Price 1382 (K). —

Kwangtung: Double I. near Swatow, Dalziel 67 (BM. E); Swatow, Dalziel s.n.

(E). — Hong Kong: Taam 1695 (L, US); Matthew 291 (E); Hance 44 (B); Chan

84 (K); Hillebrand s.n. (S-PA); Seemann s.n. (BM); Walker s.n. (BM); Lamont

1049 A (BM). — ‘Yiinnan’ (corr.??): Henry s.n. (BM).

2. Sphenomeris chinensis (L.) Maxon J. Wash. Ac. Sc. 3 (1913) 144; Ohwi FI.

Jap. (1965) 40; Kramer Acta Bot. Neerl. 15 (1967) 565; Blumea 15 (1968) 572;

Fosberg Taxon 18 (1969) 596; Kramer Blumea 17 (1970) 163. — Trichomanes

chinense L. Sp. Pl. 2 (1753) 1099. — Odontosoria chinensis (L.) J. Smith Bot. Voy.

Herald (1857) 430; Nakai J. Coll. Sc. Imp. Un. Tokyo Bot. 31 (1911) 403;

The Lindsaeoid Ferns of the Old World VI 5

Merrill Lingn. Sc. J. 5 (1927) 12. — Stenoloma chinense (L.) Beddome Ferns

Br. Ind. (1883) 70. Type: Osbeck s.n., China (S-PA). — Adiantum chusanum

L. Sp. Pl. 2 (1753) 1095. — Sphenomeris chusana (L.) Copeland Bull. Bish. Mus.

59 (1929) 69; Tagawa Col. fll. Jap. Pterid. (1959) 54, 257, fig. 88; Holttum Rev.

Fl. Mal. 2 (1954) 341. — Stenoloma chusanum (L.) Ching Sinensia 3 (1933) 337;

Tardieu-BJot & Christensen Fl. Gén. I.-C. 7 (1939) 130; Ito Fil. Jap. Ill. (1944)

pl. 13; Ching Fl. Reip. Pop. Sin. 2 (1959) 275; Chun, Chang & Chen Fl. Hain.

1 (1964) 59. Type: coll.?, China (not seen.).— Adiantum tenuifolium Lam. Encycl.

1 (1783) 44. — Davallia tenuifolia (Lam.) Swartz Schrad. J. Bot. 1800? (1801) 88;

Hope J. Bomb. Nat. Hist. Soc. 13 (1900) 35; Dun & Tutcher Kew Bull. Add.

Ser. 10 (1912) 337; Gibbs Common Hongkong Ferns (1927) 8, pl. 4. Type:

Sonnerat s.n., ‘Inde’ (P.) — Hymenophyllum ramosissimum Ham. ex D. Don Prodr.

Fl. Nepal. (1825) 12. Type: Hamilton s.n., Nilkantha, Nepal [not seen; this species

acc. to Salomon, Nomencl. Gefisskrypt. (1883) 201, and Christensen, Ind. Fil.

1906) 367]

For further synonymy and description see Fl. Mal. (gen. 1 sp. 3).

Distribution. Very widespread in the tropical and northern subtropical parts of the

Old World.

Ecology. Terrestical in open or not too shady places; apparently quite euryoecious.

a. Var. divaricata (Christ) Kramer Blumea 15 (1968) 672. — Odontosoria

chinensis (L.) J. Smith var. divaricata Christ Journal de Bot. Sér. 2, II (1909) 23.

Type: Chevallier 14309, Sao Tomé (P.) — Sph. chusana (L.) Copeland: var.

tenuifolia auct. non (Lam.) Holttum, Rev. Fl. Mal. 2 (1954) 341, fig. 198.

Characterized by the shape of the ultimate segments which are cuneate,

abruptly spathulately broadened at the sorus, slightly narrowed at the + rounded

apex, the apical margin not rarely erose, the sides often corniculate, the base

often $4 mm wide, 1-14 mm wide at the sorus; the sori are not rarely paired

in a segment, mostly uninerval, or, if binerval, on two connivent vein-ends;

spores mostly 55-60 , long.

This variety is not quite sharply distinct from the following; in its typical

form it is widespread, and often much more common than var. chinensis, in the

larger Malesian Islands. The following more or less typical specimens may be

cited from the area of the present revision:

CHINA. Yiinnan: Wang 76956 (A). — Prov.? Harland 43 (E).

INDO-CHINA: Tonkin: Balansa 105 (B, K).

THAILAND. Kerr 9656 (K, SING).

_ BURMA. Kingdon-Ward 20407 (BM).

SIKKIM. Hope (?) s.n. (L).

NEPAL. Wallich s.n. (E, K).

INDIA. Assam: Schiller 19 (B); Mann s.n. (L).

b. Var. chinensis

See Fl. Mal. (gen. 1 sp. 3, 3). Segments cuneate, gradually broadened from

the base, often about twice as long as wide, the apical margin not or scarcely

erose; sori not rarely uninerval but most often bi- or tri-, rarely to quadrinerval,

most often 3-14 mm long. Spores mostly 42-48 » long.

Distribution. Very widespread, but absent from continental Africa.

6 Gardens’ Bulletin, Singapore — XXVI (1972)

Geographically representative or widespread collections:

KOREA. Prov. South Kyéngsang: Uno 22953a (GH). — Cheju D6 (Quelpaert I):

Faurie 92 (B, BM, E, MICH), 2165 (W); Taquet 2328 (B, E), 3540 (E, K),

3684 (BM, S-PA).

JAPAN. Honshu: Tagawa 7149 (E, GH, K, L, Pic-Ser, U, US): Togashi 363

(E, K, L, U); Ohwi & Okuyama NSM 19] (B, BM, E, L, S-PA); Togasi NSM 433

(B, BM, E, K, L, MICH, S-PA, US) — Kyushu: Maximowicz s.n. (BM, SING,

S-PA, W); Schottmiiller 155 (B, S-PA); Engler 7171 (B); Ichikawa 200140 (GH).

— Goto I.: Warburg s.n. (B). — Shikoku: Tagawa 6862 (US); Beattie & Kurihara

10135 (US); Faurie 4610 (B). — Hachijo Jima: Oldham 108 (GH). — Nakashima:

Schwarz 162 (Pic-Ser). — Oshima: Faurie 4607 (B); Kurihara, Kurihara & Ohba

361 (U). — Ryukyu. Yakushima: Furuse s.n. (TOFO). Amami Oshima: /watsuki

4927 (E, K, L, U). Iheya: Suzuki s.n. (US). Okinawa: Field & Loew 23 h (SING);

Walker, Sonohara, Tawada & Amano 6052 (K, MICH, US); Conover 948, 971,

1013, 1799 (US). Iriomote: Nishida 653 (US); Walker & Tawada 6745 (US).

Yaeyama Retto: Unger’s coll. 8 (B).

TAIWAN: Tanaka & Shimada 13523 (BM, L, MICH. US); Gressitt 221] (B, BM,

K, S-PA, U); Faurie 620 (B, BM, W); Warburg 9477 (B, E), 9584, 10924 (B).

CHINA. Kiangsu: Chavel 596 (E). — Anhwei: Ching 8743 (GH, US). — Hupeh:

Wilson 2663 (B, BM, E, HBG, K, US, W); Henry 4373 (B, BM, US). — Chekiang:

Ching 1370 (B, BO, E, US), 1816 (BO, E, GH, K, US); Steward 2375 (E, K, US);

Chiao 14104 (K, US), 14543 (BRI, SING, US). — Kiangsi: Schindler 375 (B,.BM,

E, K, W); Handel-Mazzetti 213 (W). — Hunan: Fan & Li 12 (BO, L, W), 685

(A, BO, L); Tsang 23756 (A, L). — Szechuan: Chow 4668 (E, US); Fang 2108

(E, GH, K, US), 3818, 4671 (E, K), 5795 (E, GH, K, US), 9931 (E, SING);

Wilson 5307 (BM). — Kweichow: Cavalerie 1373 (E); Tsiang 6256 (GH); DeVol

563 (GH). — Fukien: Chung 2/37 (BO, K), 2857 (E, GH, K, W); Tang 6916

(BM, MICH), 7/01 (SING, S-PA); Norton 1081, 1082 1083, 1084 (US); Metcalf

& Tang 543 (US), 6097 (BM). — Kwangtung: Tsang 16574 (S-PA, US), 20001

(K, MICH), 20579 (BO; K, MICH, W), 20708 (B, BO, K, L, MICH, SING, W),

21243 (A, K, MICH, S-PA), 2620] (A, MICH); Lau 684 (GH, MICH), 20135

(K, US); Levine 511 (BO, GH); Gressitt 1235, 1314 (BM, E). — Kwangsi: Tsang

23069 (GH), 28175 (A, SING, US). — Yiinnan: Rock 7698 (GH, K, US); Wang

66809, 71825, 71871, 71962, 73922, 74146, 77198 (A); Handel-Mazzetti 5935

(K, US, W); Forrest 18840 (BM, E, K), 26019 (E, K); Henry 9018 A (E, K, US);

Cavalerie 1373, 4023 (E, K); Maire 2744 (E, K). — Tibet: Ludlow, Sheriff &

Taylor 6728 (BM, MICH). — Hainan: Tsang 16778 (B, BO, K, US); Gressitt

1030 (E, GH); Eryl Smith 1452 (BM, K), 1453 (K, SING, S-PA, US). — Hong

Kong: Fortune 16 (B, E, L); Taam 1336 (US), 1448 (BO, US), 2023 (MICH US);

Jelinek 31 (B, W); Palmer & Bryant 41 (US); Eryl Smith 1456 (SING). — Macau:

Warburg 5490 (B). :

INDO-CHINA. Tonkin: Tsang 27256, 30152 (K); Chevalier 29327, 29495 (SING).

— Laos: Poilane 16926 (K). — Annam: Clemens 3565 (BM, VU); Poilane 1617

(K, SING). — Cochin-China: Gaudichaud s.n. (B).

THAILAND. Eryl Smith 585 (BM, K, SING), 398, 1214, 2292 (K); Smitinand 417

(K); Tagawa, Iwatsuki & Fukuoka T 588 (L, U), T 1273 (UV).

BURMA. Dickason 6721 (A, E), 9105 (A, E, L); Rock 2078 (US), 2152 (S-PA,

US); Forrest 52394 (K); McKee 6018 (BM).

The Lindsaeoid Ferns of the Old World VI |

BANGLA DESH. Wallich s.n. (K).

BHUTAN. Griffith s.n. (B, K).

SIKKIM. Meebold 2224 (B); Hope s.n. (K, L, US); Gamble 7019 (K); Hooker s.n.

(GH, B); Engler 3543 (B).

NEPAL. Stainton, Sykes & Williams 5189 (BM, E, GH, Pic-Ser), 8977 (BM, E);

Fleming 893, 905, 915 (BM).

INDIA. Assam: Mann s.n. (BO, E, HBG, L, SING, S-PA, US); Schlagintweit s.n.

(B, BM, S-PA); Hooker & Thomson s.n. (B, BM, E, GH, K, S-PA, U, W);

Watt 10322 (L, US); N.E.F.A., Panigrahi 17045 (K). — Manipur: Watt 6050

(B, E). — West Bengal: Schelpe 3670 (BM); Bir s.n. (U, US); Warburg 989 (B);

Gamble 6856 A, 7325 (K), 7307 (E, K). — Bihar: Haines 652 (K); Mooney 138

(K.) — Uttar Pradesh: Kumaon, Strachey & Winterbottom 2 (BM, GH, K):

Stewart 320 (BM). — Himachal Pradesh: Bliss 95, 168 (K); Trotter 255 (K). —

Orissa: Mooney 695, 3869 (K). — Pondicherry: Perrottet 591 (B, L), 1397 (B).

— Madhya Pradesh: Mooney 1296 (K). — Madras: Perrottet 591, 1171, 1172

(W); Bourne 4843, 4998, 5139 (K); Hohenacker-exsicc. 1260 (B, BM, E, L, S-PA,

W); Thomson s.n. (B, BM, E, MICH, SING, S-PA, U, US, W); Hiigel 3176 (W);

Engler 3636, 3641 (B). -— Mysore: Meebold 9621 (B); Blanford s.n. (US). —-

Kerala: Wallich ‘245’ (US).

CEYLON; Thwaites CP 983 (B, BM, BO, E, SING, S-PA, W); Pfaeltzer &

Abeyguna Wardena 35 (U); Freeman 17, 42 (BM); Gardner 20 (W); Schmid 1039

(BM): Naylor Beckett 202 (B, E, K).

Not seen from the Andaman and Nicobar Islands.

DOUBTFUL AND INSUFFICIENTLY KNOWN SPECIES

Sphenomeris gracilis (Tagawa) Kurata J. Geob. 13 (1965) 101.— Stenoloma gracile Tagawa

Acta Phytotax, Geob. 6 (1937) 227. Type: Koidzumi s.n., Iriomote, Ryukyu (KYO, not seen);

one other collection cited from the same island.

No authentic material seen. Judging from the description it seems to fall within the

variability of Sph. chinensis, except for the arrangement of the leaves with are described as

subremote. The author compared it with Stenoloma eberhardtii, in the present author’s

opinion a Lindsaea (L. dissectiformis, see below), but the description of the scales of St.

gracile shows that it is a Sphenomeris and not related to the group of L. dissectiformis.

Sphenomeris minutula Kurata J. Geob. 13 (1965) 101. Type: M. Saté 4243, Amami Oshima,

Ryukyu (TOFO, not seen).

a The original description, in a journal which is perhaps not very widespread, is as

ollows:

Rhizoma repens, ca. 1 mm crassum, squamis setaceis fuscobrunneis cylindricis

articulatis ca. 1 mm longis dense obtectum; frondibus subremotis. Stipites straminei

deorsum subcastanei, glabri, graciles, 5-10 mm longi, supra sulcati. Lamina ovata

vel oblongo-ovata, apice in ambitu obtusa, 1-1.5 cm longa, 0.7-1 cm lata, bipinnata

vel tripinnatifida, chartacea, utrinque glabra; rachi gracili leviter flexuosa; pinnis

2-4 jugis, ascendentibus, alternatis, inferioribus oblique flabelliformibus, usque ad

5 mm longis 5 mm latis, costis supra late sulcatis, segmentis ultimis cuneatis apice

dilatatis truncatis, 1.5-2 mm longis 0.5-1 mm latis, venulis in segmentis ultimis 1-2.

Sori marginales, indusiis submembranaceis subintegris, pallide griceis [sic] interdum

brunnescentibus, ca. 0.5 mm longis, 0.3-0.9 mm latis...This is the tiniest species of

the genus Sphenomeris. Notwithstanding the minuteness, it + a mes produces

perfectly developed sori on the terminal margin of ultimate lobes.

Through Prof. Kurata’s courtesy a specimen identified by him as bveleattnns to this

species was deposited in the Utrecht herbarium: Sako 5214, Amami Oshima. I am not

satisfied that it is specifically distinct from Sph. chinensis, nor can I at present decide its

status. It may be a juvenile, yet fertile form of that species; the specimens were collected

on mossy rocks, like L. odorata var. japonica, which I think is also a permanently (pheno-

typically or genotypically ?) juvenile form.

8 Gardens’ Bulletin, Singapore — XXVI (1972)

2. Tapeinidium

(Presl) C. Christensen Ind. Fil. (1906) 631; Tardieu-Blot & Christensen Fl. Gén.

I.-C. 7 (1939) 133; Ching Fl. Reip. Pop. Sin. 2 (1959) 278; Kramer Blumea 15

(1968) 545.

For further synonymy and description see Fl. Mal. (gen. 2).

A genus of seventeen species, almost confined to the Malesian-Melanesian

area.

KEY TO THE SPECIES

(1) Petiole concolorous, usually quite pale.

(2) Lamina pinnate & pinnatifid or more dissected ......... 2. T. luzonicum

(2) Lamina simply pinnate, with serrate or crenate pinnae .....................

1. Tapeinidium gracile (Blume) v.A.v.R. Handb. (1909) 315; Kramer Blumea

15 (1968) 551. — Davallia gracilis Blume En. Pl. Jav. (1828) 233. Type: Blume

1731 or s.n., Java (L). — ? Tapeinidium lineare (Cav.) C. Chr. Dansk Bot. Ark.

9 (1937) 26; Tardieu-Blot & Christensen Fl. Gén. I.-C. 7 (1939) 133. — Dicksonia

linearis Cavanilles Descr. (1802) 274. Type: Née s.n., Philippines (MA, not seen;

photogr. in U).

For notes on the synonymy see Kramer, loc. cit., and for the description

Fl. Mal. (gen. 2 sp. 8).

A West Malesian species extending to the Moluccas, known from the present

area by two collections.

TAIWAN. Orchid I, Huang & Kao 7525 (U).

INDO-CHINA. Annam: Nhatrang, Poilane 3400 (BM, K, P).

2. Tapeinidium luzonicum (Hooker) Kramer Blumea 15 (1968) 552. — Davallia

luzonica Hooker Sp. Fil. 1 (1846) 174, pl. 60 B 2, 3, 5. Type: Cuming 139,

Luzon (isotypes B, L). — T. biserratum auct. non (Blume) v.A.v.R.; Holttum

Rev. Fl. Mal. 2 (1954) 339, fig. 197.

For further synonymy and description see Fl. Mal. (gen. 2 sp. 11).

A West and Central Malesian species recently collected in southern Peninsular

Thailand.

THAILAND. Tagawa, Iwatsuki & Fukuoka T 4830 (U, US).

3. Tapeinidium pinnatum (Cav.) C. Christensen Ind. Fil. (1906) 631; Ogata

Ic. Fil. Jap. 1 (1928) pl. 44; Ito Fil. Jap. Ill. (1944) pl. 15; Holttum Rev. FI.

Mal. 2 (1954) 339, fig. 196; Ching Fl. Reip. Pop. Sin. 2 (1959) 278, pl. 25 fig. 7-11;

Kramer Blumea 15 (1968) 553. — Davallia pinnata Cavanilles Descr. (1802) 277;

Hooker Sp. Fil. 1 (1846) 173, pl. 60 B. Type: Née s.n., Philippines (MA, not

seen; photogr. U).

For further synonymy and description see Fl. Mal. (gen. 2 sp. 12).

Western and Central Malesia, extending sporadically to Continental Asia

and Japan. Reports from elsewhere are due to confusion with other species.

The Lindsaeoid Ferns of the Old World V1 9

JAPAN. Ryukyu: Okinawa, Amano 6297 (US). Ishigaki, Masamune & Suzuki s.n.

(US); Tagawa & Iwatsuki 4480 (US); Nishida 447 (US); Fosberg 37391 (L, US).

Iriomote, Tagawa & Iwatsuki 4566 (K, L, U, US), 4657 (US); Walker & Tawada

6705 (BISH, K, MICH, US); Ogata 273 (BM); Oka 13751 (TOFO).

TAIWAN. Tagawa 990 (K).

THAILAND. Eryl Smith 1656 (BM), 1566, 1912, 1913, 2190 (K); Marcan 1241

(SING), 1332 (BM, SING); Kerr 6803, 9299 (K); Smitinand 2164 (K); Murton 8

(K).

INDIA. Kerala: Palghat Hills, Gen. Johnston s.n. (K).

The single old collection from India, which I did not find cited in the

literature, and the absence of the species from Ceylon are quite remarkable.

3. Lindsaea

Dryander in J. E. Smith Mém. Ac. Turin 5 (1793) 401; Trans. Linn. Soc. 3

(1797) 39; Beddome Ferns S. India (1863/64; 1873) 39; Handb. Ferns Br. India

(1892) 72; Tagawa Acta Phytotax. Geob. 6 (1937) 24; Tardieu-Blot & Christensen

Fl. Gén. I.-C. 7 (1939) 118; Holttum Rev. Fl. Mal. 2 (1954) 321; Ching FI. Reip.

Pop. Sin. 2 (1959) 257; Tagawa Col. Ill. Jap. Pteridoph. (1959) 52, 226; Ohwi

Fl. Jap. (1965) 39; Kramer Blumea 15 (1968) 557. — Schizoloma Gaudichaud

Ann. Sc. Nat. 3 (1824) 507; p.p.; Beddome loc. cit. (1863/64, 1873) 9; loc. cit.

(1892) 77, p.p.; Tardieu-Blot & Christensen loc. cit. (1939) 128; Holttum loc. cit.

(1954) 342; Ching loc. cit. (1959) 272. — Stenoloma Fée sensu Tardieu-Blot &

Christensen loc. cit. (1939) 130, p.p.; Ching loc. cit. (1959) 275 p.p.

For further synonymy and description see Fl. Mal. (gen 4).

About 150 species, pantropic and subtropic. For the subdivision of the genus

see Kramer (1968, loc. cit. and Fl. Mal.).

KEY TO THE SPECIES*

(1) Lamina bipinnate, the secondary rachises at least in the adaxial groove with

short pubescence (SECT. Aulacolindsaea) ........................ 25. L. caudata

(1) Secondary rachis glabrous; or lamina not bipinnate.

(2) Rhizome epiphytic, long-scandent, with broad, triangular scales and

strongly dorsiventral stele; leaves remote (SUBGEN. Odontoloma).

(3) Lamina of full-grown plants bipinnate .................. 35. L. parasitica

(3) Lamina simply pinnate.

(4) Sori continuous; rhizome scales chocolate brown ..................

Be Aegan Pub shan ies addos exh MEE Ye Ae raeee tna 34. L. oblanceolata

(4) Sori interrupted; rhizome scales golden brown.

(5) At least some of the fertile lobes of the pinnules erose-

SD aD, 6 Bee areal nc ines ncaa: 33. L. merrillii

(5) Fully fertile lobes of pinnules not erose-denticulate.

*The reader is reminded that in the present author’s terminology the term pinnule

always designates a free ultimate segment, regardless of the degree of dissection of the

amina.

Gardens’ Bulletin, Singapore — XXVI (1972)

(6) Larger pinnules 10-12 mm long; pinnules incised to 4 or 3,

the lobes evenly narrowed from base to apex ............

}. CUES, behaweRtde) DO, DR 31. L. glandulifera

(6) Larger pinnules 15 mm or more long, with truncate lobes,

much less deeply incised, the lobes about parallel-sided

ee TEE ne Te oe eee ee 2 ee 32. L. repens

(2) Rhizome terrestrial, mostly short-creeping and with clustered leaves,

rarely more long-creeping and with remote leaves, but then the scales

narrow; stele radially symmetric or nearly so (SUBGEN. Lindsaea).

(7) Lamina bipinnate or more dissected, with gradually reduced upper

(primary) pinnae; or at least the basal pinnules of the almost conform

terminal pinna enlarged, lobed (SECT. Schizoloma). (p. 12)

(8) Veins scantily to freely anastomosing ......... 13. L. heterophylla

(8) Veins quite free.

(9) Pinnules not or hardly dimidiate, deeply incised, consisting

of cuneate or flabellate, ‘+ divaricate lobes.

(10) Rhizome not very short-creeping, the petioles not

clustered, several mm apart; sori various.

(11) Sori on 1 or 2 veins; all ultimate lobes linear-cuneate

#o..Gok apemeteidt toll aal 11. L. schizophylla

(11) Sori rarely on 1, mostly on 2-5 veins; very few or

no ultimate lobés''linear-cuneate’'2..27...2 Rie

Ji)... oo... aeeteeeoreb: Loe... vane 12. L. cambodgensis

(10) Rhizome very short-creeping, with clustered petioles; sori

uni- to quadrinerval.

(12) The greater part of the lamina fully tripinnate;

ultimate lobes abruptly spathulately broadened at

the sorus, there much broader than the wings that

CONNEGL, ANGI Nib o. wstiug dhe « BT oy a a 1. L. dissectiformis

(12) Lamina fully tripinnate only at the base; ultimate

lobes gradually broadened to the sorus, there not

much wider than the wings connecting them, or

scarcely broadened.

(13) Sori on 1 or 2 veins; larger ultimate lobes 1 mm

wide at the apex (fig. 4) ...... 2. L. kawabatae

(13) Sori on 1-4 veins; larger ultimate lobes 3-5 mm

wide at the apex (fig. 5) ..... 3. LL. annamensis

(9) Pinnules distinctly dimidiate, or, if only indistinctly or not at

all so, not deeply dissected into cuneate or flabellate lobes.

(14) Larger, dimidate pinnules incised to the middle or

beyond, the lobes not broadened at the sorus and not

apreulate viii. t2. A200. ae A ne see 10.

(14) Larger, dimidiate pinnules incised to 4, occasionally to

the middle; some or most lobes broadened at the sorus

and apiculate (Hg. 0) i...-..0--.-:ss2-uneaeeeeee 9. L. chingii

The Lindsaeoid Ferns of the Old World VI 11

(14) Larger, dimidiate pinnules not or much more shallowly

incised.

(15) Petiole and basal half of the primary rachis abaxially

Cos iat OM) ST a ee 7. L. austro-sinica

(15) Petiole and basal half of the rachis abaxially

bi-angular, often pale-margined.

(16) Sori strongly interrupted, even though the

incisions of the margin do not, or some just

barely, reach the level of the receptacle;

indusium reaching and often here and there

Surpassing the margin; receptacle sometimes

seemingly decurrent onto some of the veins

supporting it; texture often subcoriaceous

(16) Sori less strongly interrupted, or some or all

incisions reaching to beyond the receptacle,

or the indusium remote from the margin.

(17) Lamina simply pinnate, with suborbicular-

flabellate, scarcely or not incised larger

pinnules, or bipinnate, the unipinnate

apical portion relatively very long, at the

base with suborbicular-flabellate pinnules

Lg. Ie ee oe 5. L. orbiculata

(17) Pinnules not suborbicular-flabellate, or,

if some approach this shape, the lamina

bipinnate with the unipinnate leaf-apex

relatively less predominant.

(18) Terminal pinnule of lateral pinnae

very large, asymmetrically deltoid,

much larger than any of the

(paucijugate) lateral pinnules of the

same pinna ......... 8. L. javanensis

(18) Terminal pinnule or segment of

lateral pinnae lanceolate, rhombic,

subelliptic, or, if deltoid, in size

comparable to the larger lateral

pinnules in the same pinna; or

lamina simply pinnate.

(19) Pinnules in the basal portion

of the simply pinnate leaf-apex

(but above those that are

transitional in shape between

pinnate pinnae and non-pinnate

pinnules) rhombic, with + pro-

tracted apex, their sori broken

by incisions reaching consi-

derably beyond the receptacle.

Gardens’ Bulletin, Singapore — X XVI (1972)

(20) Terminal pinnule/segment

large, well-developed, free

or nearly so; no upper

pinnules of lateral pinnae

(if any) so strongly reduced

as to be denticuliform;

indusium mostly close to ~

the margin; lamina not ~

rarely unipinnate ........... :

eerearct ord 8. L. javanensis

(20) Terminal segment nar-

rowly triangular, in lateral

pinnae (if any) mostly at

the base confluent with

some denticuliform re-

duced upper pinnules;

pinnule-lobes (except

sometimes the outer ones)

with little or not convex

outer margin; indusium

strongly intramarginal;

rarely unipinnate (fig. 7)

eee ee 4. L. chienii

(20) Terminal segment as in the

preceding species; lobes of

fertile pinnules convex on

the outer margin; indusium

almost or quite reaching

the margin; bipinnate,

rarely more dissected ......

Mis teaver 10. L. bouillodii

(19) Pinnules in the basal portion of

the leaf-apex (but above those

that are transitional from

pinnate pinnae to non-pinnate

pinnules) _ parallelogrammoid,

rectangular, or flabellate, not

rhombic with protracted apex,

their sori continuous or with

very few incisions that scarcely

reach beyond the receptacle or

do not reach its level (fig. 8)

AMR, Ah: 5. L. orbiculata

(7) Lamina simple, simply pinnate, or, if bipinnate, without gradual

transition from pinnae to pinnules, and with conform terminal pinna

without larger, lobed basal pinnules.

The Lindsaeoid Ferns of the Old World VI 13

(21) Lamina simply pinnate (rarely simple), with non-dimidiate,

lanceolate or linear pinnules, with conform terminal one, and

oe 0 ne 14. L. ensifolia

(21) Lamina simply pinnate, with lanceolate pinnules, without free

conform terminal one, and with irregularly reticulate venation

eT fe" ee 13. L. heterophylla

(21) Lamina simply pinnate with free veins or bipinnate with dimidiate

pinnules, or simply pinnate with reticulate veins and dimidiate

pinnules.

(22) Veins sparingly to copiously anastomosing (SECT. Synaphle-

bium). (p. 15)

(23) Veins of larger fertile pinnules irregularly anastomosing,

sometimes in some of the pinnules nearly or quite free.

(24) Sori continuous, or, if interrupted, the incisions

not deeper than 1 mm.

(25) Sori continuous or interrupted; pinnules

10-12 mm long, 3-4 mm wide; petiole pale,

abaxially flat or convex, bi-angular, the

angles evanescing downward; sterile pinnules

broadly crenate-sinuate ...... 16 L. napaea

(25) Sori interrupted; pinnules 10-20 mm long,

4—6 min wide; petiole pale, abaxially angular

to the base, sulcate at least near the apex;

serie ipinnglesbicrenate ......3.. We.

(24) Sori interrupted, the larger incisions 2 mm deep

WE CORN ts. escapee. cuss sia 17. L. malabarica

(23) Veins of larger fertile pinnules regularly anastomosing,

at least in the basal 2.

(26) Sori of larger, fully fertile pinnules continuous.

(27) Pinnules twice as long as wide; petiole usually

POUGiy DEW iss. sca deva vanes 21. L. integra

(27) Pinnules 24-3 X as long as wide; petiole

SEPAUIICOUS ooo oe eee ee ns 20. L. cultrata

(26) Sori of larger, fully fertile pinnules interrupted

by incisions of the margin.

(28) Incisions of pinnules going to or slightly

beyond the level of the receptacle; outer or

all sori short, on 1-3 vein-ends; indusium

falling short of the margin by less than its

width to slightly surpassing it (fig. 2) ......

Tee ae es MIP E NE eb eT ee viecew lens 23. L. venusta

Gardens’ Bulletin, Singapore — XXVI (1972)

(28) At least most incisions of the pinnules deeper;

sori usually on more than 3 veins, and/or the

indusium more strongly intramarginal.

(29) Pinnules opaque, hardly narrowed to the

obliquely truncate apex; outer margin

distinct, with an incision; pinnae rather

abruptly narrowed below the + caudate,

pinnatifid apex; pinnule-bearing

rachises abaxially brown, sulcate and

Gale-Mareined «2.20.2 <i.0ssas-2ne

ic iacii etd est 19. L. parallelogramma

(29) Pinnules with subacute or rounded apex

and/or distinctly narrowed to the apex;

or, if truncate, the outer margin not

incised; pinnules often translucent;

pinnae more gradually narrowed;

pinnule-bearing rachises various.

(30) All pinnule lobes and receptacles

distinctly convex towards the

margin; indusium reaching to the

margin or nearly so (fig. 1) ......

ee eee ee 22. L. lobata

(30) These characters not combined, at

least the inner pinnule lobes with

straight outer margin.

(31) Pinnules with a distinct outer

margin joining the upper at

an angle of less than 90°, its

sorus sometimes continuous

with that of the upper; at

least the inner incisions reach-

ing considerably beyond the

level of the receptacle ......

Peis dose ees 20. L. cultrata

(31) Pinnules without a distinct

outer margin, this rounded

into the upper; most or all

incisions reaching consider-

ably beyond the level of the

receptacle .... 18. L. obtusa

(31) Pinnules without a distinct

outer margin, this rounded

into the upper; incisions

reaching to the level of the

receptacle, or shallower ......

“sis ioe eka eee 16. L. napaea

The Lindsaeoid Ferns of the Old World VI LS

(22) Veins quite free.

(32) Pinnules not dimidiate.

(33) Pinnules at least 6 X as long as wide; rachis

dark-sclerotic, abaxially terete.

(34) Pinnules articulate at their insertion .........

PL). 0. Sa. JOU aad... 29. L. divergens

(34) Pinnules not articulate at their insertion ...

eM REET Accs hee te recs’ 28. L. walkerae

(33) Pinnules relatively much shorter; rachis abaxially

bi-angular, dark or not. see 7 (SECT. Schizoloma).

(32) Pinnules dimidiate.

(35) Rachis abaxially terete, except near the apex.

(36) Pinnules incised, or, if entire, less than 12 mm

long; simply pinnate; spores monolete.

(37) Pinnules herbaceous to subcoriaceous,

without a distinct outer margin; rachis

not dark-sclerotic ...... 26. L. odorata

(37) Pinnules subcoriaceous or coriaceous,

with a distinct outer margin; at least

the basal half of the rachis dark-

sclerotic (fig. 3) ...... 27. L. himalaica

(36) Pinnules entire (except if incompletely fertile),

usually 15-20 (—35) mm long; spores trilete;

simply pinnate or bipinnate ..................

D. MEMES, We, Sew 2... one 24. L. doryphora

(35) Rachis abaxially bi-angular.

(38) Pinnules erose; indusium 0.5 mm wide ......

So eee, es a 5. L. orbiculata

(38) Pinnules not erose; indusium narrower.

(39) Lamina simply pinnate, with reduced

and somewhat remote basal pinnules;

most incisions of the pinnules reaching

to about the level of the receptacle ...

Beebe Eonar eee 30. L. lucida

(39) Lamina bipinnate, or, if simply pinnate,

the basal pinnules not reduced and little

Or not remote; incisions of pinnules

reaching considerably beyond the level

of the receptacle ... 17. L. malabarica

16 Gardens’ Bulletin, Singapore — X XVI (1972)

SECTION Schizoloma (Gaud.) Kramer

1. Lindsaea dissectiformis Ching Sinensia 1 (1930) 52. Type: McClure 18312,

Hung Mo Shan, Hainan (isotype A). — Odontosoria eberhardtii Christ Journal

de Bot. 21 (1908) 235, 266, nom. nud.; Merrill Lingn. Sc. J. 5 (1927) 12. —

Stenoloma eberhardtii Ching Sinensia 3 (1933) 338; Tardieu-Blot & Christensen

Fl. Gén. I.-C, 7 (1939) 132, fig. 16 1/2; Ching Fl. Reip. Pop. Sin. 2 (1959) 277,

pl. 24 fig. 1/2; Chun, Chang & Chen FI. Hain. 1 (1964) 58, fig. 26. — Lindsaea

eberhardtii Kramer Acta Bot. Neerl. 6 (1957) 135. Type: Eberhardt 115, 116,

Annam (P).

Rhizome very short-creeping, 1-14 mm in diam.; scales castaneous, very

narrowly triangular, the cell partitions laterally somewhat protruding, almost

the whole apical half uniseriate, to 6-seriate at the base, to 24 mm long. Leaves

clustered; petioles ca. 10-25 cm long, slightly exceeding the lamina to 14 X as

long, castaneous to atropurpureous, scarcely lustrous, quadrangular with subterete

base, upward pale-margined. Lamina deltoid to oblong, acute, ca. 8-15 cm long,

at the base tri- or quadripinnate & pinnatifid, upward gradually of simpler

structure, with up to 10 primary pinnae to a side that are more than once

pinnate & pinnatifid; colour dark or olivaceous green when dry, texture herbaceous.

Primary rachis like the petiole, upward gradually stramineous. Primary pinnae

slightly ascending, contiguous, the largest (basal) ones 4-7 cm long, 24-34 cm

wide; upper pinnae gradually shorter; at least the basal pinnae distinctly inequila-

teral, basally broader; secondary rachises abaxially narrowly sulcate, bi-angular at

the base, upward gradually green-marginate. Secondary pinnae up to 10 to a side,

slightly ascending. Penultimate divisions ascending, variously once or twice bifid,

the ultimate lobes obcuneate-spathulate, uni- or the largest binerval, 3-5 mm

long if almost free, the base ca. 0.4-0.5 mm wide, upward gradually cuneately,

then at the sorus abruptly spathulate-broadened, there 4-14 mm wide, their apices

triangular-acute, obliquely truncate, or rarely rounded, in addition erose-denticulate.

Upper lobes/segments gradually confluent into pinnatifid pinna-apices. Sterile

segments narrower, lanceolate, acute. Veins immersed, scarcely evident. Sori uni-

or less often binerval; indusium pale, 4-2 mm long, not rarely not at right angles

with its vein, its base straight or faintly concave or convex, laterally free, its

outer edge sinuate-erose, 0.4-0.5 mm wide, not reaching the margin by an equal

or often considerably larger distance, rarely by only about half its width. Spores

light yellowish brown, trilete, verruculose, ca. 27 up.

Ecology: In moist forests. ca. 1500 m alt.; very few data.

Distribution: Confined to Annam (Central Viet-Nam) and Hainan.

INDO-CHINA. Annam: Eberhardt 115, 116 (P, type of S. eberhardtii); Poilane

3692 (BM, BO, K, MICH), 4156 (SING), 4360 (K, P), 22047 (GH); Jacquet s.n.

(BO, P).

CHINA. Hainan: Eryl Smith 1457 (BM, K); McClure 778 (Lingnan Univ. 1831/2)

(A, isotype).

Notes. Closely related to the Japanese L. kawabatae and the Madagascan

L. millefolium; see also Kramer (1957, loc. cit.). The basionym of Stenoloma

eberhardtii, the name most often used for this species, was published as a nomen

nudum and the first description did not appear until 1939 in the Flore Générale

de l’Indo-Chine. Therefore Ching’s name L. dissectiformis takes precedence.

The Lindsaeoid Ferns of the Old World VI

gq ah

7 C17?

BEE?

soe

a 7

Vises

Figs. 1-8. LINDSAEA

I. Lindsaea lobata var. hainaniana. Detail of pinna (Liang 63789, US). 2. L. venusta.

Detail of pinna (Wall 20.37, GH). 3. L. himalaica. Detail of pinna (Griffith s.n.,

Assam, K). 4. L. kawabatae. Basal pinna (holotype KYO). 5. L. annamensis. Lower

pinna (Poilane 3567, P). 6. L. chingii. Basal pinna (Tsang 29330, P). 7. L. chieniti.

Lamina apex (Kurata s.n., Honshu, TOFO). 8. L. orbiculata var. commixta.

Lamina apex (Kawabata 807, TOFO).

18 Gardens’ Bulletin, Singapore — X XVI (1972)

2. Lindsaea kawabatae Kurata J. Geobot. 13 (1965) 100. — Type: Kawabata

991, Yakushima, Japan (TOFO). Fig. 4

As the journal in which this species was described is not likely to be found

in many smaller libraries, the original description is here quoted verbatim.

Rhizoma breviter repens, squamis minutis lineari-lanceolatis dilute castaneis

obtectum; frondibus approximatis. Stipites graciles, usque ad 14 cm _ longi,

rufo-castanei vel sursum brunneo-straminei, quadrangulares, supra sulcati. Lamina

8-11 cm longa, basi 5-9 cm lata, deltoidea vel ovata, acuminata, tripinnata

sursum bipinnata vel simpliciter pinnata; pinnis ca 10-jugis, erecto-patentibus,

breviter petiolatis, alternatis vel inferioribus suboppositis, inferioribus inter se

1-2 cm remotis, deltoideo-lanceolatis, acuminatis; pinnulis 5-7 jugis, distincte

anadromice dispositis, infimis posterioribus longioribus, subdimidiato-lanceolatis,

ad 1.2 cm longis 5 mm latis, pinnatis, segmentis 2—3-jugis, anguste cuneatis ad

3 mm longis, ad 1.5 mm _ latis, apice acute denticulatis, segmentis infimis

anterioribus flabellato-cuneatis, pauci-incisis, pinnulis superioribus et anterioribus

omnibus dimidiato-rhomboideis vel oblique flabellato-cuneatis, venulis in segmentis

ultimis 1—2; textura herbacea, viridi. Sorus unus pro lobulo ultimo, venas 1-2

Occupans, indusio griceo [sic] e margine magis remoto.

To this good description the following points may be added: Major, bipinnate

pinnae 3 to a side, the upper ones very gradually of simpler structure, the

uppermost confluent into a _ pinnatifid leaf-apex. Primary rachis abaxially

bi-angular like the petiole; secondary rachises abaxially narrowly sulcate, narrowly

green-marginate. Ultimate divisions apically erose or denticulate, the narrower

with convex outer margin, the broader ones sometimes subtruncate. Sori 1-1 mm

long, the longer ones basally concave; indusium pale, delicate, + erose, laterally

free, 0.3-0.4 mm wide, not reaching the margin by an approximately equal distance,

not reflexed at maturity. Spores as in L.. chienii, g.v. Only known from the type

collection; no ecological data.

Notes. As already noted by Kurata, this species is very closely related to

L. dissectiformis (“‘eberhardtii’’), with which it shares i.a. the inequilateral, basitonic

basal pmnae. For the differences see the key; more material may show the two

species to be one.

3. Lindsaea annamensis Kramer, spec. nov. Fig. 5

Rhizoma breviter repens, squamis ferrugineis, angustissime triangularibus,

acuminatissimis obtectum. Folia approximata; petiolus badius, quadrangularis.

Lamina triangularis, basi bipinnata et pinnatifida vel tripinnata, chartacea vel

subcoriacea. Pinnae majores pro latere 6-10, adscendentes, triangulares, acutae

vel acuminatae. Pinnulae majores pro latere 3-6, adscendentes, + profunde

pinnatifidae, acroscopice segmentis 1-3, basiscopice segmento unico vel nullo; lobi

ultimi valde adscendentes, cuneato-flabellati, majores bifidi, minores simplices,

usque ad 8 X 5 mm, basi ca 1 mm lati, apice eroso-denticulati. Apices laminae

pinnarumque sensim angustati et structura simpliciore, pinnatifidi; saepe acuminati

vel caudati. Venae binae ad quaternae vel in lobis minoribus singulae. Sori venis

1-4 insidentes, majores basi saepe concavi. Indusium marginem non attingens.

Sporae tetraédricae.

Rhizome short-creeping, ca. 24 mm in diam.; scales reddish brown, very

narrowly acuminate-triangular, with a considerable uniseriate apical portion, to

6-seriate at the base, to 24 mm long. Leaves close; petioles castaneous, somewhat

shining, quadrangular, upward + pale-margined, adaxially sulcate, 10-30 cm long,

The Lindsaeoid Ferns of the Old World VI 19

equaling to almost twice as long as the lamina. Lamina ca 10-22 cm long,

triangular, bipinnate & pinnatifid at the base, or in large leaves there tripinnate

and otherwise bipinnate & pinnatifid; colour medium green or olivaceous when

dry, texture chartaceous to subcoriaceous. Rachis like the upper part of the petioie,

upward gradually green. Major pinnae 6-10 to a side, (subfalcately) ascending, con-

tiguous, triangular, acute or acumimate, subsessile, to 8 XxX 3 cm; secondary

rachises stramineous, abaxially shallowly sulcate, upward gradually narrowly

green-marginate. Pinnules ascending, ‘+ continguous, 3-6 larger ones to a side;

larger pinnules asymmetrically ovate to trapeziform in outline, ca. 15-20 xX

5-7 mm, + deeply pinnatifid, with 1-3 segments on the acroscopic side, the

largest also with one on the basiscopic side, in the largest incised to a narrow

costal wing; basal pinnae sometimes with 1 or a few basiscopic fully pinnate

& pinnatifid pinnules. Ultimate lobes strongly ascending, cuneate-flabellate, the

larger ones bifid, the smaller ones entire, the largest ca 8 X 5 mm, ca 1 mm

wide at the base, the apical margin protracted into 1 or several apiculi

separated by incisions, or in broader lobes coarsely erose. Upper pinnae, pinnules,

etc., gradually and strongly reduced, confluent into narrow, lobed, lanceolate,

often acuminate or caudate lamina and pinna apices. Veins immersed, evident,

2-4 per lobe, single in the smailest. Sori on 1-4 vein-ends, usually basally concave

if long; receptacle often laterally exceeding its veins, not rarely not at right

angles with them; indusium delicate, pale, subentire to erose, laterally free, 4 mm

wide, falling short of the margin by an at least equal, often larger distance.

Spores medium brown, trilete, slightly verruculose, ca. 25-28 uw.

Type and only known collection: Poilane 3567, Nhatrang, Annam, Viet-Nam

(P, 2 sheets), on poor sandy soil in forest, 1800 m. “‘Bia di’’ (Mois).

Note. This species combines characters of L. chingii, L. dissectiformis, and

L. chienii but is probably closest to the last-named species. Its spores seem to be

normally developed, but it might, of course, still have originally arisen by

hybridization.

4. Lindsaea chienii Ching Sinensia 1 (1929) 4; Hu & Ching Ic. Fil. Sin. 1 (1930)

pl. 19; Tagawa Acta Phytotax. Geob. 6 (1937) 34, fig. 3 A-C; Col. ll. Jap Pterid.

(1959) 53, 226, fig. 86; Ching Fl. Reip. Pop. Sin. 2 (1959) 270; Kurata, Satake

& Mashiko Pterid. Idzu Penins. (1960) 23, map II. — L. tenera Dryander var.

chienii (Ching) Tardieu-Blot & Christensen Fl. Gén. L-C. 7 (1939) 127; Iwatsuki

Acta Phytotax. Geob. 19 (1961) 5. — L. orbiculata (Lam.) Mett. ex Kuhn var.

chienii (Ching) Ohwi FI. Jap. Pteridoph. (1957) 35 (not seen). — Schizoloma chienii

(Ching) Tardieu-Blot Am. Fern J. 48 (1958) 34. Type: Ching 7184, Yao-mar Shan,

N.W. Kwangsi, China (isotype US). — L. orbiculata auct. non (Lam.) Mett. ex

Kuhn; Ito Fil. Jap. Ul. (1944) pl. 20; Ohwi FI. Jap. (1965) 40, p.p. Fig. 7

Rhizome short- to very short-creeping, 14-2 mm in diam.; scales castaneous,

narrowly lanceolate, the upper 4 or 4} uniseriate, up to 3-seriate at the base but

some entirely uniseriate, to ca. 14 mm long. Leaves clustered to a few mm apart;

petioles reddish brown to castaneous, lustrous, slender, upward gradually quad-

rangular with narrow, pale edges, not or only adaxially sulcate, ca. 10-30 cm

long, equaling the lamina to almost twice as long. Lamina herbaceous or firmly

herbaceous, dark green when dry, especially above, ca. 10-20 cm long, bipinnate,

Or tripinnate at the extreme base, deltoid, oblong (or rarely linear and simply

pinnate in juvenile but already fertile plants), with 1-6 pairs of major (fully

pinnate) pinnae to a side, without a conform terminal one; rachis like the

upper part of the petiole, upward gradually entirely pale. Pinnae narrowly

20 Gardens’ Bulletin, Singapore — X XVI (1972)

triangular, acute or shortly acuminate, very shortly petiolulate to subsessile, the

lower ones about their width apart but ascending and therefore often + contiguous,

the larger ones 5-10 cm long, 2-3 cm wide; secondary rachises pale to greenish,

abaxially flat or narrowly sulcate, at least above narrowly alate. Lower pinnae

with ca. 4-8 free pinnules to a side; pinnules ascending or the basal ones spreading,

of variable size and shape, depending on the place in the lamina, the larger ones

of basal pinnae in outline 1-elliptic, rhombic, or trapeziform, strongly dimidiate,

often ca. 8 X 4-12 X 6 mm, obtuse, incised, with 1-3 oblique incisions on the

upper and 1 or 2 on the outer side, these very oblique, of very unequal depth,

the basal pinnules often deeply pinnatipartite, with a few very unequal segments

on the anterior side, or less often fully pinnate, with a few flabellate pinnules on

both sides. Upper pinnules of lower pinnae and many or all of upper pinnae

cuneate-flabellate, gradually reduced to the pinna-apex, some confluent with it,

or sometimes, especially in larger pinnae, the pinna-apex basally pinnatilobate

but + distinct, rhombic to lanceolate, of variable size, acute or shortly acuminate,

+ distinctly serrato-lobate or basally pinnatifid, in size comparable to the larger

pinnules of the pinna or larger. Basal acroscopic pinnule cuneate-flabellate. A few

upper pinnae pinnatifid or only basally pinnate, transitional to the non-pinnate

pinnules in the base of the pinna-apex, these rhombic or trapezoidal, with several

incisions that go well beyond the receptacle, none, or only the uppermost that are

already connected with the pinnatifid leaf-apex not incised and with continuous

sori. Veins immersed, scarcely evident except in transmittent light, once or twice,

rarely 3 X, forked, +-4 mm apart, a costa scarcely or not developed. Incisions

of the margin acute, the lobes with parallel, straight or slightly convex-convergent

sides, their outer margin straight or convex in outer lobes, distinctly erose, usually

also slightly crispate; dimensions of lobes very variable. Sterile pinnules sharply

dentate; sterile leaves of simpler structure sometimes present with the fertile ones

but much less regularly so than in L. orbiculata. Sori extending to the terminal

segments, interrupted by the incisions of the margin, of variable length, often

1-3 mm long and on 2-5 vein-ends (rarely uninerval), strongly intramarginal,

with straight or, especially in outer lobes and smaller pinnules, basally concave

receptacle; indusium pale to brownish, + erose and/or crispate, 0.4-0.5 mm wide,

not reaching the margin by an equal or larger (rarely smaller) distance, not

reflexed at maturity. Spores light yellowish brown, trilete, verruculose, ca. 25 un.

Ecology. In shaded places, on rich soil, mostly at lower elevations but in the

tropics to above 1000 m. Apparently not rare in Japan.

Distribution. See below.

Geographically selected citations:

JAPAN. Honshu: Tagawa Fl. Jap. 6199 (US); Kurata 82, 1175a (TOFO). —

Shikoku: Inada 4194 (TOFO). — Kyushu: Otomasu 4928 (TOFO); Kurata &875

(TOFO). — Hachijo Jima: Sakurai s.n. (B, E). — Ryukyu: Yakushima, Jwatsuki

3427 (E, K, L, U, US); Arakane 136 (KYO); Hamaya 2784 (TOFO). Amami

Oshima, Hutch 21015 (TOFO). Tokara Gunto (Linschoten Is), Kawagoe s.n. (US).

Okinawa, Sonohara, Tawada & Amano 6293 (BISH, K. MICH, US); Conover 960,

1134, 1757 (US). Ishigaki, Oka 14720 (TOFO); Hatusima 22917 (TAI).

TAIWAN. Ogata 190 (BM); Hancock 71 (K); Suzuki 7442 (TAI); DeVol c-.s.

4475 (TAI).

CHINA. Kwangsi: Ching 7184 (US, isotype). — Hainan: Eryl Smith 1478

(K, SING). — Also reported from Kwangtung and Yiinnan by Ching (1959).

The Lindsaeoid Ferns of the Old World VI 21

INDO-CHINA. Annam: Chevalier 30685 (K, MICH, P).

THAILAND. Hennipman 3937 (L); Tagawa, Iwatsuki & Fukuoka T 1259 (L),

T 4808 (L, U, US).

Notes. The isotype seen by the author is less incised than most other specimens

but otherwise not atypical.

L. chienii is very closely related to, and probably to a certain degree inter-

fertile with, L. orbiculata var. commixta. There are a few intermediates, but

relatively so few that L. chienii is treated here as a distinct species.

5. Lindsaea orbiculata (Lamarck) Mett. ex Kuhn Miq. Ann. Mus. Bot. Lugd.-

Bat. 4 (1869) 279; Merrill Lingn. Sc. J. 5 (1927) 13; Ogata Ic. Fil. Jap. 1 (1928)

pl. 34; Tagawa Acta Phytotax. Geob. 6 (1937) 33; Tardieu-Blot & Christensen

Fl. Gén. I.-C. 7 (1939) 125; Ching FI. Reip. Pop. Sin. 2 (1959) 264; Chun, Chang

& Chen Fl. Hain. 1 (1964) 54; Iwatsuki Acta Phytotax. Geob. 19 (1961) 8; Ohwi

Fl. Jap. (1965) 40, p.p. — Adiantum orbiculatum Lamarck Encycl. 1 (1783) 41.

Type: Sonnerat s.n., Malacca (P). — L. flabellulata Dryander Trans. Linn. Soc.

3 (1797) 41, pl. 8 fig. 2; Beddome Ferns S. India (1864/65) 72, pl. 216 (poor);

Dunn & Tutcher Kew Bull. Add. Ser. 10 (1912) 337; Gibbs Common Hongkong

Ferns (1927) 10, pl. 15. Lectotype: Nelson s.n., Macao (BM). — L. longipes

Tardieu-Blot & Christensen Fl. Gén. L-C. 7 (1939) 125, fig. 15. Type: Poilane

8208, Ninh-Hoa, Annam (P, 2 sh.; dupl. in BM, K). — ? L. simulans Ching FI.

Reip. Pop. Sin. 2 (1959) 265, 371, nom. invalid. (see notes at the end of the genus).

For further synonymy and description see Fl. Mal. (sp. 3).

KEY TO THE VARIETIES

Plants usually (always ?) with simply pinnate sterile leaves beside the fertile ones;

fertile leaves simply pinnate. or, if bipinnate, at least some pinnules at the base

of the terminal pinna, above the uppermost pinnate pinnae, suborbicular; the

lamina not gradually passing from the bipinnate condition at the base to the

simply pinnate apex but with a rather abrupt transition ............... var. orbiculata

Sterile simply pinnate leaves usually wanting; no pinnules suborbicular; larger

laminas upward gradually of simpler structure, with a gradual transition from the

bipinnate base to the simply pinnate apex. .................ccccceeeeeees var. commixta

a. Var. orbiculata

Distribution. As given below, and extending to the Philippines and Java.

Geographically selected citations:

JAPAN. Shikoku: Okamura s.n. (GH). — Ryukyu: Amami Oshima, Tokashiki

s.n. (TOFO); Hutch 21150 (TOFO). Yoronjima, Ugehara s.n. (US). Okino Erabu,

Tagawa & Iwatsuki 2118 (E, K, L, U. US), 2157 (US); Hatusima & Sako 21402,

21507 (TOFO). Okinawa, Teruya 168 (SING); Walker 5923a (BISH, K, US).

TAIWAN. Tanaka & Shimada 13522 (BM, E, GH, L, MICH, SING, S-PA, US,

W, Z); Hancock 70 (BM, K, US); Faurie 619 (B, BM, W):; Ream 477 (GH.

MICH); DeVol & Huang 1213 (TAI, US); Kao 3485 (TAI).

22 Gardens’ Bulletin, Singapore — XXVI (1972)

CHINA. Hunan: Fan & Li 355 (A, BO, L, W). — Fukien: Tang S. G. 7065

(BM, S-PA); Schindler 421 (B, BM, E, K). — Kwangsi: Morse 47 (US); Tsang

24497 (A, MICH). — Kweichow: Cavalerie 3563 (E). — Kwangtung: Tsang

21753 (A, BO, E, K, L, MICH, W, Z); Canton Chr. College 12201 (E, US);

Delavay s.n. (P). — Hainan: Eryl Smith 1465 (BM, K, SING, US, p.p.); Mrs.

Hancock 70, 71 (US); Lau 1491, 4950 (MICH). — Prov.?: Osbeck s.n. (S-PA);

Meyen s.n. (B). — Hong Kong: many coll., eg. Eryl Smith 1468 (K, SING);

Matthew 308, 309 (E); Taam 1415 (US), 1882 (MICH, US); Faurie 15762 (E):

Wright s.n. (GH, K, US). — Macau: Nelson s.n. (BM, lectotype of L. flabellulata).

— Reported from Szechuan and Yunnan by Ching (1959); no material seen.

INDO-CHINA. Tonkin: Tsang 30149 (A, E, K), 30752 (A, K). — Annam:

Squires 344 (BO, BRI, E, SING, W); Pételot 3424, 3511 (P, US); Poilane 1094

(K, MICH, P), 2698 (GH, HBG, MICH, P), 5242 (GH, MICH, P), 8208 (BM,

K, P, type of L. longipes). — Cochin China: Gaudichaud s.n. (B, K).

THAILAND. Marcan 1228 (BM, SING); Kerr 2179 (BM K).

BURMA. Reported by Dickason (1946), no material seen.

INDIA. Assam: Clarke 45642 A, B (K). — Kerala: Ferguson ? (BM).

b. Var. commixta (Tagawa) Kramer Fl. Mal. II 1, 3 (1971) 207. — Lindsaea com-

mixta Tagawa Acta Phytotax. Geob. 6 (1937) 37, fig. 3 H-J; Ching Fl. Reip. Pop. Sin.

2 (1959) 268; Tagawa Col. Ill. Jap. Pterid. (1959) 54, 226, fig. 87. — L. tenera

Dryander var. commixta (Tagawa) Iwatsuki Acta Phytotax. Geob. 19 (1961) 6.

Type: Hidaka s.n., Tanage-Shima, Kyushu (KYO, not seen; photogr. in U). —

L. bonii Christ Not. Syst. 1 (1910) 187. Type: Bon 8, S. Tonkin (P). —

L. hainanensis Ching Bul. Fan Mem. Inst. Biol. N.S. 1 (1949) 298; Acta Phytotax.

Sin. 8 (1959) 141, fig. 16; Fl. Reip. Pop. Sin. 2 (1959) 267; Chun, Chang & Chen

Fl, Hain. 1 (1964) 54. Type: Lau 6149, Hainan (A, MICH, isotypes). — L. tenera

auct. non Dryander; Beddome Ferns S. India (1863/64, 1873) 7, pl. 24. Fig. 8

For further synonymy and description see Fl. Mal. loc. cit.

Distribution. See below; sporadically extending to Malesia.

JAPAN. Kyushu: Koshiki Jima, Ohyo 8913 (KYO, paratype). — Ryukyu:

Tanegashima, Tasiro s.n. (KYO, paratype). Yakushima, Ohtani E.J2 (U); Ohba

662674 (US); Kawanabe 5346 (TOFO); Hutch 19108 (TOFO); Kawabata 807, 808

(TOFO); Kudo s.n. (KYO, paratype). Nakanoshima, Hatusima 15720 (US). Amami

Oshima, Faurie 4599 (B, KYO, W, paratypes; p.p., with L. chienii); Hutch 21018,

21639, 21704 (TOFO); Koidzumi s.n. (KYO, paratype). Okinawa, Walker c:s.

6086 (US); Teruya 98, 166, 167 (SING); Ito s.n. (KYO, paratype); Sonohara,

Tawada & Amano 6294 (MICH, US); Conover 978, 1056, 1070, 1757, 1800 (US):

Ogata 19] (BM). Ishigaki, Oka 13348 (TOFO); Kawagoe s.n. (US). Iriomote,

Nishida 542 (US); Warburg s.n. (B). Yonaguni, Koidzumi s.n. (KYO, paratype).

TAIWAN. Sagae s.n. (TOFO); Hancock 71 (US); Ito s.n. (KYO, paratype);

Huang 1052 (TAI); Murphy s.n. (K); Botel Tobago, Chuang & Hsu 2448 (TAI);

Orchid I., Huang & Kao 7534 (TAI, UV).

CHINA. Hainan: Lau 6149 (A, MICH. isotypes of L. hainanensis), 1491 (GH);

Tsang & Fung L. U. 18141 (GH, K); Eryl Smith 1465 (BM, p.p.). — Hong Kong:

Lorrain 18 (K, p.p.).

The Lindsaeoid Ferns of the Old World VI 23

INDO-CHINA. Tonkin: Bon 8 (P, type of L. bonii). — Annam: Evrard 1280

(MICH, P); Hayata 582 (P); Chevalier 30640 (P); Cadiére 48 [68] (BM). —

Cambodia: Bouillod 59 (P). Cochin China: Gaudichaud s.n. (B).

THAILAND. Tagawa, Iwatsuki & Fukuoka T 1259 (US), T 637, 7153 (L, U, US);

Tagawa T 3939 (L, U, US); Sdrensen, Larsen & Hansen 2269 (E).

BURMA. Lace 4746 (E).

NEPAL. Ghose 10 (P).

INDIA. Assam: Godwin Austen s.n. (P); Clarke 45642 A, B (K); Wenger 15, 169

(K). — Uttar Pradesh: Fleming 864 (BM). — Kerala: coll.? (BM).

CEYLON. Thwaites CP 3311 (B, BM, BO, E, K, P, SING, W), CP 138] (B, BM,

BO, E, GH, K, L, P, SING, W); Wall s.n. (B, E, GH, K, P, US); Walker s.n.

(B, E, GH, P, S-PA, U, W): Sledge 1381 (K, U); Hutchison s.n. (E).

Notes. Many, not all, Ceylonese specimens have longer rhizomes, narrower

terminal segments, and more interrupted sori than typical specimens. This is

presumably due to introgression by L. schizophylla.

As stated under L. chienii, there are some intermediates between that species

and L. orbiculata var. commixta which may be of hybrid origin, e.g., Ohtani s.n.

from Yakushima (U). They have more strongly intramarginal and interrupted

sori and more incised subapical pinnules than typical var. commixta. On the other

hand, the number of intermediates between var. commixta and var. orbiculata is

considerably larger, and the differences between the two are more gradual and

not very easy to define. Therefore the two are treated as varieties of one species,

whereas L. chienii is retained as specifically distinct.

The type collection of L. hainanensis consists of large, atypical plants with

an almost conform terminal pinna, but in the absence of further material they

do not seem to represent more than an extreme form of var. commixta, typical

specimens of which have also been collected in Hainan.

6. Lindsaea repanda Kunze Bot. Zeit. 6 (1848) 541; Tagawa Acta Phytotax.

Geob. 6 (1937) 38, fig. 4 A — C. Type: Mertens s.n., Bonin Is. (LE; dupl. in L).

Rhizome short-creeping, ca. 2 mm in diam.; scales castaneous, very narrowly

triangular, about the apical 4 uniseriate, to ca. 5-seriate at the base, to ca. 15 mm

long. Leaves close to clustered; petioles quadrangular almost to the base, medium

or more often dark brown to blackish or adaxially paler, abaxially the greater

part pale-angled and scarcely sulcate, ca. 8—22 cm long, equaling to almost

twice as long as the lamina. Lamina olivaceous to brownish when dry, herbaceous

or usually subcoriaceous, ca. 8 —13 cm long, bipinnate or subbipinnate, rarely

almost simply pinnate when fertile, with 1 — 6 well-developed pinnae to a side,

rarely only the basal divisions subpinnate; no conform terminal pinna present.

Primary rachis like the petiole. abaxially largely or entirely dark and pale-angled.

Pinnae subsessile, spreading to ascending, the larger ones (14-) 3—5 cm long,

+— 14 cm wide; secondary rachises pale, abaxially bi-angular to sulcate. Pinnules

of well-developed pinnae 3—7 to a side, ‘+ narrowly and asymmetrically

obovate to flabellate-subrhombic, not very distinctly dimidiate, cuneate at the

base, sessile, + contiguous; larger fertile ones ca. 6— 10 mm long, 5—6 mm

wide; pinnules of small pinnae and of the leaf-apex trapezoid, distinctly dimidiate,

to 12 x 6 (-10) mm. Upper pinnules of lateral pinnae and of the leaf-apex

24 Gardens’ Bulletin, Singapore — X XVI (1972)

somewhat reduced, more narrowly cuneate, none or 1 or 2 connected with the

relatively large, rhombic to lanceolate, often lobed, obtuse to shortly acuminate

terminal segment (pinnule); upper (primary) pinnae rather abruptly reduced,

some transitional between pinnae and pinnules, + pinnatifid, very obtuse. Veins

immersed, not evident, free, 1 — 3 x forked, close, 4 — % mm apart. Upper/

outer margin of sterile pinnules coarsely erose-denticulate or more often irregu-

larly and sharply dentate; this margin in fertile pinnules with a few irregular,

shallow incisions ca. + mm deep, otherwise erose or not rarely sharply erose-

denticulate. Sori on the upper/outer margin, strongly interrupted even by quite

shallow incisions, on (1 —) 2—5 vein-ends, basally not rarely concave, their

interior edge usually irregular, seemingly slightly decurrent onto the apices of

the veins supporting them (see Tagawa’s figures); indusium pale, rigid, 0.4 —0.5

mm wide, its edge erose-denticulate, reaching, and, with its small lobes mostly

here and there slightly exceeding the margin, scarcely bulging at maturity. Spores

yellowish, trilete, verruculose, ca. 27p.

Ecology. No data extant.

Distribution. Confined to the Bonin Is.

JAPAN. Bonin Is: Hahashima, Hisauchi s.n. (KYO); Tuyama s.n. (KYO).

Ototoshima, Tuyama s.n. (KYO). Chichishima, Tuyama s.n. (KYO). Island?,

Hattori 272 (P); Mertens s.n. (L,LE, type); Toshima s.n. (KYO); Warburg s.n.

(B); Imp. Acad. Petersb. 43 (K, isotype ??).

Notes. A distinct species but evidently quite close to L. orbiculata var. commixta.

Among the species with sufficiently known range this is perhaps the most nar-

rowly distributed.

7. Lindsaea austro-sinica Ching Bull. Fan Mem. Inst. Biol. N.S. 1 (1949) 297;

Fl. Reip. Pop. Sin. 2 (1959) 266. Type: Tsang 22638, Shang-sze Hsien, Kwangsi,

China (GH, P, Pic-Ser, isotypes). — L. scandens auct. non Hooker; Tardieu-

Blot & Christensen Fl. Gen I.-C. 7 (1939) 121, pp.mai — UL. longipes

Tardieu-Blot & Christensen. loc. cit. 125, quoad specim. citat., p.p.,excl. typ. Fig.9

Rhizome rather short- to short-creeping, 2—4 mm in diam.; scales reddish

brown, very narrowly triangular, the apical 4 or 4 uniseriate, up to 4-seriate at

the base, to 21 mm long. Leaves close to a few mm apart; petioles lustrous,

castaneous to blackish, abaxially entirely terete and concolorous, adaxially

sulcate and upward usually narrowly pale-margined, 15—40 cm long, about

equaling to 14 x as long as the lamina. Lamina oblong, ca. 15—35 cm

long, medium to dark or olivaceous green when dry, herbaceous or chartaceous,

bipinnate (rarely tripinnate at the base), with 1 — 7, mostly 3 — 5 pinnae

to a side, the terminal pinna almost conform or passing rather abruptly

into the bipinnate part; primary rachis like the petiole or upward paler

and + sulcate, sometimes the uppermost part pale-margined. Pinnae very

narrowly triangular, shortly acuminate, or in small leaves oblong and acute

or obtuse, often distinctly petiolulate, 6 — 13 cm long, 24 —4 cm wide, relatively

broadest in paucijugate leaves, 14-5 x as long as wide, ascending, several cm

apart but sometimes contiguous through being ascending, with 2— 13 pinnules

to a side, the basal posterior pinnule of the lowermost pinnae sometimes pinnate.

The Lindsaeoid Ferns of the Old World VI 25

a few cm long, and a few pinnules next to it pinnatifid; secondary rachises abruptly

pale at their insertion, stramineous to pale brown, or the basal ones of large

leaves basally concolorous, abaxially terete at the base, upward gradually bi-

angular, sometimes shallowly sulcate, near the apex narrowly green-margined.

Pinnules less than half their width apart to contiguous, spreading or somewhat

ascending, dimidiate, otherwise variable in shape, the larger ones of paucijugate,

usually small laminas rhombic or subrectangular, 12—20 mm long, 9 —15

mm wide, less than 14 x as long as wide, often widest at the base, often with

one incision a few mm deep in the upper and another in the outer margin, the

lobes separated by them + convex; in plurijugate, mostly larger laminas, and

upper pinnules of some paucijugate ones, rounded-rectangular to ligulate (resem-

bling those of L. lancea and L. doryphora), 11— 14 mm long, 6—7 mm wide,

14 —2 x as long as wide, scarcely narrowed to the apex, the upper margin

rounded into the outer, the larger pinnules with 1 or 2 incisions up to 4+ mm

deep in the upper and one in the outer margin, the smaller ones entire; transitions

between these two shapes not rare. Lobes of smaller pinnules with straight or

laterally slightly protracted margin, scarcely erose. Upper pinnae little reduced,

the terminal pinna with larger pinnules of the shape described first and then

almost conform to the lateral pinnae, or very few pinnae more strongly reduced,

with a few transitions from pinnate pinnae to incised pinnules. Upper pinnules of

lateral pinnae little reduced, not less than 2 the size of the larger ones, the pinna-

apex triangular, acuminate, or rhombic and sometimes subobtuse in paucijugate

leaves, narrowly connected with one of the uppermost pinnules, asymmetric, broadly

cuneate or subtruncate at the base, there lobed, upward crenate, soriferous,

with interrupted sori. Sterile pinnules shallowly crenate (not sharply dentate).

Veins immersed or slightly raised, + evident, free, 2—4 x forked, }—1 mm

apart. Sori continuous in entire, interrupted in incised pinnules, on 2— 12 vein-

ends; indusium pale, 0.3 mm wide, entire, almost or quite reaching the margin,

rarely to 0.5 mm wide and not reaching the margin by about half its width, not

reflexed and scarcely bulging at maturity. Spores yellowish, verruculose, trilete,

Ca. 27 p.

Ecology. Terrestial in thickets and forests, on sand or clay, 700—41100 m.

Distribution. Southern China and Indo-China.

CHINA. Kwangsi: Tsang 22638 (GH, P, Pic-Ser, isotypes).

INDO-CHINA. Annam: Vincens s.n. (BO, K, MICH, P, SING; several coll.);

Cadiére I1 (P); Poilane 3378 (BISH, BO, MICH, P), 3472 (HBG, MICH), 44/3

(GH, HBG), 46/4 (P); Fleury (Chevalier 38646) (P); Chevalier 38819, 38834 (P).

— Cambodia: Bouillod 69 (P).

Notes. A distinctive species, and in spite of the great variability of the foliage

readily recognized by its dark, abaxially quite terete axes. Ching (1959) completely

misunderstood its affinity. The type of L. hainanensis bears some resemblance to

L. austro-sinica (see under L. orbiculata var. commixta); there may be some

hybridization, but L. austro-sinica does not seem to have been found in Hainan.

8. Lindsaea javanensis Blume En. Pl. Jav. (1828) 219; Tardieu-Blot & Christensen

Fl. Gen. I.-C. 7 (1939) 124. — Schizoloma javanense (Blume) Holttum Rev. FI.

Mal. 2 (1954) 349, fig. 202.

26 Gardens’ Bulletin, Singapore — XXVI (1972)

Fig. 9. L. austro-sinica. Left: sterile lamina (Vincens s.n., Annam, P). Right: fertile lamina

i (Bouillod 69, P).

The Lindsaeoid Ferns of the Old World VI ay

Type: Blume s.n., Java (L.) — L. flabellulata Dryander var. gigantea Hooker Sp.

Fil. 1 (1846) 211, pl. 63 c. — L. orbiculata (Lam.) Mett. ex Kuhn var. gigantea

(Hooker) Mett. ex Kuhn Mig. Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 279. — L.

tenera Dryander var. gigantea (Hooker) Holttum Gard. Bull. S.S. 5 (1930) 65. — L.

gigantea (Hooker) C. Christensen Bot. Jahrb. 66 (1933) 53. Type: Griffith s.n.,

Assam & Khasya Hills (K). — JL. orbiculata (Lam.) Mett. ex Kuhn var.

deltoidea Wu Bull. Dep. Coll. Sci. Sun Yatsen Univ. 3 (1932) 134, pl. 52. —

L. chienii Ching var. deltoidea (Wu) Tagawa Acta Phytotax. Geob. 6

(1937) 37; Col. Ill. Jap. Pterid. (1959) 54, 226. Type: Sin 180, Kwangsi, China

(not seen), — L. liangkwangensis Ching Fl. Reip. Pop. Sin. 2 (1959) 269, 372, ex

char. et synon. Type: Hsu 00813, Kwangtung, China (not seen).

For further synonymy and description see Fl. Mal. (sp. 5).

Distribution. From S. Japan and S.E. China to Assam and W. Malesia; reported

by Ching (1959) from the Chinese provinces of Kwangsi and Kwangtung.

Geographically selected citations:

JAPAN. Ryukyu: Yakushima, Kawabata 834, 973 (TOFO), 13890 (US). Amami

Oshima, Hutch 21266, 21606 (TOFO). Okinawa, Tawada 34 (TAI); T. Ito s.n.

(TAI); Naito s.n. (KAG).

TAIWAN: Ream 478 (MICH); Nakamura 527 (TAI); Simizu 3801 (TAI).

INDO-CHINA. Tonkin: Pételot 3422 (BM, US), 3646 (P). — Annam: Poilane

4375 (P); Cadiére 137 (P).

THAILAND: Hennipman 937 (L); Tagawa, Iwatsuki & Fukuoka T 1259 (VU),

1514 (L,U); Smitinand 1262 (K).

BURMA: Sidney s.n. (US).

INDIA. Assam: Griffith s.n. (K, S-PA, type of L. flabellulata var. gigantea); Mann

s.n. (B, BO, HBG, K, L, MICH, P, SING, S—PA); Jerdon s.n. (K); Godwin Austen

fan: *{P):

Notes. As noted in FI. Mal., the Continental-Asiatic and Japanese specimens of

this species are not in all respects like the Malesian ones. They are as a rule more

amply bipinnate, the large, asymmetric, terminal segment forming a relatively

smaller portion of the pinna; the two forms are sometimes present together on

one rhizome, e.g., Cadiére 137 from Annam (P). The continental form often —

but not always — has more strongly intramarginal indusia. There is probably

some hybridization with L. chienii.

9. Lindsaea chingii C. Christensen Ind. Fil. Suppl. II (1934) 121, based on:

L. chinensis Ching Sinensia 1 (1929) 5, non (L.) Mettenius ex Kuhn (1868) (=

Sphenomeris ch.); Hu & Ching Ic. Fil. Sin. I (1930) pl. 20; Ching FI. Reip. Pop.

Sin. 2 (1959) 263. Type: Ching 7968, Tsing Lung Shan, Kwangsi, China (not seen).

Fig. 6

Rhizome short- to very short-creeping, 145-2 mm in diam,; scales castaneous,

very narrowly triangular, about the apical 4 uniseriate, up to 6-seriate at the base,

up to 24 mm long. Leaves close to clustered; petioles ca. 12-20 cm long, 4-1 x

as long as the lamina, medium to dark brown or paler in the upper part, adaxially

broadly sulcate and green-margined above, abaxially in the upper part obtusely

bi-angular, + convex, scarcely marginate. Lamina oblong, ca. 15-22 cm long,

bipinnate, with ca. 6-10 well-developed pinnate pinnae to a side, without a

28 Gardens’ Bulletin, Singapore — XXVI (1972)

conform terminal one; colour olivaceous when dry, texture herbaceous; primary

rachis abaxially bi-angular, reddish brown or mottled, upward stramineous. Pinnae

subsessile, laxly ascending, very narrowly lanceolate, acuminate, the larger ones

3-10 cm long, 1-14 cm wide, with ca. 8-18 well-developed pinnules to a side, the

lower ones not, the upper ones + contiguous; secondary rachises slender,

pale, greenish, abaxially flat, upward gradually narrowly marginate. Pinnules

spreading or slightly ascending, dimidiate, sub-quadrangular or 4-elliptic

in Outline, the larger ones 4-7 mm long, 4-5 mm wide, as long as wide to almost

14 x as long as wide, cuneately subpetiolulate at the base. Larger pinnules with

straight or slightly concave lower margin, often little narrowed to the apex,

irregularly incised from the upper and outer edge, the incisions oblique, usually

one major one halfway the upper margin, reaching down to 4 or a in addition

1 or 2 shallower ones on the outer and a few on the upper margin on both sides

of the deeper one; lobes subdigitately divergent, ?-1} mm wide, 4-2 mm long,

the basal part parallel-sided, at least some subspathulately broadened at the sorus,

with convex sides, the outer edge with one or two apiculi to ca. 4 mm long, less

often only erose and rounded or subtruncate. Upper pinnae gradually reduced

with transitions to the pinnules of the leaf-apex; upper pinnules gradually and

strongly reduced, cuneate and pluri-, then uni-apiculate. some denticuliform ones

confluent with the pinnatifid, narrowly lanceolate, acute pinna-apex of ca. 1 cm.

Veins immersed, + evident, simple or once forked, 1 or 2 running to each lobe.

Sori single in the lobes, uni- or binerval; indusium pale, +-1 cm long, 0.3-0.4 mm

wide, with straight or convex or in binerval sori often somewhat concave base,

free at the sides, erose, overtopped by the apiculus, otherwise almost reaching the

outer margin of its segment. Spores like those of L. chienii, ca. 23-25p.

Distribution. Tonkin; described from Kwangsi, China, and not reported from other

provinces by Ching (1959).

Ecology. In thickets on dry sand or clay; altitude not noted.

INDO-CHINA. Tonkin: Tsang 27183 (E,K), 29330 (A,E,K,P); Balansa 121 (BM).

Notes. I have not seen the type of L. chingii, but Ching’s good description and

Hu & Ching’s figure are not open to misinterpretation. The description was

evidently based on a young specimen. The description given above was newly

made from seven presumably full-grown specimens.

L. chingii is a well-marked species and is apparently rather closely related

to the following.

10. Lindsaea bouillodii Christ Not. Syst. 1 (1909) 59. Type: Bouillod 48, Cambodia

(P). — L. tenera auct. non Dryander; Tardieu-Blot & Christensen FI. Gén.I.-C. 7

(1939) 126, and of other authors. Schizoloma tenerum auct. non (Dryander)

Holttum Rev. Fl. Mal. 2 (1954) 348, fig. 201. — L. cambodgensis auct. non

Christ; Kramer Blumea 15 (1968) 563.

For further synonymy and description see Fl. Mal. (sp. 2).

Distribution. Indo-China to Java and Borneo.

CHINA. Hainan: Eryl Smith 1478 (SING).

INDO-CHINA. Tonkin: Sollet 3512. — Annam: Pételot 3512 (US). — Cambodia:

Pierre 5767, 5787 (P); Bouillod 48 (P, type); Eryl Smith 2303 (BM.K,P), 2304 (K);

Kerr 7582(K), 15485 (BM, K).

The Lindsaeoid Ferns of the Old World VI 29

THAILAND. v. Beusekom 789, 791 (L, p.p. min.); Smitinand 5469 (K); Eryl Smith

1878 (K).

Noies. A specimen labelled “China, Forbes 506” (p.p.) in BM is perhaps from

elsewhere.

See also under L. tenera among the insufficiently known species.

11. Lindsaea schizophylla (Baker) Christ Journal de Bot. 21 (1908) 234.— Davallia

schizophylla Baker Syn. Fil. 2nd ed. (1874) 468. Type: Thwaites CP 3903, Ceylon

(BM; specimens with the same number but prob. from several collections in

B, BO, P, W). — Davallia trichomanoides auct. non (Dryander) Beddome; Beddome

Ferns Br. India 2 (1868) pl. 178. — L. orbiculata auct. non (Lam.) Mett. ex Kuhn;

Beddome Ferns Br. India (1892) 75, in part.

Rhizome rather short-creeping, 14 — 2 mm in diam.; scales castaneous to

fuscous, narrowly triangular, shortly uniseriate at the apex, up to ca. 8-seriate

at the base, to 2 mm long. Leaves not clustered, a few mm to 1 cm apart:

petioles ca. 12 — 25, often about 20 cm long, brown, dark at the base, reddish

or pale brown to stramineous near the apex, adaxially sulcate and sometimes pale-

margined upward, abaxially at least in the upper part acutely bi-angular

but scarcely sulcate. Lamina oblong or narrowly oblong, sometimes subtriangular,

12 —20 cm long, half as long as to equaling the petiole, dark or olivaceous green

when dry, herbaceous (mostly thinly), bipinnate & bipinnatifid, less often at the base

tripinnate & pinnatifid, or in small plants bipinnate & pinnatifid; primary rachis

stramineous or pale brown at the base, abaxially bi-angular, adaxially deeply sulcate.

Major pinnae 1 — 4 to a side, ascending or arcuately ascending, with a petiolule of

a few mm, 5 — 10 cm long, 14 -— 2 cm wide, acuminate, often slightly narrowed at

the base; secondary rachises abaxially flattened, greenish, upward gradually margi-

nate. Secondary pinnae (pinnules) slightly ascending, the lower ones not contiguous,

shortly petiolulate, up to 8 major ones to a side, pinnate & pinnatifid or only

pinnatifid, or the basal ones of basal pinnae & bipinnatifid; segments cuneate, bifid,

twice bifid, or simple, those on the anterior side larger and more numerous than

on the posterior side, the pinnules strongly anadromous. Ultimate divisions mostly

2— 5 mm long, at the base 0.3 — 0.5 mm wide, cuneately broadened from the base,

0.5 — 1 mm wide just below the sorus, at the sorus again and usually + spathulately

broadened, there 1 — 14 mm wide, outer margin rounded to. acute, usually erose,

sometimes truncate and then not at right angles to the vein. Upper pinnae rather

abruptly reduced, pinnate and twice bifid, those above again simpler and, like the

upper pinnules, etc., gradually confluent into the pinnatifid, acute lamina-, pinna-,

etc., apices. Sterile leaves sometimes only with short, pinnate & twice bifid pinnae,

then linear; sterile lobes subacute or acute, not spathulately broadened. Veins

immersed, single or rarely paired in the lobes, evident, especially their clavate ends

on the adaxial side. Sori uni-, less often binerval; indusium yellowish, delicate,

subentire or lobulate-erose, if short with convex base and adnate at the narrowed

sides (not pouch-shaped), if longer often with concave base and free at the rounded

sides, 4— 1 mm long, 1 —4 mm wide, not reaching the margin by its width or

more, scarcely bulging at maturity. Spores pale brown, trilete, smooth, ca. 22u.

Ecology. Terrestrial in montae forests, ca. 1600 — 2000 m.

Distribution. Endemic in Ceylon; most collections with data on exact provenance

from Adam’s Peak, only J. Smith s.n. marked “Siidspitze”’.

30 Gardens’ Bulletin, Singapore — XXVI (1972)

CEYLON. Ferguson s.n. (GH, US); Robinson 113a (K); Matthew s.n. (K); Alwis

s.n. (Z); Thwaites CP 3903 (BM, type; dupl.? in B, BO, E, K, P, W); Wall 1012

(P, S-PA); J. Smith s.n. (P. S-PA); Sledge 608 (K, U); Skinner s.n. (K); Hutchinson

s.n. (E).

Notes. The closest relative of this species is probably L. orbiculata var. commixta,

with which it seems to be interfertile to a certain degree. Putative hybrids are

reported under that taxon.

12. Lindsaea cambodgensis Christ Not. Syst. 1 (1909) 58; not of Kramer Blumea

15 (1968) 563. Type: Bouillod 61, Cam-Chay Mts., Cambodia (P). — L. parvipinnula

K. Iwatsuki Acta Phytotax. Geob. 19 (1961) 6, fig. 13. Type: Tagawa & Iwatsuki

3204, Yakushima, Japan (KYO, not seen; photogr. in U).

For a description the reader is referred to the very complete original one of

L. parvipinnula to which only a few points are added here.

Rhizome scales castaneous, very narrowly lanceolate, rather shortly uniseriate at

the apex, up to 5-seriate at the base, to almost 2 mm long. Petioles to 22 cm long.

Lamina to 15x 8 cm, not rarely subtripinnate & pinnatilobate at the base, i.e., the

basal pair(s) of pinnae with one or a few pinnate pinnules on both sides. Pinnae

subsessile. Pinnules scarcely dimidiate, the basal ones of larger pinnae always with

at least one incision going well beyond the receptacle, sometimes with more and

deeper incisions, especially if they are transitional between pinnate secondary pinnae

and non-pinnate pinnules; upper pinnules entire, flabellate-cuneate, gradually re-

duced, usually one connected with the very obtuse, cuneate-flabellate, often cleft or

lobed terminal segment. Sori on (1 —) 2—5 vein-ends, basally convex if very

short, often concave if longer, #—3 mm long. Indusium greenish, subentire or

slightly erose, 0.3—0.4 mm wide, not reaching the erose margin by the same

distance or a little less. Spores yellowish, trilete, nearly smooth, ca. 25u. Juvenile,

sterile plants with unevenly digitate-crenate pinnules.

Ecology. According to Iwatsuki (loc. cit.) terrestrial on the floor of light forest at

lower elevation.

Distribution. Yakushima and Okinawa, Japan; Cambodia. Also reported from

Amami-Oshima by Iwatsuki (loc. cit.); no material seen by the present author.

JAPAN. Ryukyu: Yakushima, Saiki sn. (KYO, paratype of L. parvipinnula);

Iwatsuki 2978 (KYO, id.); Ohtani E-12 (U); Furuse s.n. (TOFO); Satake s.n.

(TOFO): Kawabata 988 (TOFO); Hutch 18970 (TOFO); Yamaguchi 6 (TOFO);

Kawanabe 5045 (TOFO, isoparatype of L. parvipinnula), 4960 (KYO, TOFO, para-

types of L.p.); Ohba 66727 (U). Okinawa, Tawada 1556 (KYO, TAI, iso-paratypes

of L.p.), 114 (KYO, paratype of L.p.); Tamaki 723 (KYO, id.); Ito sn. (KYO),

INDO-CHINA. Cambodia: Eryl Smith 2304 (K); Bouillod 61 (P, type).

Notes. The disjunct distribution of this species is remarkable. The Cambodian

specimens are a little coarser in texture and dissection than the Japanese ones

which may be infraspecifically distinct.

The rather long-creeping rhizome, the small, cleft pinnules and the delicate

lamina characterize this species rather well.

The Lindsaeoid Ferns of the Old World VI 31

13. Lindsaea heterophylla Dryander Trans. Linn. Soc. 3 (1797) 41, pl. 8 fig. 1;

Beddome Ferns S. India (1863/64) 70, pl. 206; Dunn & Tutcher Kew Bull. Add.

Ser. 10 (1912) 338; Gibbs Common Hongkong Ferns (1927) 14, pl. 7; Tagawa Acta

Phytotax. Geob. 7 (1938) 197; Ito Fil. Jap. III. (1944) pl. 22; Iwatsuki Acta

Phytotax. Geob. 19 (1961) 3.—Schizoloma heterophyllum (Dryander) J. Smith

Hook. J. Bot. 3 (1841) 414; Beddome Ferns S. India (1863/64) 9, pl. 26; Blatter &

d’Almeida Ferns of Bombay (1922) 55, fig. 18; Merrill Lingn. Sc. J. 5 (1927) 13;

Ogata Ic. Fil. Jap. 3 (1930) pl. 148; Ching Fl. Reip. Pop. Sin. 2 (1959) 273, pl. 23 fig.

7 — 11; Chun, Chang & Chen FI. Hain. 1 (1964) 57. Type: Robertson s.n., Malacca

(BM). — Schizoloma intertextum Ching Fl. Reip. Pop. Sin. 2 (1959) 274, 374. Type:

Herb. Biol. Kwangsi Univ., Kwangsi, China (not seen).

For further synonymy and description see Fl. Mal. (sp. 7).

Distribution. Madagascar and Mascarenes, S. India and Ceylon to Ryukyu and

Central Malesia.

Geographically selected collections:

JAPAN. Ryukyu: Okinawa, Conover 958, 1173 (US); Sonohara 65 (US); Ogata 195

(BM); Tawada 130 (U); Tagawa & Iwatsuki 4845 (US). Iriomote, Nishida 540 (US);

Walker & Tawada 6635 (MICH, US). Yonaguni, Hatusima 24424 (TOFO).

TAIWAN. Steere s.n. (MICH); A. M. Evans 089 (TENN).

CHINA. Kwangtung: Canton Chr. Coll. Herb. 1573 (US); Tsang 26045 (A);

Gressitt 1213 (BM, E, GH); Petersen s.n. (L).— Kwangsi: Morse 6 (US); Ching

7758 (US; paratype of Sch. intertextum); Tsang 22106 (GH, Pic-Ser; id.), 23909

(MICH).— Hainan: Eryl Smith 1500 (K, SING, US); Gressitt 966 (E, GH);

Hancock 4 “‘p. p.”’ (BM). — Hong Kong: Cadiére 92 (MICH, SING); Eryl Smith

1479 (K, SING); Taam 2210 (US); Wright s.n. (GH, K, US); Matthew 311, 312 (E);

Hance & Simson 667 (W).— Reported from Yunnan by Ching (loc. cit., as Sch.

intertextum).

INDO-CHINA. Tonkin: Bourret 7 (MICH); Balansa 125 (K), 1972 (K, MICH);

Tsang 29370 ((A, p.p.).— Laos: Poilane 2117 (GH, MICH, S-PA). — Annam:

Poilane 5241 (GH, MICH); Cadiére 49 (GH).— Cambodia: Eryl Smith 2306 (XK).

THAILAND. H. M. Smith 202, 212 (US); Tagawa, Iwatsuki & Fukuoka T 592

(U, US), T 1311, 1831 (U); Tagawa 3951 (U);Hennipman 3588, 3937a (L); Kerr 779,

2372, 6841 (K).

INDIA. Bombay: Meebold 9613 (S-PA).— Mysore: Blanford s.n. (E); Law s.n.

(GH); Meebold 9614 (S-PA). — Madras: Beddome s.n. (BM, E, K); Faucheux s.n.

(BM). — Kerala: Beddome s.n. (BM).

CEYLON. Thwaites CP 386] (B, BM, E, K); J. Smith s.n. (S-PA); Wail s.n. (B,

GH).

Note. The Chinese specimens have mostly broader, more rigid and erose indusia,

but are connected with the typical form by intermediates.

32 Gardens Bulletin, Singapore — XXVI (1972)

14. Lindsaea ensifolia Swartz Schrader, J. Bot. 1800 (1801) 77; Dunn & Tutcher

Kew Bull. Add. Ser. 10 (1912) 338; Gibbs Common Hongkong Ferns (1927) 12, pl.

6; Tagawa Acta Phytotax Geob. 7 (1936) 197, 198; Ito Fil. Jap. IIL. (1944) pl. 23;

Iwatsuki Acta Phytotax. Geob. 19 (1961) 2; Kramer Blumea 15 (1968) 564. —

Schizoloma ensifolium (Swartz) J. Smith Hook. J. Bot. 3 (1841) 414; Beddome Ferns

S. India (1863/64, 1873) 9, pl. 25; Ferns Br. India (1892) 80, pl. 41; Blatter &

d’Almeida Ferns of Bombay (1922) 53, fig. 17; Merrill Lingn. Sc. J. 5 (1927) 13;

Ogata Ic. Fil. Jap. 6 (1935) pl. 293; Tardieu-Blot & Christensen Fl. Gén. I.-C. 7

(1939) 129, fig. 15 1-2; Ching Fl. Reip. Pop. Sin. 2 (1959) 273, pl. 23 fig. 1-6; Chun,

Chang & Chen FI. Hain. 1 (1964) 56, fig. 24. Type: coll. unknown, Mauritius (S-PA).

— L. griffithiana Hooker Sp. Fil. 1 (1846) 219, pl. 68 B. — Schizoloma griffithianum

(Hooker) Fée Gen. Fil. (1852) 108. Type: Griffith s.n., Mergui, Burma (K).

For further synonymy and description see Fl. Mal. (sp. 8).

Of the three subspecies recognized by Kramer (loc. cit.) only one occurs in the

area of the present paper, viz. ssp. ensifolia.

Distribution. Old World tropics to Malesia, western Melanesia, and tropical

Australia.

Geographically selected citations:

JAPAN. Ryukyu: Iriomote, Nishida 534 (US).*.

TAIWAN. Henry 1497 (E), 1502 (B); Faurie 130 (S-PA).

CHINA. Kwangsi: Tsang 24699 (A, MICH).—Kwangtung: Lau 20178 (A);

Merrill 10899 (GH); Dalziel s.n. (E).— Yiinnan: Wang 81126 (A). — Hainan:

Lau 6369 (MICH); Wang 34040 (E, GH), 34261 (MICH.— Hong Kong: many

coll., e.g., Eryl Smith 1501 (SING, US); Taam 1409 (MICH, US), 221] (US);

Topping 605 (GH, US), 608 (US); Fortune 3 (BM, E), 27 (E, L); Matthew 304 (E).

INDO-CHINA. Tonkin: Balansa 107 (K).— Laos: Colani 4111 (BM); Poilane

2116 (BM, HBG, MICH).— Annam: Cadiére 50 [67] (K).— Cambodia: Pierre

5794 (BM, HBG, K). — Cochin China: P. Condor, Harmand 875 (MICH, SING);

Pierre 5743, 5801, 5802 (MICH), 5794 (E, HBG).

THAILAND. Eryl Smith 1236, 1879, 1880, 1904 (K, SING); Molesworth-Allen

2154 (S-PA); H. M. Smith 599 (GH, MICH, US); Tagawa 3937 (U); Tagawa,

Iwatsuki & Fukuoka T 594, 1262, 1309 (L, U); v. Beusekom 316 (L); Larsen,

Smitinand & Warneke 659, 896 (L).

BURMA. H. M. Smith 575 (US); Dickason 8007, 8026 (A); Sidney 33 (BM, US);

Brandis 274 (K); Mergui, Griffith s.n. (K, type of L. griffithiana).

SIKKIM. Gamble 347 A (E); Clarke 36796 B (EB).

NEPAL. Wallich 93 or s.n. (B, L, US).

BANGLA DESH. Gamble 7909 (E, K, US); Cowan 924 (E); Hooker & Thomson

s.n. (B, GH, K, W); Thomson s.n. (B, E, S-PA, VU).

4 A specimen labelled “Yokohama”, Schottmueller 122 “p.p.” (B), is probably mislabell-

ed; the species was not reported from Honshu by Tagawa (1959).

The Lindsaeoid Ferns of the Old World VI 33

INDIA. Assam: Mann s.n. (B. E, HBG, K, L, SING, S-PA, US); Clarke 42947 B

(K, US); Meebold 7390 (K); Hooker s.n. (B). — Rajasthan: Gamble 2387 A (K).

— West Bengal: Gamble 409 A and s.n (K), 409 B (E).— Madras: Gamble 15974

(K, US). — Kerala: Wight 214 (B).

CEYLON.Thwaites CP 1382 (B, BM, E, GH, S-PA, U, W); Hance 32 (W); Walker

s.n. (B, BM, GH, K, W); J. Smith s.n. (S-PA); Naylor Beckett 61 (E, GH);

Koenig s.n. (L); Freeman 51, 52, 89 (BM); F. Schmid 1090 1125 (BM); Macrae

s.n. (EB).

PUTATIVE HYBRIDS IN SECTION SCHIZOLOMA

Four specimens in herb. TOFO combine characters of two species each of section

Schizoloma and are presumably of hybrid origin. Yamaguchi 14 from Yakushima has a

short rhizome and the leaf pattern of L. orbiculata var. commixta but pinnules rather like

L. cambodgensis; its spores are abortive. Yamaguchi 9 from the same island is somewhat

like the preceeding but looks more like a hybrid between L. chienii and L. cambodgensis;

it also has abortive spores. Satuke s.n., also from Yakushima, is about midway between L.

kawabatae and L. chienii. Oka 13890 from Iriomote has very irregularly dissected subbipin-

nate leaves with features of both L. orbiculata and L. heterophylla. The spores of the two

last-named collections are well-developed.

For supposed hybrids between L. orbiculata var. commixta and L. schizophylla see

under the former.

SECTION Synaphiebium (J. Smith) Diels

15. Lindsaea malayensis Holttum Gard. Bull. S. S. 5 (1930) 69, fig. 8; Rev. FI.

Mal. 2 (1954) 335, fig. 194. Type: Md. Haniff 4032, G. Kerbau, Perak, Malaya

(SING).

For description see Fl. Mal. (sp. 15).

Distribution. Malay Penisula: Malaya and southern Peninsular Thailand.

THAILAND. Tagawa, Iwatsuki & Fukuoka T 4781 (VU).

16. Lindsaea napaea v.A.v.R. Bull. Jard. Bot. Btzg. II. 20 (1915) 19, pl. 3;

Holttum Gard. Bull. S. S. 5 (1930) 66; Rev. Fl. Mal. 2 (1954) 330, fig. 189. Type:

Teijsmann 16616, Mt. Dai, Lingga Is. (BO).

For description see Fl. Mal. (sp. 16).

Distribution. Sumatra, Malaya, and adjacent islands; southern Peninsular Thailand.

THAILAND. Kerr 15227 (K).

17. Lindsaea malabarica (Beddome) Baker ex Christensen Contr. U. S. Nat.

Herb. 26 (1931) 295; Baker Syn. Fil, 2nd ed. (1874) 545 (in indice; invalid

combination, species not accepted). —- Schizoloma malabaricum Beddome Ferns

Br. India (1868) pl. 268, with descr. — Sch. lobatum (Poiret) Beddome var.

malabaricum (Beddome) Beddome Ferns Br. India (1883) 79, (1892) 79, pl. 39 (err.

‘malabarica’). Type: Beddome s.n., South Canara, Mysore, India (K).

Rhizome short-creeping or rather so, 1-14 mm in diam.; scales not seen.

Leaves close to almost 1 cm apart; petioles stramineous to pale reddish brown,

quadrangular except at the extreme base, scarcely sulcate except adaxially, ca.

10-20 cm long, 4 to about as long as the lamina. Lamina simply pinnate or

bipinnate with one or two pairs of lateral pinnae and a conform terminal one,

medium green when dry, herbaceous, ca. 15-35 cm long; pinnule-bearing rachises

like the upper part of the petiole. Pinnules ca. 20-35 to a side, 4-1 x their width

apart or the upper ones closer, ascending (often strongly) or the lower ones almost

34 Gardens’ Bulletin, Singapore — XXVI (1972)

spreading, parallelogrammoid or subtrapezoid, the major ones 10-12 mm long,

34-5 mm wide, 2-3 X as long as wide; margins straight or the upper and/or lower

outward somewhat convex, a distinct outer margin developed or not; basal pinnules

sometimes a little reduced, upper pinnules gradually and strongly reduced, a few

denticuliform ones connected with the almost linear terminal segment. Veins

immersed, ‘+ evident, once or twice forked, rather close, 1 mm apart, free or

usually the inner ones here and there connivent, rarely truly anastomosing but

never regularly so. Upper margin with 3 or 4 oblique incisions to # mm deep,

reaching or slightly surpassing the level of the receptacle, the lobes laterally

narrowed, usually with convex outer margin. Sterile lobes subacute. Sori inter-

rupted, on 2—3 (— 5) vein-ends, with straight or laterally slightly convex, or in the

outer sori slightly concave receptacle; indusium pale or greenish, rigid, entire or

crenulate, 0.3 mm wide, reaching the margin or nearly so, scarcely reflexed at

maturity. Spores not seen.

Distribution. Southern India; recorded from Burma by Christensen (1931) and

Dickason (1946), probably in error.

INDIA. Madras: Ghatak 440 (K). — Mysore: Beddome s.n. (K, type).

Notes. With the very limited material at hand I feel uncertain about the status of

this species. Yet it does not seem to fit any of the others of the present section. It

might be taken for a juvenile form of L. venusta, but the juvenile specimens of

that species I saw which were in size comparable to the material of L. malabarica

have more regularly crenate pinnules, shorter sori, and much less rarely anastomos-

ing veins. More material is required in order to establish the variability of ZL.

malabarica.

A collection that may represent the mature form is Thomson s.n., “Mt.

Nilghiri & Kurg’ (B, E, GH, K, S-PA, U, US, W). It has uni- or bijugate-

bipinnate leaves, larger pinnules (up to 16 X 7 mm), + parallel-sided lobes with

straight outer margin, incisions to 3 mm deep going to 4 or in the apical half to

the middle of the pinnule, and in the basal half of larger pinnules more or less

regularly, elsewhere irregularly anastomosing veins. In some characters it is quite

divergent from the material on which the above discription is based, but part of

the material in S-PA is incompletely fertile and much more like it. It bears some

resemblance to L. lobata var. epirotes but has more deeply and irregularly incised,

less elongate, more truncate pinnules and much less regularly anastomosing veins

that are closer in the pinnule lobes.

18. Lindsaea obtusa J. Smith in Hooker Sp. Fil. I (1846) 224, Type: Cuming 394,

Malacca (K; dupl. in B, GH, W). — ZL. davallioides auct. non Blume; Ito Fil. Jap.

Ill. (1944) pl. 21.

For further synonymy and description see Fl. Mal. (sp. 18).

Distribution. Taiwan and Malesia to western Melanesia; to be expected in southern

Thailand.

TAIWAN. Faurie 245 (S-—PA); Suzuki s.n. (TAI 6179); Sasaki s.n (TAI 6183).

ANDAMAN IS. South Andaman, Mann s.n. (P, S-PA); Kurz s.n. (K).

Notes. The specimens from South Andaman are not typical, having smaller and

more shallowly incised pinnules than usual. They may represent a local form, but

L. obtusa is as a whole very variable. —— See also the note after L. lobata var.

epirotes, p. 36.

cee

Ae eh one ease tment hin ee scteneceeripesiee one

The Lindsaeoid Ferns of the Old World VI 35

19. Lindsaea parallelogramma v.A.v.R. Bull. Jard. Bot. Btzg III. 5 (1922) 212;

Holttum Gard. Bull. S. S. 5 (1930) 70, fig. 9; Rev. Fl. Mal. 2 (1954) 335, fig. 193.

Type: Biinnemeijer 7359, P. Singkep, Lingga Is (BO; dupl. in L).

For further synonymy and description see Fl. Mal. (sp. 22).

Distribution. S. Peninsular Thailand and Malesia.

THAILAND. Kerr 14497 (K).

20. Lindsaea cultrata (Willdenow) Swartz Syn. Fil. (1806) 119; Kramer Blumea

15 (1968) 565; not of other authors. Type: coll.?, ‘““Malabaria”’ (B, herb. Willdenow).

—— L. decomposita Willdenow Sp. Pl. (1810) 425; Holttum Gard. Bull. S. S. 5

(1930) 66, fig. 5; Rev. Fl. Mal. 2 (1954) 333, fig. 192, and of other authors, at least

in part. Type: coll.? “India” (B. herb. Willdenow). L. recurvata Hooker Sp.

Fil. 1 (1846) 222, pl. 70 A. Synaphlebium recurvatum (Hooker) J. Smith Hook.

J. Bot 3 (1841) 415; Hist. Fil. (1875) 268, pl. 18 c. Type: prob. Wallich sin. dat. (K).

For further synonymy and description see Fl. Mal. (sp. 23).

Distribution. Southern India (?), Ceylon; southern Thailand; Malesia; Orchid I.

and Botel Tobago; ? Hainan.

TAIWAN. Botel Tobago, Chuang & Hsu 2453 (TAI); C. C. Hsu s.n. (U). Orchid

I., Huang & Kao 7532 (TAI, U); Fukuyama s.n. (TAI 6178, 6184); Sasaki s.n. (TAI

6181).

CHINA. Hainan: Hancock (Kew no. 119) (K; mislabelled 7).

THAILAND. Eryl Smith 2301 (BM, K), 2301 B (K); Hennipman 3937b (L);

v. Beusekom 791 (L).

INDIA. S. India, without exact loc., Gough s.n. (K, poor, uncertain). ——- The

exact provenance of the types of L. cultrata and L. decomposita, as cited above,

is uncertain.

CEYLON. Thwaites CP 982 p.p. (with L. venusta) (B, BM, E, GH, K, L, P,

S—PA, W); Sledge 543, 619, 1229 (K); Holttum S.F. 39201 (SING); Henderson s.n.

(K); Freeman 47, 49 (BM), 50 (BM, prob.); Gardner 1122 (BM, E, K), 1260, 126]

(B, K); Ballard 1077 (K); and others.

Notes. Not all Ceylonese specimens agree with the Indo-Malayan ones. Some are

quite typical, others have more strongly reduced upper pinnules, and in many the

outermost sorus of the upper margin is not or only in some pinnules, continuous

with that of the outer, a fairly constant character in Malesia. As some Ceylonese

collections are not divergent, it does not seem advisable to treat the local

from as infraspecifically distinct. There may be some introgression of L. venusta.

The specimens from the islets near Taiwan are very similar; they can be told apart

from L. lobata var. epirotes by i.a. less elongate pinnules, from var. hainaniana by

less deeply incised pinnules and straighter outer edges of pinnule lobes.

21. Lindsaea integra Holttum Gard. Bull. S. S. 5 (1930) 67, fig. 6. Type: Holttum

20934, Pahang, Malaya (SING; dupl. in BM, BO, K, US). — L. nitida auct. non

_ Copeland; Holttum Rev. Fl. Mal 2 (1954) 333, fig. 191.

For description see Fl. Mal. (sp. 25).

Distribution. Malay Peninsula, Sumatra, Borneo; southern Peninsular Thailand.

THAILAND. Eryl Smith 1877, 1878 (K), 2298, 2299 (BM, K).

36 Gardens’ Bulletin, Singapore — XXVI (1972)

22. Lindsaea lobata Poiret in Lamarck Encyl. Suppl. 3 (1813) 448; Kramer

Blumea 15 (1968) 565. Type: Commerson s.n. Java (P.) — L. davallioides Blume

En. Pl. Jav. (1828) 218; Hooker Sp. Fil. 1 (1846) 224, pl. 68 A; Tardieu-Blot &

Christensen Fl. Gén. I-C. 7 (1939) 127; Ching Fl. Reip. Pop. Sin. 2 (1959) 271;

Chun, Chang & Chen FI. Hain. 1 (1964) 56; not of Ito Fil. Jap. Il. (1944) pl. 21.

Type: Blume s.n., Java (L).

For further synonymy and description see Fl. Mal. (sp. 21).

Distribution. Malesia to western Melanesia and Micronesia. Two continental

groups of plants seem referable to this species, but they differ decidedly from the

Malesian material and are therefore here described as new varieties. They may prove

to be distinct species, although they are certainly very close to L. lobata.

a. Var. epirotes Kramer var. nov.

Petiolus brunneus vel raro nigrescens. Lamina unijugo-bipinnata vel unipinnata.

Pinnulae herbaceae, pellucidae, parallelogrammoidales vel paulo angustatae et sub-

trapezoidales, 11-16 mm longae, 3-5 mm latae, margine exteriore saepe distincto.

Margo anterior/exterior incisus, incisionibus basalibus minime duplum spatii a

receptaculo ad marginem aequantibus, exterioribus plerumque sensim profundi-

oribus. Lobi basales margine exteriore receptaculoque rectis. Indusium 4+ mm latum,

marginem attingens vel fere. __

Type: Tsang 29002, Taai Wong Mo Shan, Tonkin (A; dupl. in E, P).

Petiole medium to dark reddish brown, rarely black, occasionally stramineous.

Lamina unijugate-bipinnate or unipinnate, drying medium green. Pinnules her-

baceous, translucent, parallelogram-shaped or usually somewhat narrowed to the

apex and then subtrapezoidal, 11-16 mm long, 3-5 mm wide, 23-34 (-4) x as long

as wide; a distinct outer margin usually developed. Inner incisions of the pinnules

at least twice as deep as the distance from the receptacle to the margin, outer ones

as deep or usually progressively deeper; inner lobes hardly, outer ones somewhat or

very little divergent. Inner lobes with straight outer margin and straight receptacle,

outer ones usually with weakly convex margin and receptacle. Upper pinnules

strongly and gradually reduced. Veins mostly regularly anastomosing, occasionally

with an incomplete second series of areoles. Indusium + mm wide, nearly or quite

reaching the margin.

Ecology. Terrestrial in thickets and woods, 700-1500 m.

Distribution. Indo-China; to be expected in S. E. China.

INDO-CHINA. Tonkin: Tsang 27107 (A, K), 29002 (A, E, P, type), 27310 ?

[sic] (E).— Annam: Poilane 1525 (GH, MICH), 3384 (GH. MICH, SING), 7006

(US), 7169 (BO, K, MICH).

Notes. In many respects close to the Malesian form. The most important difference

is in the margin of the lobes and the receptacle which are straight at least in the

proximal lobes; the petioles are usually darker and there is often a distinct outer

margin. Certain forms of L. obtusa are also not unlike this variety; they have

wider indusia (0.4-0.7 mm) which are more strongly intramarginal, and usually

dry blackish.

The Lindsaeoid Ferns of the Old World V1 37

b. Var. hainaniana Kramer var. nov. — L. decomposita auct. non Willdenow;

Merrill Lingn. Sc. J. 5 (1927) 12. — L. cultrata auct. non (Willd.) Swartz; Merrill

loc. cit. 13 (prob.). Fig. 1.

Petiolus stramineus. Lamina unijugo-bipinnata vel unipinnata. Pinnulae her-

baceae, paulo pellucidae, subligulato-parallelogrammoidales, parce vel haud

angustatae, margine exteriore saepe distincto, majores 12-15 mm longae, 5-6 mm

latae. Margo anterior/exterior incisus, incisionibus minime triplum spatii a recep-

taculo ad marginem aequantibus, Lobi margine exteriore et receptaculo parce sed

distincte convexis, etiam basales divergentes. Indusium =-4 mm latum, mar-

ginem attingens vel fere.

Type: Liang 63789, Hainan (US; dupl. in K).

Petiole stramineous. Lamina unijugate-bipinnate or unipinnate, drying bright

to dark green. Pinnules herbaceous, little translucent, subligulate-parallelogram-

moid, little or not narrowed to near the apex, a distinct outer margin usually

developed; major pinnules 12-15 mm long, 5-6 mm wide, over 2 to 24 x as long

as wide. Incisions going to 4 of the width, rarely less but at least 3 x the distance

from the margin to the receptacle. Lobes somewhat narrowed, even the inner ones

distinctly divergent, most or all, including the inner ones, with weakly but

distinctly convex outer margin and receptacle. Veins regularly anastomosing, not

rarely with an incomplete second series of areoles. Indusium = or 1 mm wide,

almost or quite reaching the margin.

Ecology. Terrestrial in forests, sometimes on rocks, in moist places, ca. 1000 m.

Distribution: Hainan.

CHINA. Hainan: Lau 3817 (BISH, GH, S-PA), 5227 (A, MICH); Eryl Smith

1469 (K, SING); McClure Canton Chr. Coll. 9482 (BISH, BM, K); Wang 35711

(GH): Liang 63789 (K, US, type).

Notes. In general appearance not unlike L. cultrata, especially the Ceylonese form.

It differs in the more deeply incised pinnules with convex outer margin and

receptacle, all typical features of L. lobata. In most specimens the upper pinnules

are less gradually and strongly reduced than in var. Jobata and var. epirotes, but

more so than in typical L. cultrata. — See also the notes after the last-named

species.

23. Lindsaea venusta Kaulfuss ex Kuhn Linnaea 36 (1869) 79. Type: Thwaites

CP 982, Ceylon [dupl. (all?) in B, BM, BO, E, GH, K, SING, S-PA, W].—

Schizoloma recurvatum auct. non (Wall. ex Hooker) Moore; Beddome Ferns S.

India (1863/64; 1873) 9, pl. 27. Fig. 2.

Rhizome slender, 1-2 mm in diam., not very short-creeping, soon almost

scaleless; scales light brown, lanceolate, relatively shortly uniseriate at the apex,

up to ca. 7-seriate at the base, up to 14 mm long. Leaves not close, up to 24 cm

apart; petiole stramineous to pale or reddish brown, below obtusely, above acutely

quadrangular, adaxially upward sulcate, 8-30 cm long, in small, usually simply

pinnate leaves 2) in large, bipinnate ones up to almost 14 xX as long as the

lamina. Lamina ca. 10-25 cm long, bright medium or dark to olivaceous green

when dry, chartaceous, simply pinnate or more often bipinnate, with 1 or 2 pairs

of lateral pinnae and a conform terminal one; primary rachis stramineous or

pale reddish brown, abaxially bi-angular. Pinnae (if any) or simply pinnate lamina

10-20 cm long, 14-24 cm wide (the terminal sometimes larger), ascending or

38 Gardens’ Bulletin, Singapore — XXVI (1972)

spreading, subsessile, the lateral pairs (if any) several cm apart, not contiguous,

acuminate; secondary rachises stramineous, abaxially bi-angular, shallowly sulcate.

Pinnules ca. 25-35 to a side, spreading or slightly ascending, almost their width

apart to subcontiguous, parallelogrammoid, trapezoid, or less often subligulate,

mostly with + parallel upper and lower margins, i.e. hardly narrowed to the apex,

both margins straight, or the uper slightly convex, the lower slightly concave, the

inner straight, the outer distinct, straight or a little convex, meeting the upper

at right angles or at a smaller angle; major pinnules 10-15 mm long, 5-6 mm

wide, 2-24 (—3) x as long as wide. Upper and outer margin very shallowly crenate,

regularly so if sterile, less so if fertile, with rounded to subacute, uninerval lobes,

the incisions up to 1 mm deep but usually only + or 4 mm, reaching or slightly

surpassing the level of the receptacle. Lobes with slightly convex or straight outer

margin. Upper pinnules gradually and strongly reduced, one or a few + denticuli-

form ones connected with the small, narrowly lanceolate terminal segment. Veins

immersed, '+ evident, mostly twice forked, rather regularly anastomosing, forming

a series of areoles 4-1 mm wide. Sori interrupted by the incisions, on (1 -)

2-4 (—5) vein-ends; indusium with at least laterally convex base, pale, entire,

4 mm wide, not reaching the margin by a smaller distance to equaling or

slightly surpassing it, scarcely reflexed at maturity, the laminal lobe oppsite it

often pale on the adaxial side, Spores pale browish, trilete, smooth, ca. 21 ug.

Ecology. Terrestrial in forests, 200 — 700 m (very few data).

Distribution. Ceylon, apparently not rare; southern Peninsular India.

INDIA. Madras: Beddome 40 (E, K), s.n. (BM, K). — Kerala: Beddome s.n. (K).

CEYLON. Thwaites CP 982 p.p. (B, BM, BO, E, GH, K, SING, S-PA, W,

some or all isotypes); Moon s.n. (BM); Lenormand s.n. (B, fragm.); F. Schmid

1110, 1116, 1133 (BM); Wall s.n. (BO, GH, S-PA); Rawson 554a 3320 (BM);

Ferguson 36 (GH), 44 p.p. (US); v. Fridau s.n. (HBG); Sledge 997 (K), 1377

(K, U); Robinson 116 p.p. (K); Naylor Beckett s.n. (K); Haycock s.n. (BM);

Randall s.n. (BM); Bradford s.n. (BM); Emerson s.n. (E); Koenig s.n. (L).

Notes. In spite of the excellent original description, the wide distribution of the

type collection, and the distinctness of the species, L. venusta almost fell into

oblivion, the name appearing on hardly any herbarium label. The short, sub-

marginal sori and regularly crenate sterile margin are unique in the section but

are reminiscent of L. repanda of the Bonin Islands (sect. Schizoloma), and aiso

of L. kirkii of the Seychelles, a similarity already noted by Kuhn (loc. cit.). The

affinities of L. venusta in section Synaphlebium are not clear.

SECTION Lindsaea

24. Lindsaea doryphora Kramer Blumea 15 (1968) 566. Type: Alston 13358,

Permantang, Kalimantan, Borneo (U; dupl. in BM).— L. scandens Hooker var.

terrestris Holttum Rev. Fl. Mal. 2 (1954) 327, nom. invalid. (not typified). —

L. lancea auct. non (L.) Beddome of various authors, as to Asiatic material, at

least in part.

For description see Fl. Mal. (sp. 32).

Distribution. Malay Peninsula to Borneo and the Philippines.

THAILAND. Kerr 14458 (K); Eryl Smith 463 (K, SING), 1874 (K), 1875 (K,

SING), 2269, 2295, 2296 (K).

BURMA. Tenasserim: Wallich s.n. (K); Wight 219 (B).

The Lindsaeoid Ferns of the Old World VI 39

SECTION Aulacorhachis Kramer, sect. nova.

Squamae rhizomatis nodulis corticis oblongis insidentes; lamina bipinnata,

rhachides saltem sulco adaxiali tomentum perbrevem sed sine lente manifestum

gerentes; pinnulae dimidiatae, haud incisae, venulis liberis, soris continuis.

Species typica (adhuc unica): Lindsaea caudata Hooker.

25. Lindsaea caudata Hooker Sp. Fil. 1 (1846) 215; Beddome Ferns S. India

(1863/64) 73, pl. 217.— L. trapeziformis Dryander var. caudata (Hooker) Cesati

Atti Ac. Sc. Fis. Mat. Napoli 7 (1877) 14.—JZ. lancea (L.) Beddome var.

caudata (Hooker) v.A.v.R. Handb. 1 (1908) 273. Type: Mrs. Walker s.n., Adam’s

Peak, Ceylon (K).—L. lancea auct. non (L.) Beddome; Beddome Ferns Br.

India (1892) 75.

Rhizome terrestrial (prob. short-creeping, only small pieces seen), up to 4

mm in diam.; scales medium brown, narrowly triangular, long-acuminate to the

rather shortly uni-bi-seriate apex, to 3 mm long, inserted on oblong mounds of

cortical tissue from which they are not quite sharply differentiated at the ca.

6-seriate base. Leaves presumably close; petioles dark brown to blackish, +

lustrous, rather stout, up to 3 mm in diam. at the apex, abaxially terete, adaxially

flattened but scarcely sulcate, ca. 25 — over 60 cm long, about as long as to

14 x as long as the lamina. Lamina usually dark olivaceous to blackish when dry,

herbaceous or chartaceous, ca. 25—50 cm long, bipinnate, with 3—9 pinnae

to a side and a conform terminal one; primary rachis like the petiole, occasionally

laterally + puberulent just below the pinna bases or rarely throughout, adaxially

sulcate, in the groove persistently or fugaciously puberulent. Pinnae laxly as-

cending, often alternate throughout, with a petiolule of a few mm to 2 cm, Ca.

10 —25 cm long, 23 —44 cm wide, basally not or only anteriorly slightly nar-

rowed, rather abruptly and strongly narrowed near the apex, with ca. 15 —25

well-developed pinnules to a side; upper pinnae little or not reduced; secondary

rachises dark reddish brown or occasionally darker or pale, mostly abruptly

discolorous at their insertion, abaxially terete, adaxially sulcate, permanently

puberulent in the groove, in the basal part often also laterally, sometimes the

posterior pinnule-bases also slightly puberulent; hairs 0.2 —0.4 mm long, with

up to 5 cells, the terminal cell rounded; rachis of the terminal pinna not or not

abruptly discolorous. Pinnules spreading or slightly ascending, close to contiguous

or slightly overlapping, not rarely touching or overlying the rachis with their

inner margin, i-elliptic to subligulate, in shape very much like those of

L. parasitica, L. doryphora, and L. lancea, cuneate-subpetiolulate at the base,

the inner margin straight, the lower straight or basally concave or apically convex,

the upper margin outward increasingly convex, broadly rounded into the outer

which is not truly distinct, transition between outer and lower margin shortly

rounded to subacute; larger pinnules 1322 mm long, 7—10 mm wide, 2

(- 24) x as long as wide. Upper pinnules rather abruptly reduced, a few denti-

culiform, one or a few connected with the small, narrowly lanceolate, in her-

barium specimens often lost pinna-apex (but one specimen, Fendler s.n., GH,

with scarcely reduced upper pinnules and a large, flabellate, free terminal

pinnule, the terminal pinna with a normal apex). Upper/outer edge of pinnules

shallowly and regularly crenate in sterile, quite entire in fertile pinnules; lower

base usually pale and distinctly sclerotic. Veins evident, abaxially elevated at the

base or throughout, free, 1—3 x forked, rather close, ca. ? mm apart. Sori

continuous around the upper and outer margin (except in incompletely fertile

40 Gardens’ Bulletin, Singapore — XXVI (1972)

pinnules), with abundant filiform pluricellular paraphyses; indusium brownish,

rigid, entire, narrow, 0.2 —0.3 mm wide, almost or quite equaling the margin,

strongly reflexed and quite concealed at maturity. Spores medium brown, trilete,

minutely granulate, ca. 35 —40 un.

Ecology and distribution. Confined to Ceylon; hardly any data on habitat and

distribution on the island. Probably only in the mountains of the central massif;

very few recent collections, perhaps now very rare through destruction of forests.

CEYLON. Thwaites CP 1380 (B, BM, BO, E, GH, L, P, SING, W), s.n. (K, W);

Ferguson 37 (GH); Beddome s.n. (K); Thomson s.n. (B, GH, P); Hooker &

Thomson 302 (BM); Gardner 22 (P.W.), s.n. (BM, K); Wall s.n. (B, BO, K, S-PA);

Finlayson s.n. (BM); Nietner s.n. (HBG); Mrs. Walker s.n. (K, type); J. Smith s.n.

(S-PA); Robinson s.n. (K); v. Fridau s.n. (HBG); Sledge 821 (U); Emerson s.n.

(E); Gower s.n. (E); Wight 1916 (E).

Notes. In spite of its similarity to L. parasitica, L. doryphora, and L. lancea,

L. caudata is readily distinguishable by the pubescence in the adaxial grooves of

the leaf axes. It is surprising that this character has not been described before.

It is almost unique in the Lindsaeoid ferns, being otherwise only found in the

Antillean Odontosoria uncinella.

SECTION Osmolindsaea Kramer

26. Lindsaea odorata Roxburgh Calc. J. Nat. Hist. 4 (1846) 511; Kramer

Blumea 15 (1968) 567. Type: a plant from the Garrow Hills, India; no specimen

extant, Plate 2578 of Icones Roxburghianae (K) to be regarded as type. —

L. cultrata (Willd.) Swartz var. minor Hooker Sp. Fil. 1 (1846) 204 in part, excl.

lectotype. — L. cultrata (Willd.) Swartz var. pallens Hooker Sp. Fil. 1 (1846) 204;

Kunze Linnaea 24 (1851) 276. Lectotype: de Silva 151 or s.n., Silhet, Assam (K).

— L. cultrata (Willd.) Swartz var. attenuata Hooker Sp. Fil. 1 (1846) 204.

Lectotype: Griffith 862, Assam (K). — L. neocultrata Ching & Wang Acta

Phytotax. Sin. 8 (1959) 165, pl. 19 fig. 15; Ching FI. Reip. Pop. Sin. 2 (1959) 260,

pl. 22 fig. 8-9; Chun, Chang & Chen FI. Hain. 1 (1964) 54, fig. 23. Type:

E. Hainan Exped. 893, Tia-lo Shan, Hainan (not seen). — L. cultrata auct. non

(Willd.) Swartz of nearly all authors, e.g., Beddome Ferns S. India (1863/64, 1873)

7, pl. 23; Nakai J. Coll. Sc. Imp. Un. Tokyo 31 (1911) 403; Ogata Ic. Fil. Jap. 2

(1929) pl. 79; Tagawa Acta Phytotax. Geob. 6 (1937) 26, fig. 1 A-C; Tardieu-Blot

& Christensen Fl. Gén. I.-C. 7 (1939) 120; Ito Fil. Jap. Ill. (1944) pl. 16; Holttum

Rev. Fl. Mal. 2 (1954) 328, fig. 188; Ching Fl. Reip. Pop Sin. 2 (1959) 260, pl. 22

fig. 1-7; Tagawa Col. Ill. Jap. Pterid. (1959) 53, 226, fig. 84; Chun, Chang & Chen

Fl. Hain. 1 (1964) 55.

For further synonymy and description see Fl. Mal. (sp. 33). Two varieties in

the present area: *

a. Var. odorata.

Pinnules incised, sori interrupted; upper pinnules gradually and strongly reduced.

Distribution. Tropical and subtropical Asia from Ceylon, Tibet, and Japan to

western Melanesia, Rhodesia and Madagascar. Very common in many parts of

Asia.

* A third variety, var. darjeelingensis Sen & Sen, was recently described from West

Bengal [Am. Fern J. 61 (1971) 14].

Se ee ee

The Lindsaeoid Ferns of the Old World VI 4]

Geographically selected citations:

JAPAN. Honshu: Tagawa 7385 (E, GH, K, L, U); Tagawa & Iwatsuki 718

(E, GH, K, L, Pic-Ser, U, US); Ito 82 (SING). — Kyushu: Sugimoto s.n. (A). —

Hachijo Jima: Ohba s.n. (TOFO). — Ryukyu: Yakushima, Tagawa 7&9 (K,

SING); Faurie 4596, 4597 (B, BM, W); Togasi TNS 1479 (E, K, S-PA, US).

TAIWAN, Ream 461 (MICH); Hancock 6 (B, BM, K, US); Oldham s.n. (GH,

K, W); Faurie 618, 619 (S-PA); Kao 4116 (TAI).

CHINA. Szechuan: Wilson 2671 (BM, E, HBG, K, US, W); Fang 3233 (E, GH,

US), 3918, 7998 (E, K); H. Smith 2065 (S-PA). Yiinnan: Delavay 21 (K):

Handel-Mazzetti sn. (K); Cavalerie 1769, 7060 (E, K); Forrest 9358 (E, K),

11756 (BM, E, K, W); Rock 7189, 7359 (US); Tsai 52429 (US), 55242 (GH, US),

58989 (GH), 60747 (GH, S-PA); Maire 66 (S-PA), s.n. (SING, US, W, Z). —

Kweichow: Tsiang 4584 (E, GH, SING, S-PA, US), 4726 (GH, SING). —

Kwangsi: Ching 5787 (US); Tsang 22715 (A, W). — Kwangtung: Tsang 20226

(A, K, US, W), 25219, 25385 25475 (A); Metcalf 17512, 17738 (MICH); Merrill

1607 (GH).

TIBET. Forrest 19953 (E); Ludlow, Sheriff & Taylor 7032 (BM).

INDO-CHINA. Tonkin: Pételot 581 (US), 5401 (GH). — Annam: Chevalier

30690 (K), 30776 (SING).

THAILAND. Soérensen, Larsen & Hansen 2335 (E, K); Smitinand 395 (KK); Floto

7405 (K); Tagawa, Iwatsuki & Fukuoka T 595 (L, U).

BURMA. Kingdon Ward 21160 (BM, GH); Parish s.n. (E); Buchanan s.n. (E);

Toppin 4294 (E).

BHUTAN. Ludlow & Sheriff 997 (BM, E); Griffith s.n. (B, K, W); Cooper 2828

(BM).

SIKKIM. Bor & Ram 19730 (BM, SING); Meebold 2117 (B); Gamble 6998 (E),

9961 (K); Engler 5483 (B).

NEPAL. Wallich 148 or s.n. (B, BM, E, GH, K, US); Bonner 134 (BM); Stainton,

Sykes & Williams 6941 (BM, E, Pic-Ser).

BANGLA DESH. Clarke 19626 (BM).

INDIA. Assam: many coll., e.g., Sinclair 2950 (E); Kingdon Ward 18663 (BM),

18794 (A, BM); Hooker & Thomson s.n. (B, BM, E, GH, K, S-PA, U, US, W);

Mann s.n. (BO, SING, S-PA); Biswas 4101 (GH). — West Bengal: Gupta 17

(SING); Bir s.n. (U, US); Pichi-Sermolli 4604, 4635 (Pic-Ser). — Mysore:

Thomson s.n. (B, BM, E, S-PA, U, W). — Madras: Ysander s.n. (S-PA); Wight

220 (E); Bembower 3 (MICH); Gamble 17365 (E); Noyes s.n. (GH); Faucheux

sn. (BM). — Kerala: Meebold 13334 (B, S-PA).

CEYLON. Holttum SFN 39212 (SING); Wall s.n. (BO, E, GH); Freeman 43,

44, 45 (BM); Thwaites CP 3070 (B, BM, BO, E, GH, L, W): Hance 47 (W);

Nietner s.n. (B, HBG); Parish s.n. (E); Gardner 1121 (E).

Notes. Some of the Chinese and Japanese specimens have more widely creeping

rhizomes than usual but are not otherwise divergent.

42 Gardens’ Bulletin, Singapore — XXVI (1972)

As noted in Fl. Mal., freshly dried leaves of L. odorata have a distinct

coumarin-like odour. I am much indebted to Dr. K. Iwatsuki, Kyoto, for the

following information otherwise not accessible to me. According to Shimada,

Sawada, Kozuka, and Kojima (Jap. J. Pharmac. 22, 1968, 37-38; Japanese)

coumarin (I) was found in the methanol extract of L. odorata (‘‘cultrata’’) from

the Kii Peninsula, Honshu. It is supposed to be present in the living plant

[which lacks the odour] as a glycoside of coumaric acid.

b. Var. japonica (Baker) Kramer comb. nov. Basionym: L. cultrata (Willd.)

Swartz var. japonica Baker Syn. Fil. 1st ed. (1867) 105. — L. japonica (Baker)

Diels in E. P. Pfl. Fam. I4 (1899) 221; Tagawa Acta Phytotax. Geob. 6 (1937) 27,

fig. 1 d-e; Ito Fil. Jap. Ill. (1944) pl. 17; Tagawa Col. Ul. Jap. Pterid. (1959) 53,

226, fig. 85; Ching Fl. Reip. Pop. Sin. 2 (1959) 259. Type: Oldham s.n., Nagasaki,

Japan (K; dupl. in GH).

Rhizome not very short-creeping, 4-1 mm in diam.; scales not seen. Petioles

at least in the basal half, often throughout, castaneous, abaxially terete, 2-7 cm

long, mostly shorter than the lamina. Lamina 2-9 cm long, 1-24 cm wide, with

3-15 pinnules to a side, these their width apart to subcontiguous, spreading,

herbaceous, mostly olivaceous when dry, asymmetrically triangular to dimidiate-

ovate, obtuse or less often subacute, 4 x 3 to 11 x5 mm, the basal ones

slightly or not reduced, sterile and crenate or with a short sorus near the apex,

the middle ones entire, with an unbroken sorus, or rarely with one shallow

incision interrupting the sorus. Upper pinnules little reduced, about half as long

as the lower ones, the leaf-apex consisting of a cuneate-flabellate, distally always

truncate, free or almost free pinnule; or in larger specimens more strongly reduced,

some denticuliform ones confluent with the narrow, lobed leaf-apex. Indusium as

in var. odorata, or in small specimens a little narrower; spores as in var. odorata.

Distribution. See below.

KOREA, Cheju D6 (Quelpaert I: Faurie 90 (B, BM, E, MICH); Taquet 51

(E, W), 2339 4754 (E, K), 3542 (E, S-PA), s.n. (GH, MICH, W).

JAPAN. Honshu: Jto 82 (SING). — Hachijo Jima: Ohsuga s.n. (TOFO). —

Kyushu: Maximowicz 120 (B, BM, GH, K, L, S-PA, W); Oldham s.n. (GH,

K, type), 477 (B); Kido 2663 (TOFO): Tagawa & Iwatsuki 1077 (E, GH, K, L, VU);

Nameégata 11560 (TENN), 11796 (US). — Ryukyu: Yakushima, Ohba 66182 (U);

Ito 83 (SING). Amami Oshima, Hosoyamada s.n. (US). Okinawa, Sonohara,

Tawada & Amano 7096 (BISH, MICH, US); Conover 931 (US), 1798 (BISH, US);

Ito sn. (US); Tashiro sn. (US); Ogata 194 (BM). Ishigaki, Walker & Tawada

7283 (US). — Iriomote, Nishida s.n. (US); Walker & Tawada 6757 (US); Ito s.n.

(US); Hatusima 23046 (TAI).

TAIWAN. Ito s.n. (BM, GH); Swinhoe s.n. (B); Oldham s.n. (BM, W, p.p.);

Wilford 466 (K); Faurie 254 (S—PA); Ford s.n. (K).

CHINA. Szechuan: H. Smith 13614 (BM, E, S—PA).

Notes. At first sight this might be taken for a small phenotypic form of L. odorata,

and I am not quite sure that this is not the case, i.a. in view of the disjunct

occurrence in western China. After comparison with a large series of typical

L. odorata it seemed that the combination of small size, dark petiole, unbroken

sori, reduced and quite or almost sterile basal pinnules could serve for distinguishing

a taxon, but, especially in view of the great variability of L. odorata proper,

at the varietal level. Small specimens of L. odorata from the Philippines look

very much like var. japonica but have paler axes, broken sori, or both.

The Lindsaeoid Ferns of the Old World VI 43

27. Lindsaea himalaica Kramer spec. nov. — L. cultrata (Willd.) Swartz var.

minor Hooker Sp. Fil. 1 (1846) 204, p.p., excl. lectot. — L. cultrata (Willd.)

Swartz var. assamica Hooker loc. cit., ex char. (no material cited). Fig. 3

L. odoratae valde affinis, differt petiolo et saltem dimidio basali rhachidis

obscuris, pinnulis margine exteriore distincto, incisionibus tantum singulis vel

binis. Type: Simons s.n., Bhutan (BM).

Rhizome short-creeping, 14 mm in diam.; scales reddish brown, very narrowly

triangular, the basal half or less biseriate, the apical half uniseriate, up to 4-seriate

at the extreme base, up to 14 mm long. Leaves close to clustered; petiole

atropurpureous to blackish, + lustrous, abaxially rounded, shorter than the

lamina. Lamina linear, simply pinnate, ca. 15-35 cm long, slightly longer than

to ca. 3 X as long as the petiole, 2-34 cm wide, with (10—) 20-40 pinnules to a

side; rachis abaxially rounded or narrowed-rounded, at least in the basal half

dark reddish brown to blackish. Pinnules spreading or little ascending, half their

width apart to contiguous, subcoriaceous to coriaceous and olivaceous when dry,

subtrapezoidal or subparallelogrammoid, little or not narrowed to the truncate

or very shortly rounded apex, the base cuneiform-subpetioluliform, abruptly pale-

offset from the rachis; larger pinnules 10-18 mm long, 44-9 mm wide, 2-24 X as

long as wide. Lower margin straight, upper margin straight or slightly convex

towards the apex, meeting the outer margin at right angles or very shortly rounded

into it. Upper margin with 1 or 2 very shallow incisions to 4 mm deep; outer

Margin entire, its sorus continuous with the outermost of the upper margin.

Veins immersed, scarcely visible even in transmittent light, free, close, 3-?} mm

apart, 1-3 X forked. Upper pinnules gradually and + strongly reduced, ane or

a few denticuliform ones connected with the small, narrow, lobed, lanceolate-linear

leaf-apex; sometimes a few basal pinnules + remote and reduced. Sori 2 or 3

per pinnule, usually somewhat convex on the inner margin near the incision(s)

of the pinnule; indusium brownish, rigid, entire or nearly so, scarcely narrowed

at the ends, 0.4-0.5 mm wide, almost or quite reaching the margin. Spores medium

brown, monolete, bean-shaped, smooth, ca. 40 x 25 un.

Ecology. No data.

Distribution. See below.

BHUTAN. Griffith 135 or s.n. (B, GH, K, MICH, syntype of ZL. cultrata var.

minor), Simons s.n. (BM, type).

INDIA. Assam: Simons s.n. (BM); Griffith sn. (K, type of L. cultrata var.

assamica ?); March s.n. (K). N.E.F.A., R. S. Rao 17471 (K).

Notes. Two specimens from N.E. India, Meebold 4787 from Manipur (B) and

Watt 11039 from Assam (GH), approach L. himalaica in texture and in outline

of pinnules, but have only basally dark rachises and more interrupted sori. They

are better regarded as an extreme form of L. odorata. With the material at hand

L. himalaica seems sufficiently distinct to merit specific rank.

SECTION Psammolindsaea Kramer

28. Lindsaea walkerae Hooker Sp. Fil. 1 (1846) 209, pl. 69A; Beddome Ferns

S. India (1863/64) 72, pl. 215; Ferns Br. India (1892) 76; Kramer Blumea 15

(1968) 560, 17 (1970) 177. — Schizoloma walkerae (Hooker) Kuhn Chaetopt.

(1882) 346; Tardieu-Blot & Christensen Fl. Gén. I.-C. 7 (1939) 128; Holttum

Rev. Fl. Mal. 2 (1954) 344. Type: Mrs. Walker s.n., Ceylon (K; dupl. in B).

For further synonymy and description see Fl. Mal. (sp. 35).

44 Gardens’ Bulletin, Singapore — XXVI (1972)

Distribution. Ceylon and Indo-China to Malesia, Queensland, and Micronesia.

INDO-CHINA. Tonkin: Balansa s.n. (P).—Cochin China: Talmy s.n. (P);

Bois 2226 (BM, P).

CEYLON. Thwaites CP 1379 (B, BM, BO, E, GH, K, S-PA, U, W); Mrs. Walker

s.n. (B, K, type); McKenzie s.n. (K); Ferguson 43 (US); Walker s.n. (GH); Fendler

2028 (GH); Mrs. McDonnell s.n. (K); Wall s.n. (BO, GH); Gardner 1379 (K): Sledge

1376 (K); Wight 105 bis (E); Emerson s.n. (BE).

SECTION Isoloma (J. Smith) Kramer

29. Lindsaea divergens Hooker & Greville Ic. Fil. (1831) pl. 226; Beddome Feins

Br. India (1892) 76, fig. 38. — Isoloma divergens (Hooker & Greville) J. Smith

Hook. J. Bot. 3 (1841) 414; Holttum Rev. Fl. Mal. 2 (1954) 337, fig. 195. Type:

a Wallich collection from herb. Roxburgh (K, herb. Hooker, ?).

For further synonymy and description see Fl. Mal. (sp. 41).

Distribution. Malay Peninsula, Malesia to Borneo and Palawan.

THAILAND. Kerr 15935 (K).

SECTION Stenolindsaea Kramer

30. Lindsaea lucida Blume En. PI. Jav. (1828) 216; Tardieu-Blot & Christensen

Fl. Gén. I-C.7 (1939) 122; Holttum Rev. Fl. Mal. 2 (1954) 328, fig. 187; Ching FI.

Reip. Pop. Sin. 2 (1959) 263; Chun, Chang & Chen Fl. Hain. 1 (1964) 55; Kramer

Blumea 15 (1968) 567. Type: Blume s.n., Java (L). — L. gracilis Blume En. PI.

Jav. (1828) 217. Type: Blume s.n., Java (L). —- L. concinna J. Smith Hook. J. Bot.

3 (1841) 415, nom subnud.; Hooker Sp. Fil. 1 (1846) 205, pl. 61 B; Tagawa Acta

Phytotax. Geob. 6 (1937) 29, fig. G-I; Ito Fil. Jap. Ill. (1944) pl. 19; Ching FI. Reip.

Pop. Sin. 2 (1959) 262. Type: Cuming 198, Luzon (K; dupl. in B, E, GH, HBG, L,

SING, W). — L. lobbiana Hooker Sp. Fil. 1 (1846) 205, pl. 62 C. Type: Lobb s.n.,

Java (K). — L. kusukusensis Hayata Ic. Pl. Form. 4 (1914) 211, fig. 143; Ching

Fl. Reip. Pop. Sin. 2 (1959) 262. — L. concinna J. Smith var. kKusukusensis (Hayata)

Tagawa Acta Phytotax. Geob. 6 (1937) 30. Type: Hayata & Sasaki s.n., Kusukusu,

Taiwan (not seen), — L. minima Ching Sinensia 1 (1930) 52, non Copeland (1929).

— L. changii C. Christensen Ind. Fil. Suppl. 3 (1934) 121; Ching Fl. Reip. Pop.

Sin. 2 (1959) 261. Type: Chang s.n., Kuling, Kiangsi, China (dupl. in K). — L.

cultrata (Willd.) Swartz var. attenuata Hooker Sp. Fil. 1 (1846) 204, p.p., excl.

lectot.

For further synonymy and description see Fl. Mal. (sp. 42).

Distribution. In continental Asia etc. only ssp. lucida, which occurs East to Malesia

and the Palau Is.

JAPAN. Ryukyu: Ishigaki, Walker & Tawada 7182 (MICH, US); Nishida 312

(US); Tagawa & Iwatsuki 4486 (US), 4761 (E, K, L, U). Iriomote, Hatusima 18532

(US); Bandai sn. (TOFO); Koidzumi s.n. (US).

CHINA. Kiangsi: Chang s.n. (K, isotype of L. minima = L. changii). — Kwang-

tung: Tsang 26755 (A, K). — Hainan: Eryl Smith 1466 (K, SING, US); Lau 3596

(BISH, GH, MICH, S-PA), 5107 (MICH); How 72036, 72725 (GH); Wang 34571

(E. GH, MICH).

INDO-CHINA. Tonkin: Pételot s.n. (BM); Tsang 29900 (A, E, K, TAI), 30036,

30236 (A, E, K). — Annam: Sallet 5 (BO); Cadiére 51 (MICH); Matthew 25 (K),

s.n. (BM). — Cochin China: Gaudichaud s.n. (B).

The Lindsaeoid Ferns of the Old World V1 45

THAILAND. H. M. Smith 602 (GH, MICH, US); Hansen & Smitinand 12264 (L);

Eryl Smith 922 (K), 1701 (SING), 1559, 1876 (K), 2279 (BM, K); Kiah 24297

(K. SING); Marcan 1260, 1264 (BM); Kerr 9235, 17029 (K); Sdérensen, Larsen &

Hansen 182 (E, K); Murton 7 (K); Iwatsuki & Fukuoka T 7389 (VU).

BURMA. Rock 750 (BM, US); Sidney 32 (U, US). Tenasserim, Falconer s.n.

(BO, L); Wight 220 (US); Parish s.n. (BM).

BHUTAN. Griffith 1670 (E).

BANGLA DESH. King 61 (US).

INDIA. Assam: Mann s.n. (E, HBG, L, S-PA, US); Griffith s.n. (B, K); Day s.n.

(GH); Barnard 41 A (BM); Wenger 167 (K). — Manipur: Watt 6925 (E). — West

Bengal: Thomson s.n. (B). — Madras: Gamble 17365 p.p. (US). — Andaman Is:

Kunstler s.n. (B). — Not seen from Taiwan. To be looked for in Ceylon.

SUBGENUS Odontoloma (Hooker) Kramer

SECTION QOdontoloma

31. Lindsaea glandulifera v.A.v.R. Bull. Jard. Bot. Btzg Il. 1 (1911) 9. Type:

Koorders 15415 B, Besuki, Java (BO; dupl. in K, L). — L. repens (Bory) Thwaites

f. minor Thwaites En. Pl. Zeyl. (1864) 388. — L. repens (Bory) Thwaites var.

minor (Thwaites) Beddome Ferns S. India (1863/64) 72, pl. 214. Type: Thwaites

CP 1389 p.p mai., Ceylon [K; dupl. (all?) in B, BM, BO, E, GH, W].

For further synonymy and description see Fl. Mal. (sp. 44).

Distribution. East Java and Lesser Sunda Is.

CEYLON. Thwaites CP 1389 p.p. mai. (B, BM, BO, E, GH, K, W, type coll.

of L. repens f. or var. minor); Hooker & Thomson s.n. (BM); Robinson 110a (K);

Skinner s.n. (K); J. Smith s.n. (S-PA).

Note. The strongly disjunct distribution is of an unusual pattern. The two popula-

tions do not seem to be even infraspecifically distinct.

32. Lindsaea repens (Bory) Thwaites En. Pl. Zeyl. (1864) 388; Beddome Ferns

S. India (1863/64) 72, pl. 209; Ogata Ic. Fil. Jap. 5 (1933) pl. 232, as to plant

depicted; Tardieu-Blot & Christensen Fl. Gén. I.-C. 7 (1939) 120; Kramer

Blumea 15 (1968) 568, 17 (1970) 180. — Dicksonia repens Bory Voy. 2 (1804)

323. Type: Bory s.n., Réunion (P). — L. macraeana auct. non (Hooker & Arnott)

Copeland of later authors, e.g., Holttum Rev. Fl. Mal. 2 (1954) 324, fig. 185;

not of Ching Fl. Reip. Pop. Sin. 2 (1959) 266.

For further synonymy and description see Fl. Mal. (sp. 46). In our area this

species is only represented by var. pectinata (Blume) Mettenius ex Kuhn Mig. Ann.

Mus. Lugd. -Bat. 4 (1868) 277; Kramer Blumea 15 (1968) 568. — L. pectinata

Blume En. Pl. Jav. (1828) 217; not of Holttum Rev. Fl. Mal. 2 (1954) 324.

Type: Blume s.n., Java (L).

For further synonymy and description see Fl. Mal. (sp. 46, 3).

Distribution. Assam and Indo-China to the Greater Sunda Is. and the Philippines.

INDO-CHINA, Annam: Evrard 1872 (BO, GH, K), Poilane 3513 (MICH),

3533 (SING), 21852 (BO, GH, K, MICH, SING), 22343 (MICH): Fleury

(Chevalier 38797) (K); Vincens s.n. (GH, K, MICH, SING); Anet 1919 (BO) —

Cochin China: Pierre 5707 (BO, E, GH, HBG, K, MICH, SING), 5707

A (E, K, MICH). — Cambodia: Poilane 15175 (K, MICH, SING).

46 Gardens’ Bulletin, Singapore — XXV]I (1972)

THAILAND. Smitinand 860 (K); Eryl Smith 634 (BM, K, SING), 2305 (K).

SIKKIM. Clarke 36800 C, E, F, 37002 G, M (E).

INDIA. Assam: Mann s.n. (B, BO, BRI, E, HBG, L, SING, S-PA, US); Gammie

68 (B, W).

CEYLON. Wail 1012 (S-PA); Thwaites CP 1389 p.p. min. (E), 3389 (B, BM, BO,

E, GH, K, S-PA, W); Ferguson 29 (GH, US).

Not seen from Burma. A specimen marked “‘Hongkong in locis umbrosis silvaticis

Dec. 67 Dr. M. . . [?]” in herb. P probably mislabelled.

33. Lindsaea merrillii Copeland Perkins Fragm. (1905) 181; Tagawa Acta

Phytotax. Geob. 6 (1937) 33, fig. 2 E, F. Type: Merrill 1774, Mindoro, Philippines

(MICH; dupl. in B, GH, K, US).

The typical subspecies is confined to the Philippines. In the present area

only ssp. yaeyamensis is found.

Ssp. yaeyamensis (Tagawa) Kramer stat. nov. Basionym: Lindsaea yaeyamensis

Tagawa Acta Phytotax. Geob. 6 (1937) 31, fig. 2 C, D; Ito Fil. Jap. 111. (1944)

pl. 18. Type: Koidzumi s.n., Iriomote, Ryukyu (KYO, not seen; dupl. in US).

-— LL. macraeana auct. non (Hooker & Arnott) Copeland; Ching Fl. Reip. Pop.

Sin. 2 (1959) 266.

In most respects similar to ssp. merrillii (see Fl. Mal. sp. 48). The petioles

tend to be more rounded abaxially and are sometimes a little longer. The main

difference is in the teeth of the fertile pinnule lobes, which in typical ssp.

yaeyamensis have a minutely erose to subentire apical margin with a tooth at each

end, whereas they are protracted and + dentiform in the middle in ssp. merrillii.

This was described and very clearly illustrated by Tagawa (loc. cit.). The sori of

ssp. yaeyamensis are more often binerval, and the indusia may almost reach the

margin. Some of the specimens from Taiwan are very close to ssp. merrillii.

JAPAN. Ryukyu: Ishigaki, Nishida 374 (US); Kawagoe s.n. (US); Fosberg

37409 (L); Masamune & Suzuki sn (TAI 6198). Iriomote, Walker & Tawada

6733 (BISH, K, L, MICH, US) 6871 (US); Gressitt 580 (B, BM, GH, U); Tagawa

& Iwatsuki 4762 (E,.K,L,U); Masamune s.n. (TAI); Oka 13743 (TOFO); Banda

s.n. (TOFO); Koidzumi s.n. (US, isotype).

TAIWAN. Beattie & Kurihara 1039la (US); Ogata 196 (BM); Tagawa 955 (BM),

976 (K, paratype); Henry 1362 (B); Simizu 2789 (TAI); Hukuyama 664 (TAD);

Kudo & Mori s.n. (TAI 6169); Chuang & Kao 3409 (TAI, U). Botel Tobago, Hsu

s.n. (U); Chuang & Hsu 2452 (TAI); Yamamoto s.n. (TAI 6162). Orchid L, Sasaki

s.n. (TAI 6161); Huang & Kao 7533 (TAI,U).

SECTION Pseudolancea Kramer

34. Lindsaea oblanceolata v.A.v.R. Bull. Jard. Bot. Btzg II. 23 (1916) 15. Type:

Ajoeb 100, Sumatra (BO; dup!. in L). — L. pectinata auct. non Blume; Holttum

Rev. Fl.Mal, 2 (1954) 324.

For further synonymy and description see Fl. Mal. (sp. 45).

Distribution. Malesia, East to the Philippines and the Moluccas.

INDO-CHINA. Annam: Evrard 1295 (MICH, SING); Fleury 38797 (BO).

THAILAND. Eryl Smith 1871 (K, SING); Kerr 7541 (K), 13287 (K); Tagawa,

Iwatsuki & Fukuoka T 4781 bis (U); Smitinand 875 (K, doubtful).

The Lindsaeoid Ferns of the Old World VI 47

35. Lindsaea parasitica (Roxburgh ex Griffith) Hieronymus Hedwigia 62 (1920)

14; Kramer Blumea 15 (1968) 570. — Vittaria parasitica Roxburgh ex Griffith

Calc. J. Nat. Hist, 4 (1844) 510. Type: Roxburgh s.n. Pulau Penang, Malaya (not

seen). — L. scandens Hooker Sp. Fil. 1 (1846) 205, pl. 63 B; Beddome Ferns Br.

India 2 (1868) pl. 298; id. (1892) 74, fig. 37; Holttum Rev. Fl. Mal. 2 (1954) 325,

fig. 186; not of Tardieu-Blot & Christensen Fl. Gén. I.-C. 7 (1939) 121. Lectotype:

Cuming 405, “Luzon” (K). — L. lancea auct. non (L.) Beddome of various authors,

as to Asiatic material, at least in part.

For further synonymy and description see Fl. Mal. (sp. 52).

Distribution. Malay Peninsula, Sumatra, Borneo.

THAILAND. v. Beusekom 804 (L); Tagawa, Iwatsuki & Fukuoka T 5277 (L. UV);

Eryl Smith 780, 1873 (K); Kerr 7909 (K).

DOUBTFUL AND INSUFFICIENTLY KNOWN SPECIES

Lindsaea conformis Ching FI. Reip. Pop. Sin.2 (1959) 372 269. Type: Tsang 24950, Kwang-

tung, China (not seen). — judging from the description this is a form of L. chienii.

Lindsaea longipetiolata Ching Sinensia 1 (1930) 51; Bull. Fan Mem. Inst. Biol. Bot. Ser 10

(1940) 174; Fl. Reip. Pop. Sin.2 (1959) 270. Type: C. C. Chang s.n., Kiangsi, China (not

seen). — This is presumably L. orbiculata var. commixta.

Lindsaea recedens Ching Fl. Reip. Pop. Sin.2 (1959) 373, 269. Type: K. L. Ling 197, Fukien,

China (not seen); also reported from Kwangsi and Japan. — The citation of Ito’s plate

(Ic. Fil. Jap. pl.20) and the description show that this is almost certainly L. chienii.

Lindsaea simulans Ching Fl. Reip. Pop. Sin.2 (1959) 371, 265. Type: several syntype collec-

tions cited from Kwangsi, Kwangtung, and Yiinnan. In the absence of a designated holotype

the name is not validly published. One syntype seen: Lau 2130 from Kwangtung (MICH),

which is L. orbiculata.

Lindsaea taiwaniana Ching FI. Reip. Pop. Sin. 2 (1959) 327, 267. Type: Hancock 10; also 128

(paratype), Taiwan (not seen). — Seems to be a form of L. orbiculata.

Lindsaea tenera Dryander Trans. Linn. Soc.3 (1797) 42, pl.10; non Kaulfuss 1824, nor of

later authors.—Schizoloma tenerum (Dryander) Holttum Rev. Fl. Mal. 2 (1954) 348, as to

type only. Type: ‘Habitat in India Orientali: Missionarii Societatis Unitatis Fratrum” (BM).

As stated before, the plants referred by all later authors to L. tenera do not match the

type (Kramer 1968, where erroneously the name L. cambodgensis instead of L. bouillodii is

adopted for them). The exact provenance of the material being unknown, it seemed that the

identity of Dryander’s type would remain a puzzle. Then three specimens came to the

author’s attention that seemed to shed some light on the problem, as they had several

important characters in common with the type of L. tenera. They are Kurz 26016 from the

Nicobar Is. (K), Kurz s.n. (K) and Parish s.n. (E) from the Andaman Is. The differences

between this material and what has always been called L. tenera (L. bouillodii in the present

author’s concept) may be summed up as follows: petiole stramineous to medium brown (not

dark brown), texture thinly herbaceous with very lax venation (not chartaceous with venation

of average density), pinnules rounded-rhombic, the incisions of very irregular depth (not

rhombic-ovate or dimidiate-ovate and much more regularly incised), the upper pinnules very

little incised, with long sori (this is not the case in L. bouillodii). The terminal segments are

broader and more obtuse than those of L. bouillodii in the above-cited specimens but not in

Dryander’s type.

One of the collectors in the service of the “Societas Unitatis Fratrum’” (= Moravian

Brothers or Herrenhuters) was Koenig, as stated by Burkill (1965) *). As the Society had

contact with the Nicobar Is. at a very early date, Dryander’s specimens may very well have

been collected by Koenig on that archipelago. As the material at hand is too scanty for a

sound judgment of the status of the taxon and its variability, L. tenera is here kept among

the insufficiently known taxa, although it may well be an endemic taxon, though not

necessarily a species, of the Andaman and Nicobar Is.

Lindsaea yunnanensis Ching FI. Reip. Pop. Sin. 2 (1959) 373, 271. Type: K. M. Feng 13646,

Yiinnan, China (not seen). — The description is unfortunately not accompanied by a figure.

This may well be a distinct species, presumably related to L. bouillodii and/or L. chingii.

Vittaria resecta Roxburgh ex Griffith Calc. J. Nat Hist. 4 (1844) 510. Type: no specimen

cited; said to be “nat. of Chittagong”. The extremely brief description is vaguely reminiscent

of L. javanensis which I saw from nearby Assam. It seems fairly certain that it applies to a

species of Lindsaea, Roxburgh having described several other, mostly equally obscure

species of Lindsaea under Vittaria.

*TI am indebted to Dr. F. M. Jarrett, Kew, for helpful advice and the bibliographic

reference pertaining to this problem.

48 Gardens’ Bulletin, Singapore — X XVI (1972)

EXCLUDED SPECIES

Lindsaea lanuginosa Wallich ex Hooker Sp. Fil. 1 (1846) 210, pl. 69 B; Beddome Ferns

Br. India (1892) 77, etc. = Nephrolepis acutifolia (Desv.) Christ.

ACKNOWLEDGEMENTS

The author is greatly indebted to the Director of the Flora Malesiana Foundation who

provided him with the opportunity to undertake the present study. Furthermore he is most

grateful to the Directors and Curators of the herbaria cited in the text who made the material

available to him. Special thanks are due to Dr. Kurata, Institute of Forest Botany, Tokyo, for

sending many important specimens on loan and providing numerous duplicates, and to Dr.

K. Iwatsuki, Department of Botany, Kyoto University, for providing photographs of types.

Grants from the Flora Malesiana Foundation, Leiden, and the Miquelfonds, Utrecht, enabled

the author to visit the herbaria and libraries of Kew, the British Museum, Berlin, and Paris.

Unfortunately the time available for study in the last-named herbarium was insufficient for

studying the whole material, and the citations from Paris are therefore incomplete. Cordial

thanks are also extended to Dr. Ding Hou, Leiden, for kindly providing translations of

otherwise inaccessible Chinese texts, and to Miss E. Hupkens van der Elst who prepared the

illustrations.

REFERENCES

BuRKILL, I. H. 1965. Chapters on the history of Botany in India. Delhi. 245 p.p.

CHING, R. C. (ed.). 1959. Flora Reipublicae Popularis Sinicae II. Peking. xvi +

406 pp.

CHRISTENSEN, C. 1931. Asiatic pteridophyta collected by Joseph F. Rock 1920 —

1924. Contr. U.S. Nat. Herb. 26: 265 — 337, pl. 13 — 29.

CHRISTENSEN, C., & TARDIEU-BLOT, M. 1936. Les fougéres d’Indo-Chine (VI):

Lindsayeae. Not. Syst. 5: 260 — 267.

Cuun, W. Y., CHANG, C.C., & CHEN, F. H. (ed.) 1964. Flora Hainanica J. Peking.

x + 517 + 26 pp., 285 figs. (Pteridophyta by T. T. Wang & C. H. Wu).

DICKASON, F. G. 1946. The ferns of Burma. Ohio J. Sc. 46: 109 — 141.

IWATSUKI, K. 1961. Taxonomic studies of Pteridophyta VI. 7. Schizoloma in Japan

and the adjacent regions. Acta Phytotax. Geob. 19: 1— 8.

KRAMER, K. U. 1957. A revision of the genus Lindsaea in the New World with

notes on allied genera. Acta Bot. Neerl. 6: 97 — 290.

1968. The Lindsaeoid ferns of the Old World. III. Notes on

Lindsaea and Sphenomeris in the Flora Malesiana area. Blumea 15:

557 — 574.

1971. Flora Malesiana Series II — Pteridophyta. Vol. 1, part 3: Lindsaea

Group. Groningen. 78 pp., 56 figs., 10 maps.

KuraTA, S., SATAKE, K., & MASHIKO, S. 1960. The Pteridophyta in the Idzu Penin-

sula, Central Japan. Tokyo Regional Forestry Office. 45 pp.

OxHwI, J. Flora of Japan (in English). Washington, D. C. 1067 pp.

Tacawa, M. 1937. The genus Lindsaya in Japan. Acta Phytotax. Geob. 6: 24 — 41.

1938. On Lindsaya heterophylla Dry. and L. ensifolia Sw. Acta Phytotax.

Geob. 7: 196— 198.

1959. Coloured illustrations of the Japanese Pteridophyta. Osaka. 270 pp.,

72 plates (Japanese).

TARDIEU-BLOT, M., & CHRISTENSEN, C. 1939. (— 1941). Cryptogames vasculaires,

in: LECOMTE, H., Flore Générale de 1’Indo-Chine. VII, 2, 6 (— 9). Paris.

544 pp.

Notes on the Systematy of Malayan

Phanerogams XI- XVII*

from

FOREST RESEARCH INSTITUTE, KEPONG, MALAYA

Abstract

Xylopia malayana var. obscura is a new variety.

The genus Tabernaemontana s.1. is preferred to its segregates.

New synonyms are proposed in Leptopus australis, Glochidion brunneum,

G. penangense, G. wallichianum, G. zeylanicum var. malayanum, Baccaurea

motleyana, Blumeodendron tokbrai, B. borneense and Suregada multiflora.

Glochidion trilobum is a mixtum compositum. Three species are reduced to

varieties: —Austrobuxus nitidus var. montanus, Agrostistachys longifolia var.

leptostachya and Trigonostemon verticillatus var. salicifolius. Endospermum

diadenum (formerly malaccense) has peltate leaves in youth. Alchornea tiliifolia is

annotated.

Scaphocalyx parviflora is reduced to S. spathacea, making the genus monotypic.

Burkilliodendron was published 8 months earlier than Alloburkillia as a new

name for Burkillia Ridley.

Glycosmis calcicola and var. kelantanica are new from limestone.

Pentace excelsa, P. grandiflora and Schoutenia furfuracea are new ‘Tiliaceae

from the east coast.

XI. Annonaceae (b)

K. M. KOCHUMMEN

Xylopia malayana Hk. f. et. Thoms. var. obscura Kochummen var. nov.

A varietas typica in folliis apice latiore breviore, subter nervis lateralibus et

reticuliis obscuris differt.

TRENGGANU: Ulu Brang FRI 12594 (holotype of var. KEP!; A, K, L,

SING).

PAHANG: Raub KEP 20309.

JOHORE: Rengam F. R. FRI 2188; G. Arong F. R. FRI 2761.

XII. Tabernaemontana (sens. lat.), Apocynaceae

T. C. WHITMORE

I follow Hallier (Bot. Jahrb., 49, 1913, 372-5), Merrill (Contrib. Arn. Arb. 8,

1934, 143-4) and Corner (Gdns’ Bull. Str. Settl. 10, 1939, 276) in recognising

Tabernaemontana L. in the broad sense and not its segregates Ervatamia (DC.)

Stapf and Pagiantha Mef.

* Continued from Gdns’ Bull. Sing. 24, 1969, 1-11; Fed. Mus. J. 13,1970,133-7.

50 Gardens’ Bulletin, Singapore — XXVI (1972)

I agree with these authors that there are species in the East, including Malaya,

which transgress the differences between the segregate genera some of which are

weak. There is also an important practical obstacle to the recognition of the

segregates, that without careful re-examination of type material it is not clear

to which segregate genus a species belongs; Markgraf (Notizbl. Berl. Dahlem 12,

1935, 540-52) who favours splitting has not himself seen all the types, few are

represented in Singapore, and three species of Ervatamia (E. curtisii King &

Gamble, E. jasminiflora Ridley and E. pauciflora Ridley) are not represented

at all.

Malesian Tabernaemontana is clearly in need of revision and for the Tree

Flora of Malaya only the trees can be satisfactorily described.

XIII. Euphorbiaceae

T. C. WHITMORE

Minor adjustments to taxonomy which need no commentary appear in the

text of the account in vol. 2 of Tree Flora of Malaya. More substantial changes

and those needing discussion are made below. Other changes have been made by

Airy Shaw and will appear in the series Notes on Malesian and other Asiatic

Euphorbiaceae in the Kew Bulletin. The genera are in the sequence of the

Pflanzenreich monograph.

Leptopus

Leptopus australis (Zoll. & Mor.) Projarkova Not. Syst. Herb. Inst. Bot. Acad.

Sci. URSS 20 (270) 1960

L.. hirta (Ridley) Projarkova loc. cit. 271 sym. nov.

L. calcareus (Ridley) Projarkova loc. cit. 271 syn. nov.

Andrachne hirta Ridley Kew Bull. 1923, 361 (type Ridley 14883 ! K, ! SING)

Andrachne calcarea Ridley loc. cit. 362. (syntypes Ridley 8203, Robinson 6201,

Annandale 1835 ! K)

This genus has been known in Malaya as Andrachne. I am unable to

distinguish Ridley’s two species. The type of L. (A.) hirta is indeed very hairy

(as the epithet implies) on leaf and twig but this condition merges via a long

series of collections into the glabrous condition of L. australis. L. (A) hirta is also

a small-leafed form; this is of no taxonomic significance. L. (A) calcarea has the

leaves rather blunter at base and apex but I can see no sharp distinction against

L. australis. 1 therefore formally propose the reduction of these optimistically

proposed entities. Gage in J. & Proc. As. Soc. Beng. 75 (1936) 522-3 cited all

the collections under Andrachne fruticosa Decne. (non Linn.) which is L. australis

and Henderson (J. Mal. Br. Roy. As. Soc. 17, 1939, 68-9) doubted if Ridley’s

species could be maintained.

Glochidion

Too many species have been based on too few collections. The following

rationalisations are possible.

Glochidion brunneum Hk. f. Fl. Brit. Ind. 5 (1887) 312.

G. goniocarpum Hk. f. loc. cit. sym. nov., type King’s Collector, Singapore (! SING).

G. pedunculatum Ridley Kew Bull. (1923) 364, non Merrill Philip. J. Sci. Bot. 11,

1916, 67 syn. nov., type Ridley 8952 (!SING); 8952 a (! SING).

These two rare species both known from single collections come well within

the range of variation of the commoner G. brunneum.

Notes on the Systematy of Malayan Phanerogams 51

G. penangense (M.A.) Airy Shaw Kew Bull. 23 (1969) 6.

G. villicaule Hk. f. Fl. Brit. Ind. 5 (1887) 326 syn. nov., syntypes Griffith (K.D.)

4842, Scortechini, Perak (! K).

G. coronatum Hk. f. loc. cit. nomen illegit., see Airy Shaw loc. cit., type Wallich

1849 (! K).

I have compared the Perak syntype of G. villicaule with the type of G.

coronatum Hk. f. and can see no difference.

G. trilobum Ridley Kew Bull. (1923) 364.

This is a mixtum compositum. The two syntypes on which the species is

based are correctly placed as follows:

Ridley 8440, Singapore Garden, is G. microbotrys Hk. f., differing only from

typical material of the latter in its densely pubescent fruits, which however is a

condition approached by some of the many sheets of G. microbotrys at K.

Burkill SFN 7004, Negri Sembilan, Tampin, is typical G. wallichianum M.A.

Baccaurea

Baccaurea motleyana (M.A.) M.A. in D.C. Prodromus 15 (1866) 461. B. pubescens

P. & H. Pflanzenreich 1V x V (1922) 251 syn. nov.

B. pubescens is known solely from Wawra 297a, Singapore. No one has

ever found it since and the collection, at Berlin, is now burned. The description

could well be of B. motleyana, as Ridley (Flora 3, 1924, 251) points out, which

is a species found in cultivation throughout the Peninsula and offshore islands.

I agree with Ridley and go one step further in reducing B. pubescens to synonymy.

That Pax and Hoffmann put it in section Calyptroon, not Pierardia, is of no

consequence, for the sections differ in the bracts of the male inflorescence and

Wawra 297a was from a female tree.

Austrobuxus

Two species of Austrobuxus have been recognized in Malaya where they

have been called Longetia. L. malayana (Benth.) P. & H. is a widespread species

of the lowlands. L. montana Ridley was described from 3300 ft, on G. Tahan;

it differs in having smaller, retuse, very leathery leaves and in its typical form

is highly distinctive; I extend its range to G. Benom and G. Ulu Kachau, also in

Pahang, and observe that on G. Jerai (Kedah), at 4000 ft, on G. Padang (Trengganu)

and at 3400 ft in the Ulu Nenggiri (Kelantan) L. malayana grows, so L. montana

does not replace the other in the mountains. Moreover KEP 15085 from near

the sea at Baloh F.R. Pahang approaches L. montana in leaf.

Austrobuxus is an earlier name than Longetia and a move to conserve the

latter failed. L. malayana now becomes A. nitida Mig. For the first time I

transfer L. montana to Austrobuxus, but because it differs solely in leaf characters

which are to some extent overlapping I consider it to be merely a variety; a few

specimens cannot clearly be placed in one taxon or the other.

Austrobuxus nitidus Mig. var. montanus (Ridley) Whitmore stat. nov.

Basionym — Choriophyllum montanum Ridley in J. Linn. Soc. Bot. 38. (1907)

322. Type Robinson 5434.

Synonym — Longetia montana (Ridley) P. & H. in Pflanzenreich XV (1922) 291.

52 Gardens’ Bulletin, Singapore — X XVI (1972)

Agrostistachys

Agrostistachys longifolia (Wight) Benth. Gen. Pl. 3 (1880) 303.

A. longifolia var. malayana Hk. f. Fl. Brit. Ind. 5 (1890) 407.

A. longifolia var. latifolia Hk. f. loc. cit.

A. borneensis Becc. Nelle foreste Borneo (1902) 331.

(1902) 331.

A. sessilifolia (Kurz) P. & H. Pflanzenreich IV 147 VI (1912) 100.

A. latifolia (Hk. f.) P. & H. loc. cit.

The full synonymy is given in the Planzenreich account. Hooker recognized

two varieties and these were later raised to species by Pax & Hoffmann. The

abundant Malayan material now to hand at Kepong and Singapore shows that

there is a continuous range of variation, which also includes the Indian material

at Kew including the type of A. longifolia.

A. longifolia var. leptostachya (P. & H.) Whitmore stat. nov.

A. leptostachya Pax & Hoffmann loc. cit. 102 syn. nov.

This is a very distinctive form with short filiform spikes, which tend to be

in cauline tufts, these features being more marked on male trees, All intermediates

to typical A. longifolia with long, stout, solitary, axillary spikes exist. I therefore

reduce A. leptostachya but retain it as a variety to draw attention to the

polymorphism of the species, which my forest observations indicate might correlate

with geographical distribution, and tree construction. KEP 94699 is almost exactly

intermediate.

A. leptostachya was described from Sarawak and is well represented at Kew.

Typical collections are:

MALAYA, PERAK: FRI 625, 647, 671.

PAHANG: SFN 28763, FRI 4734, 4907, Wong & Wyatt-Smith

35, KEP 8955.

KEDAH: KEP 4641.

Locality unknown: KEP 98304.

Trigonostemon

Trigonostemon verticillatus (Jack) Pax var. salicifolius (Ridley) Whitmore stat. nov.

Trigonostemon salicifolius Ridley Kew Bull. (1923) 366 syn. nov. (type Ridley s.n.

Selangor Batu Caves, ! K).

Ridley’s species seems to be no more than a central Selangor form of

T. verticillatus with leaves oblong (as opposed to elliptic) and often small, for

we now have collections from Kanching, Bt. Lagong and Ulu Gombak as well as

Batu Caves. It is interesting that two collections from the hills of Lower Perak

have oblong (but big) leaves. Varietal status keeps this form in view, and a

biosystematist might find it interesting to investigate it more closely, especially

in view of its disjunct distribution. It grows on hills once lapped by the Pleistocene

sea.

SELANGOR Batu Caves: Ridley s.n. (type) Bt. Lagong: Poore 1062 (K):

Kanching: KEP 21258 (KEP) Ulu Gombak: KEP 18167 (KEP, K), Hume 9015

(SING) PERAK Scortechini 702 (K) (locality?); Briah Larut: Wray 4204 (K,

SING); Telok Kopia F.R. FRI 3112 (KEP).

Notes on the Systematy of Malayan Phanerogams 53

Alchornea

Alchormnea tilifolia (Benth.) M.A.

I reject Evans s.n. Aug 1917 Pahang, Labong, Endau but add Kunstler 166,

mounted at Kew with Scortechini (received 1888), which is A. villosa. Thus the

species is still known from one collection only. The label of the Kunstler sheet

has no locality and merely says Flora of the Malay Archipelago, but is probably

one of his first collections from Johore, G. Panti.

Blumeodendron

Blumeodendron tokbrai (Bl.) J. J. Smith Meded. Dept. Landbouw. 10 (1910)

12, 460.

Mallotus? vernicosus Hk. f. Fl. Brit. Ind. 5 (1890) 443.

B. vernicosum (Hk. f.) Gage Rec. Bot. Svy. Ind. 9 (1922) 244 syn. nov.

B. elateriospermum J. J. Smith Bull. Jard. Bot. Buitenz. sér. 3, 28 (1912) 56.

B. tokbrai is a widespread polymorphic species, first recorded for Malaya by

Henderson (Gdns’ Bull. Str. Settl. 7, 1933, 124) into which merges B. vernicosum.

I can find no disjunctions in habitat or morphology amongst the abundant material

at Kepong to justify retaining two species.

B. borneense Pax & Hoffmann in Engler Pflanzenreich IV 147 XIV (1919) 14.

B. concolor Gage Rec. Bot. Svy. Ind. 9 (1922) 244 syn. nov.

The latter is known in Malaya from only two collections, Curtis 1368 Pangkor

(type !K) and Haniff 15472 Langkawi (!SING, !K). It differs from the type of

B. borneense (Beccari 2976 !K) in its slightly larger, more ovate, more strongly

nerved leaves, but this is well within the range of B. borneense now abundantly

represented at Kew from northern Borneo.

Endospermum

Endospermum diadenum (Mig.) Airy Shaw Kew Bull. 14 (1960) 395.

E. malaccense M.A. Flora 47 (1864) 469.

I am able amply to confirm Airy Shaw’s suspicions that Melanolepis diadena

Miq. Fl. Ind. Bat. Suppl. (Fl. Sum.) (1865) 455 is indeed none other than ‘‘an

extreme, perhaps juvenile, form of E. malaccense’’, which has spreading straight

hairs and no stellate hairs. The confirmation is based on four pieces of evidence:

(1) the, by now numerous, collections made by the Kepong staff of young

E. malaccense.

(2) the rich collection at SING of juvenile E. malaccense made by Corner,

and which Airy Shaw did not see and K has not got in duplicate.

(3) the thorough analysis by Corner in Gdns’ Bull. Str. Settl. 10 (1939) 296

of variation with age of leaf hairiness, an analysis apparently not known

to Airy Shaw when he wrote.

(4) two of the three Malayan collections of E. diadenum cited by Shaw

are indeed from juveniles.

There is therefore no alternative but to rename the common well known

sesendok of Malaya, and use a new name based on a single unattached leaf,

a sterile specimen from a juvenile plant. The rigid application of the rule of

priority in this case is distressing and in the account of Euphorbiaceae for the

54 Gardens’ Bulletin, Singapore — XXVI (1972)

Tree Flora I have no compunction in recommending Malayan foresters to continue

to use the epithet well known for over half a century. My justification for this

course is set out in full in the Prologue to the Tree Flora, vol. 1.

Over the years collections of Endospermum from the Peninsula with peltate

leaves have accumulated. Corner stated that juvenile plants of E. malaccense

never have peltate leaves and referred these collections to E. moluccanum Becc.,

an east Malesian species, from which they differ in not possessing hollow

ant-inhabited twigs. None of these peltate-leaved collections is fertile, all are

from saplings or poles or small to medium trees to 70 ft tall. I consider them

merely a juvenile stage of E. malaccense. FRI 770 from Slim Hills Perak,

50 ft tall, 2 ft girth, seems to clinch this conclusion, it too is sterile and possesses

leaves strongly peltate, weakly peltate and apeltate. Forest observations show

that E. malaccense usually but not always has peltate leaves in youth. At Kepong

we have two seedling collections (FRI 29, 1005) and two collections of 20 ft

saplings (FRI 11, 4787) which are not peltate and FRI 336, 560, 768, 864, 2212,

KEP 98546, 115652 which are strongly peltate.

In conclusion, there is no evidence that E. moluccanum or the peltate Borneo

species EF. peltatum Merr. occur in Malaya. Our material can most simply be

accounted as juvenile E. malaccense.

Suregada

I am unable to detect any consistent differences between the four species

referred to this genus (as Gelonium) by Ridley (Flora 3, 1924, 310-2).

In the first place, Gelonium tenuifolium Ridley must be reduced.

In the second place, the widespread continental species G. multiflorum Juss.

merges into G. glomerulatum Hassk. of the Malesian archipelago here in Malaya

where the ranges overlap.

In the third place, Corner (Gdns’ Bull. Str. Settl. 10, 1939, 299) found

G. bifarium Roxb. impossible to recognize in the Peninsula, and I agree.

I refer all collections made in Malaya to S. multiflora (Juss.) Baill., the

earliest published species, but without study of types and material from the whole

range of the complex can make no formal decision except:

Suregada multiflora (A. Juss.) Baill. Et. Gén. Euphorb. (1858) 396.

Gelonium tenuifolium Ridley J. Roy. As. Soc. Str. Br. 59 (1911) 181 (type Ridley

15902 ! K,! SING) syn. nov.

In addition to the above, S. angustifolia (Miq.) Airy Shaw has once been

collected in Malaya. It is quite distinct.

XIV. Flacourtiaceae

T. C. WHITMORE

Scaphocalyx

Scaphocalyx spathacea Ridley J. Bot. 58 (1920) 149.

S. parviflora Ridley loc. cit. syn. nov.

Most of the flowers at Kepong herbarium have been eaten by insects, but

KEP 99478 from the Bt. Lagong population retains a few buds; some are 3 mm

long on 4 mm pedicels, on another cluster they are 23 mm long on 15 mm pedicels;

the collection thus spans both Ridley’s species and demonstrates that the flower

size difference is accounted for by the different stages of development of the flowers

when collected.

‘

i i i ee es eee ee eee:

Notes on the Systematy of Malayan Phanerogams 55

XV. Leguminosae (c)

T. C. WHITMORE

Burkilliodendron Sastry Bull. Bot. Surv. India 10 (1969) 243.

Burkillia Ridley Fl. Mal. Pen. 5 (1925) 304, non Burkillia West & West (1907).

Alloburkillia Whitmore Gdns’ Bull. Sing. 24 (1969) 4 syn. nov.

Burkilliodendron album (Ridley) Sastry loc. cit.

Burkillia alba Ridley loc. cit.

Alloburkillia album (Ridley) Whitmore loc. cit. syn. nov.

Sastry’s new name for Ridley’s genus was published on 3 Jan. 1969, my

own on 9 Aug. 1969. Ridley describes the plant as a shrub, and the legitimate

name is therefore a misnomer.

XVI. Rutaceae

B. C. STONE (University of Malaya)

Glycosmis

Glycosmis calcicola B. C. Stone, sp. nov. Fig. 1

G. parkinsonii var. ovatofoliolis Tanaka, in herb. KEP and SING. ? Ined.

Frutex inermis ad 1-2 m altis, ramis divaricatis, ramulis gracilibus, 1-2 mm

crassis, juventute tomentellibus, trichomiis rufidulis 1-—3-cellularibus acutis,

maturitate rufo-brunneis glabratis minute striatis; foliis imparipinnatis, petiolo

9-20 mm longo, minute tomentello glabrescentio; foliolis 1-5, alternatis vel

suboppositis, tenuiter coriaceis, ovalis vel obtuse-ellipticis, 1.7—-7 cm long, 0.9-4.3 cm

latis, apice obtuso, vel plerumque emarginato-retuso, base rotundato vel obtuso,

dorso in costa puberulento, costa infra prominulento, supra subdepresso-

canaliculato, nerviis lateralibus utrinque 4-7, lamina minute glanduloso-punctulato.

Inflorescentia rufotomentella, axillaris, parvis, cymosis, 3—7-floriferis, pedunculo

perbreve ad 2 mm longo, pedicellis 1 mm longis minute bracteolatis. Calyx

4-lobatus cupuliformis lobis 1.5 mm longis late deltoideis rotundato-obtusis.

Petala 4 ovata utrinque acuminata alba dorso rufotomentella (plerumque in

portione medio) et ciliolulata, intus glabra, 3.2 x 1.5 mm, Stamina inaequalia,

4 longiora (filamentis 3 mm longis) petalis aequantia, 4 breviora (filamentis 2.5 mm

longis); filamentis oblongis complanatis apice abrupte acuminatis, distaliter (sub

anthero) subsparse puberulo; antheris glabris late cordatis 0.7-0.9 mm_ longis.

Gynoecium disco glabro albo 2 mm alto; ovario lente quadrangulato obclavato

1.7-1.8 mm longo subsparse glanduloso supra medium puberulento; stigma truncato

non expanso 0.2—0.25 mm lato; loculis 4; ovulis elongatis solitariis pendulosis.

Fructus baccatus parvus ad 1 cm longus ovoideus vel subellipsoideus apiculatus,

pericarpio tenue, semina plerumque singulo, cotyledonis viridis crassiter plano-

convexibus punctatis 6 mm longis, plumulo perminute albo-pilosulo.

Holotypus: MALAYA. Selangor: Kanching Distr., Gua Anak Takun, lime-

stone hill near Templer Park, 500 ft. alt., 21 Sept. 1969, B. C. Stone 8789

(KLU; isotypes A, BISH, K, L, SAR, SING, TNS).

Gardens’ Bulletin, Singapore — XXVI (1972)

Fig. 1.

Glycosmis calcicola Stone (BCS 8789, type); a: habit; b: fruit; c: seed; d: hairs;

e: flower, bud and parts; f: ovary.

Notes on the Systematy of Malayan Phanerogams 57

Further specimens examined: MALAYA: Selangor; Bukit Takun, 8 May

1925, Symington KEP 39586 (KEP). Kanching, 16 March 1935, Symington

KEP 37407 (KEP); 5 Nov. 1940, Reid KEP 51681 (KEP), 23 Dec. 1963, Sow

KEP 94745 (KEP). Bukit Takun, 18 Feb. 1937, Symington & Henderson KEP

43289 (KEP). Bukit (Gua) Takun, limestone hill, 900 ft. alt. 11 July 1965,

B.C. Stone, Y. K. Wong, P. C. Lee & G. B. Evans BCS. 5907 (KLU). Gua Batu

(“Batu Caves’’), 28 Dec. 1969, B. C. Stone & P. R. Wycherley s.n. (KLU). Gua

Anak Takun, 31 July 1968, Whitmore FRI 12165 (KEP). Perak, Lenggong,

G. Runta, 300 ft. 11 June 1930 Henderson SFN 23819 (SING KEP). Kedah:

Pulau Langkawi, Kuala Kelim, 4 May 1938, Symington KEP 46792 (KEP), 46794

(KEP). Batu Ayam, 19 Nov. 1941, Corner SFN 37838 (KEP).

This species appears to be distinct from most or all others in the genus by

its small ovate-emarginate leaflets. It is similar in most floral details to G. parkinsonii

Tanaka (of Burma), though the typical variety differs in its glabrous anthers;

in addition, the usually more numerous leaflets and the emarginate or retuse

leaflet tips and different leaflet shape, seem to confer specific rather than varietal

status as concluded by Tanaka. The species apparently is restricted to limestone.

G. calcicola var. kelantanica B. C. Stone, var. nov.

A varietas typica in foliolis plerumque duplo vel triplo magnioribus, petalis

latioribus, antheris pilosulis, differt; cetera simillima.

Holotypus: MALAYA: Kelantan; S. Nenggiri, Gua Jaya at Kuala Jenera,

limestone hill, 27 July 1967, T. C. Whitmore FRI 4255 (KEP).

Further specimen examined; Kelantan, Gua Musang, 15 Aug. 70, Stone 9521

(KLU).

According to Tanaka, there is a variety of G. parkinsonii in Sumatra; I have

not seen this and thus do not know whether it might belong to G. calcicola.

G. calcicola is not uncommon where it occurs. It is probably to be found on

some if not all of the other limestone hills in the country, but many of these have

never been botanized or perhaps even climbed.

The specific epithet calcicola indicates the occurrence of this plant exclusively

on limestone.

Tanaka, who was apparently in control of much information concerning

Glycosmis, published G. parkinsonii Tan. in Studia Citrologica 2: 25, 1928, and

in the Journal of Botany (ed. Rendle) 68: 227, 1930. In both the herbaria of

Kepong and Singapore a number of sheets of the species described above are

named with tickets reading ‘“‘Glycosmis parkinsonii Tan. var. ovatofoliolis Tanaka”

but nowhere can I find any publication of this name. It is not listed in the

exhaustive “Catalog of Genera, Species and subordinate taxa in the Orange

subfamily Aurantoideae (Rutaceae)’’ by Carpenter and Reece (Crops Research,

Agricultural Research Service, U.S. Dept. of Agriculture ARS 34-106, May 1969).

In the Jour. Bot. paper mentioned above, Tanaka says: ‘A trifoliolate variety

occurs in Sumatra, which will be described elsewhere.’ However, the Malayan

plants are not mentioned. It appears that the variety ovatofoliolis is a nomen

nudum appearing only on herbarium labels.

Gardens’ Bulletin, Singapore — XXV]I (1972)

Fig. 2. Pentace excelsa Kochummen (FRI 10880).

Notes on the Systematy of Malayan Phanerogams 59

XVII. Tiliaceae

K. M. KOCHUMMEN

Colona

Colona merguensis (Planch. ex Mast.) Burret Notizbl. Bot. Gard. Berlin-Dahlem

9 (1926) 807.

Columbia curtisii Ridley J.R. As. Soc. Str. Br. 57 (1911) 26 syn. nov.

Columbia diptera Ridley ibid. 59 (1911) 78 syn. nov.

Dr. P. S. Ashton in an unpublished manuscript deposited in the Forest Office,

Kuching has noted this synonymy with which I concur. He observes that ‘Ridley’s

two species are similar to C. merguensis though its type has narrower leaves than

usual. When the achene dehisces adjacent valves from adjacent wings remain

attached. Ridley 15221, type of C. diptera, possesses only these united valves, no

entire achenes, hence Ridley’s observation that the fruit was ‘2 winged’; this is the

only significant difference between it and Curtis 3692 type of C. curtisii’.

Pentace

Pentace excelsa Kochummen sp. nov. Fig. 2

Arbor maxima, 60 m alta, foliis coriacei, ellipticis vel ovato ellipticis, apice

acuminato, basi acuto vel rotundato, supra glabra, subtus perdense stellato-

lepidotis, nerviis lateralibus 4 — paribus; paniculis minute stellato-piloso-lepidotis;

pedicellis conspicuis; phalangiis staminorum 5, staminodiis in basi abrupte rotundato-

Ovatis, parte apicale filiforme; ovario dense stellato-lepidoto, 7-8 costato, stylis

7-8 libris, lepidotis; fructibus 7-8 alatis, minutissime stellato-lepidotis, stylis libris.

TRENGGANU: Ulu Sg. Loh FRI 10686, FRI 10853, FRI 12063 (holotype KEP! ;

ee FRI 12131.

PAHANG: Ulu Sg. Keliu FRI 10880; main ridge G. Tapis FRI 10148, FRI 10149.

P. grandiflora Kochummen sp. nov. Fig. 3

Arbor 27 m magna alta, foliis chartaceis, obovatis vel obovato-ellipticis, apice

acuminato, basi cuneato, supra glabra, subtus glabra, nervis lateralibus 3-paribus,

petiolis gracilis; paniculis paucifloris floribus magnis petalis 11 mm longis stamini-

bus multitudinis (c. 150) libris, ovario 5-costato; fructibus ignotis.

The few large flowers and numerous free stamens at once distinguish this

species.

TRENGGANU: Bt Bauk FRI 3951; Ulu Dungun nr Sg. Bebir (= Ulu Sg. Loh nr

K. Datok) FRI 9667, FRI 9675, FRI 9683 (holotype KEP!; K, L, SING), FRI

12013; State land Jerangau KEP 79785.

Schoutenia

Schoutenia furfuracea Kochummen sp. nov. Fig. 4

Arbor, ramulis juvenilibus dense-pilosis, foliis coriaceis oblongis, 16 — 26.5 X

6.5 — 10.5 cm, apice acuminato, basi inaequali rotundato vel cordato, supra

glabra, costa basim versus dein piloso excepta, subtus dense-pilosis, nervis

depressis, petiolo crasso 4-8 mm longo 3 mm diam; floribus ignotis; calyce in

fructu campanulato, stellato-piloso, interius glabro, disco glabro, staminibus c. 40;

fructibus globosis dense-pilosis stylo glabro.

PAHANG: Rompin, Ulu Sg. Anak Endau FRI 8042 (holotypus KEP!; A,

K, L, SING).

Very distinctive in the red hairy twigs, short stout petioles and large leaves.

60 Gardens’ Bulletin, Singapore — XXVI (1972)

0-0 :

6 4

;

in 4410 cm —

Fig. 3. Pentace grandiflora Kochummen (fruiting twig FRI 12013 flower FRI 9683, type). .

Notes on the Systematy of Malayan Phanerogams

in 4+10

Fig. 4. Schoutenia furfuracea Kochummen (FRI 8042, type); inset: fruit with

ersistent bracts.

Studies in Macaranga V: M. lowii

T. C. WHITMORE

Forest Research Institute, Kepong, Malaya

The following brief note brings the taxonomy of the Malayan species up to

date for vol. 2 of the new Tree Flora of Malaya, now in press. Unfortunately this

discovery was made too late for inclusion in part IV of these studies (Kew Bull.

25, 1971, 237-42).

Macaranga lowii King ex Hk. f., Fl. Brit. Ind. 5 (1887) 453.

M. auriculata (Merr.) Airy Shaw, Kew Bull. 19 (1965) syn. nov.

M. lowii is the commonest member of section Pseudorottlera in Malaya, and

numerous new collections have been made from all over the country in the last

few years. These collections completely break down the subtle distinctions Airy

Shaw uses to distinguish M. auriculata from M. lowii. There is indeed a form with

the leaves drying rather grey-green with rather distant secondary nerves (e.g. FR/

4772, 5520, 6010, 6043, 10514, 10555) and some do have sparse spreading hairs on

the base of the midrib below (e.g. FRI 38159). There are also sheets of M. lowii

with leaves drying reddish brown with rather closer secondary nerves (e.g. FRI

3442, 3986, 5415, 8549, 8898). But numerous intermediates occur with these charac-

ters in different combination (e.g. FRI 5366, 7700, 8338, 10669, 12009). In the

circumstances I reduce the later described species to synonymy.

ee SO

ee ee OO Oe

The Genus Johannesteijsmannia H. E Moore Jr.

by J. DRANSFIELD

Botany School, Cambridge, U.K.

Summary

A revision of the W. Malesian genus Johannesteijsmannia (Palmae) is presented. Of the

four species described, three represent new species.

History

In 1856/57 during his collecting trip in the Padang area of the West Coast

of Sumatra, J. E. Teysmann of the Botanic Gardens, Bogor, Java, made contact

with the Assistant Resident of Lima-puloe district, K. F. Stijman, who subsequently

sent a variety of plants to Teysmann when he had returned to Java. Among these

plants were fruits of a new palm which Stijman presumably described in an

accompanying letter. Teysmann incorporated this collection in his account of his

collecting trip in Sumatra (Natuurk. Tijdscht. Ned.-Indie /4 (4, new series) (1857)

321). In Zollinger’s introduction to the description of this palm by Reichenbach f.

and himself (Linnaea 28 (1856)), the finding of the palm was credited to Teysmann,

who probably never saw it in the field but sent fruit and presumably leaves to

Europe from Bogor; they named it Teysmannia altifrons Reichb. f. et Zoll. in

honour of Teysmann. Miquel, however, described Apocynaceous specimens in

1857 as Teysmannia (versl. Med. K. Akad. Wetens. Amsterdam). The part of

Linnaea 28 containing the description of Teysmannia Reichb. f. et Zoll. (Palmae)

did not appear until 1858. In any case, Reichenbach f. and Zollinger could hardly

have described the palm in 1856 before it had been collected by Stijman and

sent to Bogor. By this time Teysmannia Mig. (Apocynaceae) had been validated

and hence has priority over the palm even though it has been sunk into Pottsia

Hook. et Arn. H.E. Moore Jr. (1961) has clarified this situation and coined the

cumbersome new name Johannesteijsmannia H. E. Moore.

In the type description in Linnaea 28, Reichenbach and Zollinger suggest

that Johannesteijsmannia should be placed next to Salacca and Wallichia and not

far from Nypa, and that it shows affinities with the ‘‘Pandanaceous genus,

Carludovica’’ (Cyclanthaceae).

According to Miquel (1868), the description in Linnaea was based on young

sterile shoots from the Botanic Garden, Bogor (possibly grown from the seed

sent by Stijman) and inflorescences sent from Sumatra, and that the description of

the fruit was erroneous. Miquel (op. cit.) gave an extensive description of the palm,

based on more material from Sumatra, and showed that in fruit structure and

leaf form, Johannesteijsmannia approached the Coryphoideae rather than

Salacca, Wallachia and Nypa.

It has not been possible to find the type specimen of Johannesteijsmannia

altifrons. As Reichenbach’s name is used with Zollinger’s as the authority for the

binomial, although the article in Linnaea 28 is titled ‘‘Uber ein neues Palmenge-

schlecht von der Insel Sumatra, von H. Zollinger” and not ‘‘von Reichenbach and

and H. Zollinger’’, it suggests that Reichenbach was included in the authority as

a courtesy, possibly because the specimen described was in his herbarium. The

64 Gardens’ Bulletin, Singapore — X XVI (1972)

palms at Vienna where Reichenbach’s herbarium was housed, were destroyed by

fire and Prof. Rechinger (in letter) states that the type of Johannesteijsmannia

was evidently lost during this fire. The specimens labelled by Miquel as collected

by Teysmann in Sumatra, formerly in Utrecht herbarium, now in Florence, fit

the original description very well and are taken as representative of Johannesteijs-

mannia altifrons in the present study.

Palm and Jochems (1924) wrote a popular account of Johannesteijsmannia

altifrons in Sumatra with information on the habitat, abundance, uses, and dis-

tribution in Sumatra. Ridley (1925) in his ‘‘Flora of the Malay Peninsula, 5,”

described Johannesteijsmannia and introduced into the description the new

information “stem very short to 7ft tall’. As a result of my researches, I can now

state that the “‘stem short” refers to Johannesteijsmannia altifrons, and “‘to 7 ft tall”

to Jt. perakensis from Gunong Kiedang, Perak.

Beccari (1931) monographed the genus with the other known Asiatic

Coryphoideae.

Johannesteijsmannia, like most palms, has been neglected by plant collectors,

and available herbarium material tends to be badly collected with little or no field

notes.

Johannesteijsmannia H. E. Moore Jr.

Description of the genus

Solitary, acaulescent to shortly trunked forest undergrowth palms. Stem

procumbent and subterranean, in one species (Jt. perakensis) becoming erect at

maturity, marked with leaf scars. Leaves large, usually at least 3 m in length at

maturity, exceptionally 6 to 7 m in length. Petiole extending to the apex of the

lamina as a midrib. Leaf sheaths short, in age rotting to form an interwoven mass

of brown fibres. Petiole channelled above, convex below, armed with short lateral

usually recurved thorns, and often bearing two conspicuous yellow lines along

either side. Scurfy brown indumentum abundant on the abaxial surfaces of the

petiole. Hastula present as a small, dry, brown, dead strip of tissue at the insertion

of the lamina on the petiole. Lamina entire, plicate, diamond-shaped to broadly

lanceolate, the upper margin with shallow lobes showing vestiges of induplicate

splitting. Lower leaf margins continuous with the margins of the petiole and

similarly armed with short recurved thorns. Inflorescence axillary with c.6 tubular,

lobed spathes all but the most distal lying in the same plane of insertion (i.e.

opposing the subtending leaf). Inflorescence branched, with 1 — 5(6) orders of

branching and 3 — 1,000 floriferous branches, with many crowded flowers each

subtended by a bracteole, flowers grouped in clusters of (1)2 —3(4). Flowers

creamy-white, sessile, scented (variously) hermaphrodite. Sepals 3, fused to form

a tube. Petals 3, valvate, fused minutely at the base. Stamens 6, the filaments

connate at their swollen bases to form a ring, minutely epipetalous, abruptly

attenuate, short. Ovary superior, composed of three uniovulate carpels, + free

at the base but connate at the apex to form a common style. Stigma punctiform.

Ovules erect, anatropous, bitegmic, crassinucellate. Fruit usually developing from

one carpel only but occasionally two or more rarely three. Pericarp corky, cracking

into pyramidal warts, endocarp hard, composed of lignified stone cells. Seed

globose, attached at the base, embryo lateral. Endosperm homogeneous and

penetrated within from the chalazal end by a convoluted mass of corky integumental

and chalazal tissue.

SPECIES: 4

DISTRIBUTION: North Sumatra, Malay Peninsula, and Borneo.

mm ee eg cre TT Tee a ee

]

}

| eee:

te 8

ifrl

Plate 1.

THE FRUITS OF JOHANNESTEIJSMANNIA Top left: Jt. altifrons; top right: Jt. peraken-

sis; bottom left: Jt. magnifica; bottom rizht: Jt. lanceolata.

Plate 2. Orang asli house thatched with daun payong Ulu Semenyih, Selangor.

The Genus Johannesteijsmannia H. E. Moore Jr. 65

Uses

The leaf of Johannesteijsmannia makes excellent thatching for aborigine huts

and temporary shelters, either used directly, or sewn into ataps. In the Malay

Peninsula it is unusual to see whole villages thatched with the leaves of Johan-

nesteijsmannia. In Ulu Semenyih, Kajang, Selangor, one such aborigine village

exists, where the thatch consists mostly of Johannesteijsmannia and some

Eugeissona (see Plate 2). In Johore, Chinese logging huts in Mersing, Jemeluang

and Kluang Forest Reserves, are very often walled and thatched with the leaves of

Johannesteijsmannia. In Sumatra, Palm and Jochems (Palm and Jochems 1924)

found the leaves of Johannesteijsmannia, in the form of ataps, used to the

exclusion of the more usual thatching material made from Metroxylon sagu and

Nypa fruticans in the area around Tandjungpura on the Batang Serangan in Atjeh,

North Sumatra. According to these authors, atap of Johannesteijsmannia lasts

from three-four years (depending on the thickness of the thatch and the pitch

of the roof) whereas that of Nypa will last up to six years or more; however,

Johannesteijsmannia thatch cost less than half the price of nipa-thatch.

According to an aborigine in Ulu Semenyih, Selangor, Johannesteijsmannia

thatch in his village lasts three years. Burkill ((1935) 1966) records six years for

nipa-thatch in Malaya and even longer for that of the sago palm.

A single leaf of Johannesteijsmannia makes a excellent umbrella (hence the

Malay name ‘“‘daun payong’’— umbrella leaf).

The colonies of Jahannesteijsmannia altifrons around Wray’s Camp on

Gunong Tahan, Pahang, have provided leaves for wind-shelters for the numerous

expeditions camping at this exposed camp-site. :

The young endosperm is reputedly edible.

Native Names

MALAY PENINSULA SUMATRA

daun payong — umbrella leaf belawan (Miquel 1868, Beccari 1931)

pokok payong— umbrella tree sang (Palm and Jochems, 1924)

daun sal siemienjak boewah maseh (Miquel 1868)

sal sieboesoeng boewah masah (Miquel 1868)

koh

pokok koh BORNEO

pokok sang

daun ekor buaya (Malay) — crocodile’s

tail leaf

sumuruk (Iban)

Natural History

Flowering in the four species of Johannesteijsmannia occurs apparently at

irregular intervals. Gregarious flowering has been observed in Jf. altifrons in

January 1968 at S. Lalang F. R., and Jt. magnifica in May 1968 at S. Lalang.

The flowering of the latter species followed the end of the long drought of January

to March 1968 and coincided with flowering in many trees —it seems reasonable

to suppose this flowering was climatically induced. In the former species it is

not obvious what factor was responsible for gregarious flowering. In all species

during 1967/68 it was unusual not to find one or two plants in flower in the

populations at any given time.

66 Gardens’ Bulletin, Singapore — XXVI (1972)

The flowers of Johannesteijsmannia are variously scented, those of Jt. altifrons

of sourmilk and sewage, those of Jt. perakensis sweetly, those of Jt. magnifica of

Tropaeolum majus, and those of Jt. lanceolata of coumarin. Large numbers of

Nitidulid beetle larvae and adults, Staphylinid beetles, Dipterous larvae, thrips,

ants, termites and spiders can be observed among the inflorescences at anthesis,

and many flowers show signs of being chewed. It is probable that pollination takes

place by the unspecialized process of “‘mess and soil” (Faegri and van der Pijl

1966).

Nothing has been discovered, directly related to the dispersal of the fruit.

Fruit occasionally show signs of having been chewed and it is possible rodents

are of importance in dispersal. During germination of the seed, the cotyledonary

sheath elongates to about 10 cm pushing the seedling into the ground; there is

no cotyledonary ligule and the seedling most closely resembles that of Phoenix

dactylifera (see Gatin 1906). The first photosynthetic leaf resembles that of

Licuala spp. i.e. entire, but with a minutely dentate apex. Tomlinson (1960)

suggests that burying of the seedling in the ground is an adaptation to dry condi-

tions; this seems unlikely in Johannesteijsmannia, and it is not known what

adaptive significance if any, this burying of the seedling in Johannesteijsmannia

has.

The Species of Johannesteijsmannia

Before the present investigations, only one species of the genus had been

recognized (Jt. altifrons (Reichb. f. et Zoll.) H. E. Moore). I have recognized three

new species in Malaya: Jt. perakensis, Jt. magnifica and Jt. lanceolata. It is

thought that these new species have remained unrecognized owing to 1) the rarity

of the species, 2) the general lack of interest in palms, 3) the difficulties of

collection which tend to inhibit botanists from making collections of palms, 4)

the difficulties in representing adequately a palm on a normal herbarium sheet,

rendering many palm specimens virtually useless for taxonomic purposes, and

5) the lack of any extended fieldwork concentrated on the genus over a whole

year.

J. altifrons is widespread but local; it is common in East Johore, and

locally abundant on ridge-tops in the north of Malaya, apparently quite common

in parts of North Sumatra (Palm and Jochems 1924), very local in West Sarawak

and unrecorded, apparently, in Kalimantan, Sabah and Brunei. Jt. perakensis is

known from two parallel mountain systems in Perak, and is the only species

known in this area. Jt. magnifica is known from Ulu Semenyih, Selangor, and

a few hills in North-eastern Negri Sembilan, and Jt. lanceolata is known from

Ulu Semenyih (where it grows with Jt. magnifica and 1.5 km away from Jt.

altifrons). There is an old collection referable to Jt. lanceolata from near

Temerloh in Pahang; Whitmore (pers. comm.) collecting just south of this area

near Temerloh in 1969 was unable to find any daun payong, and aborigines in the

area apparently did not know of its occurrence. However, it may still survive in

the area to the north (see Fig. 1 distribution map).

A Key to the Species of Johannesteijsmannia

(1) Plant tree-like with a trunk up to 4 m tall, inflorescence branches divaricate

- pues amen edie » igs’: ptiaadied’s daney teat + mete ice At Jt. perakensis J. Dransfield

(1) Plant “‘stemless’

The Genus Johannesteijsmannia H. E. Moore Jr.

Pulau Lankawi

Penang ‘ei

Province --~

Wellesley

Distribution of the genus

Johannesteijsmannia in the

Malay Peninsula

+ x PP ®@

Fig. 1.

Thailand

Selangor

Jt. altifrons

Jt. perakensis

Jt. lanceolata

Jt. magnifica

generic records

DISTRIBUTION MAP OF JOHANNESTEIJSMANNIA IN

Singapore

THE MALAY PENINSULA

Pulau

Tioman

68 Gardens’ Bulletin, Singapore — XXVI (1972)

(2) Leaves relatively narrow. 6 —8 times as long as broad. Inflorescence

with 3—6 pachycaul branches, to 1 cm in diameter, bearing spirals

of papillate-petalled flowetsin. fia: 02. is <c00n Jt. lanceolata J. Dransfield

(2) Leaves relatively broad. Inflorescence with 20 or more leptocaul branches

not more than 2.5 mm in diameter. Petals not papillate.

(3) Leaves very broad, with dense grey-white indumentum on the

lower surface of the lamina. Inflorescence with c. 1000 branches,

fruits bright red brown with corky warts 2 — 3 mm high ............

et a. AR ee ee Jt. magnifica J. Dransfield

(3) Leaves glabrous below. Inflorescence with c. 20 — 100 branches,

fruits dull brown with warts 7—9 mm high .....................

cians.) Oh. Oa Jt. altifrons (Reichb. f. et Zoll.) H. E. Moore

Johannesteijsmannia altifrons (Reichb. f. et Zoll.) H. E. Moore Jr. Linnaea 28

657-60 (1856), 1858, Principes 5(4) 116, 1961. Type locality: not known precisely

but probably in the “Lima Puloe” province, Sumatra, West Coast. Stijman —

communicated by Teysmann.

DESCRIPTION

Solitary, acaulescent, undergrowth palm. Stem subterranean procumbent to

15 cm in diameter. Leaves c. 20 — 30 erect, large, to 6 m tall. Petiole to 2.5 m x

2 cm, leaf sheaths to 40 cm rotting to form a network of brown fibres. Petiole

armed with short thorns to 1 mm; petiole with lateral yellow lines prominent

or not. Lamina to 3.5 x 1.8 m, diamond-shaped, with scurfy brown scales along

the costa and at the short marginal lobes. Plicae to 20 or more on either side of

the costa with fine anastomosing veins between the main veins of the folds.

Lower leaf margin armed with short spines, as is the petiole.

Inflorescence axillary, at first erect and then pendulous from the primary ~

axis. Spathes covered in fugaceous scurfy brown hairs, cream at first, then turning —

brown, five-six in number, 10 — 20 cm x 6—8 cm, tubular at the base, inflated ©

above, split on one side, with acute apices. Peduncle 30 — 50 cm long, tomentose,

c. 2 cm in diameter, branching to produce three orders of branches, floriferous

branches 1.5—2.5 mm in diameter, to 100 cm long, c. 20— 100 in number,

greenish and covered in dense white tomentum; densely covered in flowers.

Flowers glabrous, white, + acute in bud, solitary or grouped in 2’s or 3’s, rarely

4’s, borne on prominent tubercles, each subtended by a minute bracteole. Calyx

short, 2 mm high, fleshy, glabrous, shallowly three-lobed. Petals fleshy, white,

tc 4 mm long, twice as long as broad, triangular, fused at the base into short tube

c. 0.5 mm in height. Staminal ring minutely epipetalous, abruptly contracted ©

above into six filaments. Anthers oval. Pollen grains white. Ovary glabrous

with three carpels free at the base, at the apex connate into a common style.

0.8 mm in length, fruit 3.9—4.6 cm in diameter, usually developing from

one carpel, rarely from two—three carpels, covered in 60— 80 brown corky

warts, 6.2—8.2 mm in height. Endocarp woody to 1 mm thick. Endosperm

bony to 2.5 cm in diameter, penetrated at the base by corky integumental tissue.

Embryo lateral.

The Genus Johannesteijsmannia H. E. Moore Jr.

JOHANNESTEIJSMANNIA ALTIFRONS

inflorescence, B: flower

Fig. 2.

70 Gardens’ Bulletin, Singapore — XXVI (1972)

COLLECTIONS EXAMINED

SUMATRA

1 — 'TEYSMANN s.n. Sumatra (ex U. now in Fl). 2 — TEYSMANN s.n. Sumatra

(ex L, now in FI). 3 — No collector mentioned, (June 25/1881) Sumatra

(ex. CAL. in K). 4— TEYSMANN 3/3, Raw, W. Kust Sumatra (BO). 5—S.C.J.

JOCHEMS (1924) Batang Serangan, O. Kust, Sumatra (BO, K, L). 6 — A. H.

BATTEN-PoOLL 5—7, (July 1939) Wassernar, Atjeh, Sumatra (SING).

MALAY PENINSULA

1 — Dr. GIMLETTE s.n. Serasa, Kelantan (SING). 2 — H. N. Ripiey 3123

(1891) Tahan woods, Pahang (SING). 3—R. E. Hottrum /9//8 (1928) Gunung

Muntahak, Johore (SING). 4 — H. N. Ruiprey (1893) Tahan Woods, Pahang

(FI). 5— T. C. WHITMORE, FRI 3854 (1967) Ulu Endau, Johore/Pahang (KEP).

6 — FRI 94873, Gunong Arong Forest Reserve, Johore (KEP). 7 — DRANSFIELD

(1967) Lenggor Forest Reserve, Johore. 8 — DRANSFIELD (1967) Kota Tinggi —

Mersing Road, Johore. 9 — DRANSFIELD 9/6 (1968) Sungei Lalang Forest

Reserve, Selangor. 10 — DRANSFIELD 6/0 (1968) Tahan Woods, Pahang. 11 —

DRANSFIELD 688 (1968) Bukit Koh, Ulu S. Kenyam, Pahang.

BORNEO

1 — O. BeccarI, P. B. 1942 (1866) Mattang, Sarawak (FI). 2 — O. Beccarl, P. B.

3645 (1867) Mattang, Sarawak (FI). 3 — E. F. W. Brunic 47450 (1956) Bukit

Tambi, Bako, Sarawak (SING). 4 — E. F. W. Brunia, S. 9544 (1957) Bako,

Sarawak (SAR). 5 — Carrick AND EnNocuH, JC. 162 (1959) Bako, Sarawak

(SAR, KLU). 6 — J. W. PURSEGLOVE 5538 (1957) Bako, Sarawak (SING). 7 —

DRANSFIELD 760 (1968) Bako, Sarawak.

COUNTRY UNKNOWN

One specimen without label in U, probably from Sumatra.

Besides the above records obtained from herbarium specimens records have

been communicated by Forest Staff, Forest Department, Malaya, Dr. T. C.

Whitmore, and Malay aborigines, and some records have been obtained from

Beccari (1931) and Ridley (1925). All records with localities have been incor-

porated in the map of the distribution of the genus Johannesteijsmannia (see

Fig. 1 distribution map).

HABITAT

Jt. altifrons like the other members of the genus is a plant of primary forest;

it is never found in belukar (secondary regrowth) and it rarely survives any

clear-felling of trees. It can, however, survive in selectively logged forest, but

often sustains considerable damage from falling trees and scorching when exposed

to direct sunlight. Within primary forest Jt. altifrons is a plant of ridge-tops and

hill slopes on well-drained soils, often with a rich mor humus layer and occa-

sionally podsolized. In Johore it occurs at 65 m above sea level on the gently

sloping hills between fresh-water swamps. In Sarawak in Bako National Park,

Kuching, Jt. altifrons occurs as low as 100 m above sealevel. Palm and Jochems

(1924) record it at 25 m above sealevel. These records, however, are exceptionally

low, the majority of populations occurring on hill slopes above 300 m. On

Gunong Tahan, Pahang, Jt. altifrons grows at 1,000 m at the junction of Lower

Plate 3. JOHANNESTEIJSMANNIA ALTIFRONS Slender form from Bako National Park, Sarawak.

i Uy Yj

: o>, L | y Wty Y

Sp LGD: Wy

thoy YU

Wyyg ™

COE

f, #

Plate 4. JOHANNESTEIJSMANNIA ALTIFRONS Northern hill-top form. Bukit Koh, Kuala

Kenyam Taman Negara, Pahang.

WEE

The Genus Johannesteijsmannia H. E. Moore Jr. 71

and Upper Montane forest (sensu Richards 1952, — non Robbins 1968),

growing with Livistona tahanensis, Dipteris conjugata, Weinmannia blumei,

Dacrydium elatum and Podocarpus neriifolius. Another high record for Jt.

altifrons is on Gunong Mandi Angin on the borders of Kelantan, Trengganu and

Pahang where it grows at 1.200 m (Whitmore, pers. comm.). In most of its

localities, Jt. altifrons is found in Hill Dipterocarp forest; in the low-lying Johore

localities it is found in Lowland Dipterocarp forest, and in Sarawak in Kerangas

forest (heath forest) on deeply podsolized soils. In all localities visited, soils

are well drained. However Miquel (1861) records Jt. altifrons in Sumatra as

growing as a swamp-palm, in maritime swamps similar to the habitat of Nypa —

this seems highly unlikely when the Malayan and Bornean habitats are considered.

Palm and Jochems (1924) record Jt. altifrons as occurring abundantly on the

higher ground, but being absent from swamps. Jt. altifrons is not confined to

soils derived from any one rock type, being found on granite-, rhyolite-, shale-,

and sandstone-derived soils. Its main requirements appears to be one of well-

drained soils.

Despite the abundance of apparently suitable habitats, Jt. altifrons is absent

from large areas of Malaya. For example it is absent from more or less the

whole of the Main Range of Malaya except for one record at Sungei Lalang,

Ulu Semenyih, in Selangor. The two valley systems parallel to the Ulu Semenyih

— Ulu Gombak and Ulu Langat — are well botanized and have large aborigine

populations, and if Jt. altifrons grew there, it would have been recorded. Yet

the three valleys are more or less identical in geology, vegetation, and aspect.

The palm is such a conspicuous plant, and so well known to aborigines and

Malaya that it is fairly safe to assume that absences represent real absences, and

not lack of exploration. Similarly Jt. altifrons is absent from large areas of Taman

Negara (the National Park), Pahang, being found only on Gunong Tahan, and

on Bukit Koh near Kuala Kenyam. Bertam, Eugeissona tristis is similarly very

local in the National Park, being known only from Bukit Jeram Panjang on the

Tembeling River.

TAXONOMIC NOTES

Jt. altifrons varies considerably over its geographical range; depredations by

aborigines and Malays for thatching, and forestry practices tend to obscure the

pattern of variation in reducing the leaf size of the palm by leaf cutting and

exposure to greater light intensities. Populations in the North of Malaya, in

Pahang and Kelantan, and in Selangor on hill slopes and ridge-tops, usually

have larger, more numerous leaves, and inflorescences with a larger number of

branches than those of the populations of lowland Johore and Sarawak. It is

sometimes possible to deduce the origin of a particular specimen if it is well

collected. From the photograph in Palm and Jochems (1924) it appears that the

Batang Serangan populations are most similar to those of lowland Johore and

Sarawak. Plants at 1,000 m on Gunong Tahan approach in size those of lowland

Johore but there is an observable gradation in size from large Hill Dipterocarp

forest plants at 500 m to the small mountain forms at 1,000 m, as the mountain

is ascended. Typical mature plants from Hill Dipterocarp forest at Sungei

Lalang F. R., Ulu Semenyih, Selangor, had an average of 28 leaves per plant,

leaves 5.5 m tall, lamina 3.5 m x 1.2 m: in contrast, typical plants from Sungei

Kayu Aru, on the road between Kota Tinggi and Mersing, lowland Johore, had

20 leaves per plant, leaves 3.3 m tall, lamina 2.3 m x 1 m. Despite this variation

in leaf size, floral structure appears to be constant. Owing to the scarcity of

Gardens’ Bulletin, Singapore — XXVI (1972)

1Ocm

JOHANNESTEIJSMANNIA PERAKENSIS

Fig. 3.

inflorescence, B: flower

The Genus Johannesteijsmannia H. E. Moore Jr. 73

flowering material it has not been possible to pursue this problem further. It is

possible that some of the variation in size is environmentally induced; on Gunong

Tahan the soils at 1000 m where the palm occurs are extremely poor podsols

derived from weathering quartzite, whereas lower down the soils are derived

from sandstones and are not podsolized. Some of the variation, however, is

probably genetically controlled; many of the populations are isolated and the

chances of cross pollination with other populations are minute, so circumstances

are ideal for divergence of populations. It is only possible to speculate on the

significance of the variation: unwieldy plants such as Jt. altifrons present too

many difficulties for experimental work.

Johannesteijsmannia perakensis J. Dransfield, sp. nov.

DIAGNOSIS

Jt. altifrons multo affinis, sed differt caule ascendenti trunci instar, inflore-

scentiae ramis in ordinibus quattuor divaricatis, floribus fragrantibus alabastro

rotundatis, fructu maiore verrucis pluribus praedito.

DESCRIPTION

Palma solitaria, aetate caule ascendenti robusto ad 4 m alto et 15 cm

diametro, annulato-cicatricoso. Folia erecta grandia ad 4 m longa. Petiolus ad

1 m longus, 2 cm latus. Lamina ad 3 m longa, ad 1.6 lata, squamis furfuraceis

fuscis intra secus costam et in lobis brevibus marginalibus tecta. Plicae numerosae

ad 20 utrimque, inter plicas nervulis lateralis anastomosantibus. Margo follii

inferior spinis brevibus (ad 1 mm) armatus atque petiolus. Petiolus duobus

striis luteis notatus. Vagina folii fibrosa marcescens reticulum fuscum formans.

Inflorescentia axillaris corona foliorum arcuata. Spathae fugaceo-furfuraceae,

primo cremeae dein fuscentes, 5 —6 numero, 20 cm longae et 8 cm latae, basi

tubulosae superne inflatae, apicibus acutis, imo latere fissae. Pedunculus tomen-

tosus ad 50 cm longus et 2 cm latus, ordinibus quattuor divaricatis ramosus.

Rami floriferi 50— 100, ad 2.5 mm diametro, ad 10 cm longi, viriduli, alboto-

mentosi, floridi. Flores albi glabri, alabastro rotundati, solitarii vel bini vel terni

ageregati, in tuberculis prominentibus insidentes, unusquisque bracteola minuta

subtentus. Calyx brevis ad 1 mm altus, carnosus, glaber, paullo 3 — lobatus.

Petala carnosa alba, ad 3 mm longa, late triangularis basi in tubo breve connata.

Annulus staminalis minute epipetalus, in filamenta 6 aequalia superne abrupte

subulata. Antherae ovales. Grana pollinis alba. Ovarium glabrum carpellis 3 basi

liberis, apice in stylo communi 0.8 mm longo connatis. Fructus ad 5 cms diametro,

plerumque a carpello uno evolutus, raro a 2 —3 carpellis, c. 60 verrucis suberosis

fuscis sordidis ad 9 mm altis, tectus. Endocarpium lignosum ad 1 mm crassum.

Endospermium osseum ad 2.5 cm diametro, basi integumentorum contextu

suberoso penetratus. Embryo lateralis.

Malay Peninsula: Perak, in monte Gunong Kledang in Hill Dipterocarp

Forest. J. Dransfield 871, Holotypus, June 1968; (K) et in monte Gunong Bubu,

Whitmore FRI 0673.

Very similar to Jt. altifrons Reichb. f. et Zoll. but differs from this in the

ascending stem, forming a trunk, inflorescence with four orders of divaricate

branching, sweet-smelling flowers, rounded bud, and the larger fruit with more

humerous warts.

74 Gardens’ Bulletin, Singapore — X XVI (1972)

Solitary palm, in age with a robust ascending stem to 4 m high and 15 cm

in diameter, marked with annular scars. Leaves large, erect to 4 m high. Petiole

to 1 m long, 2 cm wide. Lamina to 3 m long, to 1.6 m wide, clothed with brown

scurfy scales along the underside of the costa, and at the short marginal lobes.

Plicae numerous, to 20 or more on each side of the costa, with small anastomo-

sing lateral nerves between the plicae. Leaf margin armed with short spines

(to 1 mm) as is the petiole. Petiole marked with two yellow lines. Leaf base

fibrous, withering to form a brown network of fibres.

Inflorescence: axillary, arching out of the crown. Spathes covered in fu-

gaceous scurfy scales, at first cream than becoming brown, 5—6 in number,

20 cm long and 8 cm wide, tubular at the base, inflated above, split on one side,

with acute apices. Peduncle tomentose to 50 cm long and 2 cm wide, with four

orders of divaricate branches. Floriferous branches 50 — 100, to 2.5 mm in diameter,

to 10 cm long, greenish, covered in white tomentum, densely covered in flowers.

Flowers glabrous, white, rounded in bud, solitary or grouped in 2’s or 3’s, borne

on prominent tubercles, each subtended by a minute bracteole. Calyx short to

1 mm high, fleshy and glabrous, shallowly three-lobed. Petals fleshy, white, to

3 mm long, broadly triangular, connate in a short tube at the base. Staminal ring

minutely epipetalous, abruptly contracted above into six filaments. Anthers oval.

Pollen grains white. Ovary glabrous with three carpels free at the base, at the

apex, connate into a common style 0.8 mm long. Fruit to 5 cm in diameter, —

usually developing from one carpel, rarely from 2—3 carpels, covered in circa |

60 corky warts, dirty-brown in colour and 9 mm high. Endocarp woody to 1 mm

thick. Endosperm bony to 2.5 cm in diameter, penetrated at the base by corky

integumental tissue. Embrvo lateral.

Malay Peninsula: Perak; on the hills of Gunong Kledang in Hill Dipterocarp

forest. J. Dransfield 871, Holotype, June 1968: (K) and on Gunong Bubu, Whitmore ~

FRI 0673.

COLLECTIONS EXAMINED

1 — H.N. Riprey (1898) Bukit Keledang, Perak (SING). 2 — C. CurtTIs 3350

(1898) Kledang, Kinta, Perak (SING). 3 — R. Lonc (1909) Kledang Hills, Perak

(K). 4 — ex Penang Botanic Gardens (1909) source unknown (K). 5 — Mb.

HANIFF and Mb. Nor, Sing. F.N. 6958 (1920) Sungei Siput, Perak (K, SING).

6 — Macuapo s.n. & s.a. Kamuning, Perak (SING). 7 — J. DRANSFIELD 871]

(1968) Kledang Saiong Forest Reserve, Perak. (K) 8 — T. C. Wuitrmore, FRI

0673, Gunong Bubu, Perak (SING, KEP).

HABITAT

In the Kledang Saiong Forest Reserve near Ipoh, Perak, Jt. perakensis is an —

abundant palm, growing gregariously on hill slopes and ridge-tops from about

175 m to 850 m. At the southern end of the range near Parit, it is equally

abundant on hill slopes and ridge-tops; in the Virgin Jungle Reserve it is con- —

fined to slopes and is absent from ridge-tops. Soils are well drained and derived —

from granite. Much of the Forest Reserve has been selectively-logged, but Jt. ©

perakensis has survived the logging in many places. On Gubong Bubu, Whitmore —

records the palm as locally very common on ridge-tops at 800 m. |

Plate 5. JOHANNESTEIJSMANNIA PERAKENSIS Kledang-Saiong Forest Reserve,

Uy;

Plate 6. JOHANNESTEIJSMANNIA MAGNIFICA (left) AND Jt. LANCEOLATA (right) S. Lalang Forest

Reserve, Selangor. (Photo Courtesy of HO Sai Yuen, Kepong).

The Genus Johannesteijsmannia H. E. Moore Jr. 7s

TAXONOMIC NOTES

Jt. perakensis and Jt. altifrons are more closely related than the other species

of the genus. The separation of Jt. perakensis as a distinct species is justified on

the basis of the characters in the diagnosis (see above) and the great isolation of

the populations. The nearest populations of Ji. altifrons are in (1) West Kelantan

separated from the Perak Hills by 100 km and the intervening Main Range

rising to 2,400 m and nowhere in this region below 1,000 m, (2) Sungei Lalang

in Selangor 200 km to the South, and (3) the Batang Serangan in East Sumatra

300 km to the West and separated by the Straits of Malacca. Isolation was

probably even more effective during the Pleistocene at the maximum sea level,

when, with sea level circa 80 m above present levels (Walker 1954), the Bubu

massif and the Kledang-Saiong massif were probably isolated as islands off the

main Malayan landmass.

In badly collected specimens it is usually difficult to distinguish Jt. perakensis

from Jt. altifrons. Infructescences are usually represented, and these always have

divaricating floriferous branches.

Johannesteijsmannia magnifica J. Dransfield, sp. nov.

DIAGNOSIS

Differt a speciebus ceteris foliis latissimis, lamina infra indumento lacteo

tecta, inflorescentia ordinibus 5—6 ramosa, petalis angustis reflexis.

DESCRIPTION

Palma solitaria acaulescens, caule subterraneo procumbente ad 15 cm

diametro. Folia erecta grandia latissima ad 5 m longa, lamina ad 3 m longa et

2 m lata, superne glabra, infra indumento lacteo vestita, costa marginibusque

lobatis nudis. Plicae numerosae ad 20 utrinque inter plicas nervulis lateralis

anastomosantibus. Margo inferior folii spinis brevibus (ad 15 mm) armatus atque

petiolus. Petiolus duobus striis conspicuis notatus. Vagina folii fibrosa ad 25 cm

longa marcescens reticulum fuscum formans.

Inflorescentia axillaris e cumulo foliorum mortuorum et fragmentorum

plantorum aliorum in corona collectante arcuata. Spathae primo cremeae, dein

fuscentes, 5—6, 26 cm longae et 7 cm latae, basi tubulosae superne inflatae,

apicibus acutis, uno latere fissae indumento albo dense tectae. Pedunculus tomen-

tosus ad 80 cm longus, 2 cm diametro, ordinibus 5— 6 ramosus. Rami floriferi

500 — 1,000, ad 1.5 mm diametro ad 7 cm longi albi glabri, floridique. Flores

glabri, alabastro acuti, solitarii vel bini vel terni aggregati, in tuberculi prominenti-

bus insidentes, unusquisque bracteola minuta subtentus. Calyx brevis ad 0.5 mm

altus, carnosus glaberque, paullo 3 —lobatus. Petala ad 15 mm longa, anguste-

triangularia, 3— plo longiora quam latiora, carnosa alba, reflexa, basi in tubo

breve connata, intra rugosa. Annulus staminalis minute epipetalus, in filamenta

6 aequalis superne abrupte subulata. Antherae ovales. Grana pollinis alba. Ovarium

glabrum carpellis 3, basi liberis, apice in stylo communi 0.4 mm _ longo

connatis. Fructus ad 4 cm diametro, plerumque a carpello uno evolutus aliquando

a 2—3 carpellis, c. 150 verrucis humilis suberosis fusco-rufescentibus, ad 2.5

mm altis, tectus. Endocarpium lignosum ad 1 mm crassum. Endospermium

Osseum ad 2.5 cm diametro, contextu integumentorum suberoso basi penetratus.

Embryo lateralis.

Gardens’ Bulletin, Singapore — XXVI (1972)

Fig. 4. JOHANNESTEIJSMANNIA MAGNIFICA

A: inflorescence, B: flower

The Genus Johannesteijsmannia H. E. Moore Jr. ao

Malay Peninsula: Selangor, in valle humido, Virgin Jungle Reserve Sungei Lalang

Forest Reserve, Semenyih, Kajang, in Hill Dipterocarp Forest. J. Dransfield 862,

Holotypus, May 1968 (K); et Negri Sembilan in monte Bukit Tangga, Seremban,

Napier 1903 (SING) et Dransfield (1967); et in monte Gunong Angsi, Corner.

Differing from the other species by the very broad leaves with the lamina

covered with white indumentum below, the inflorescence with 5—6 orders of

branching, and the flowers with narrow reflexed petals.

Solitary acaulescent palm with procumbent underground stem to 15 cm in

diameter. Leaves erect, large and very broad, to 5 m tall, with lamina to 3 m

long by 2 m broad, glabrous above, on the lower surface covered with white

indumentum except along the costa and at the marginal lobes. Plicae numerous

to 20 or more pairs on either side of the costa, with small anatomosing lateral

nerves between the plicae. Lower leaf margin armed with short spines (to 1.5 mm)

as is the petiole, Petiole with two conspicuous yellow lines. Leaf base fibrous, to

25 cm long, withering to form a brown network of fibres.

Inflorescence axillary, arching out of the pile of dead leaves and other plant

debris accumulating in the crown of leaves, Spathes cream at first then becoming

brown, 5— 6 in number, 25 cm long and 7 cm wide, tubular at the base, inflated

above, split down one side, with acute apices, and densely covered in white indumen-

tum. Peduncle tomentose to 80 cm long, 2 cm in diameter, with 5 — 6 orders of

branching. Floriferous branches 500 — 1,000 in number, to 1.5 mm in diameter

and 7 cm long, white, glabrous, and densely covered in flowers. Flowers white

and glabrous, acute in bud, solitary or grouped in 2’s and 3’s, borne on prominent

tubercles, each subtended by a minute bract. Calyx short, to 0.5 mm high, fleshy

and glabrous, shallowly three-lobed. Petals to 1.5 mm long, narrowly triangular,

three times as long as broad, white and fleshy, reflexed, connate at the base into

a short tube, and rugose within. Staminal ring minutely epipetalous, abruptly

narrowed above into six equal filaments. Anthers oval. Pollen grains white. Ovary

glabrous with three carpels, free at the base, connate at the apex into a common

style 0.4 mm long. Fruit to 4 cm in diameter, usually developing from one carpel,

occasionally from 2— 3 carpels, covered by c.150 low, reddish-brown corky warts,

to 2.5 mm high. Endocarp woody to 1 mm thick. Endosperm bony to 2.5 cm

in diameter, with corky integumental tissue penetrating at the base. Embryo

lateral. Malay Peninsula: Selangor, in humid valley, Virgin Jungle Reserve,

Sungei Lalang Forest Reserve, Semenyih, Kajang, in Hill Dipterocarp Forest. J.

Dransfield 862, Holotype, May, 1968 (K); and Negri Sembilan on Bukit Tangga,

Seremban, Mapier 1903 (SING) and J. Dransfield (1967); and on Gunong Angsi,

Corner.

COLLECTIONS EXAMINED

1 — H. J. Napier (1903) Bukit Tangga, Negri Sembilan (SING). 2 — J.

DRANSFIELD (1967) Bukit Tangga, Berembun Forest Reserve, Negri Sembilan.

3 — WonGc YEW Kwan 95088 (1961) Cpt. 24, Sungei Lalang Forest Reserve,

Selangor (KEP). 4 — J. DRANSFIELD (1967) Cpt. 3, Sungei Lalang Forest Reserve,

Selangor. 5—J. DRANSFIELD 862, (1968) Virgin Jungle Reserve, Sungei Lalang

Forest Reserve, Selangor.

78 Gardens’ Bulletin, Singapore — X XVI (1972)

HABITAT

In all localities visited, Jt. magnifica is a plant of steep slopes and ridge-tops

except in the Virgin Jungle Reserve, Sungei Lalang Forest Reserve where it also

grows at the side of a small river in a very humid valley bottom, alongside Jt.

lanceolata; in this last locality Jt. magnifica occurs in great abundance in the

untouched forest, from the valley bottom at 150 m to the ridge-top at 500 m.

At Bukit Tangga in Berembun Forest Reserve in Negri Sembilan, Jt. magnifica

grows in much disturbed, logged forest.

TAXONOMIC NOTES

As Jt. magnifica and Jt. lanceolata grow side by side in the Virgin Jungle

Reserve (see plate 6) it is possible that hybridization between the two species could

take place. As any hybridization experiments would be extremely prolonged,

difficult to perform, and dependent on simultaneous flowering, it has only been

possible to use indirect evidence to assess the possibility of hybridization taking

place. There are no intermediates observable in mature plants; in young plants

the presence of white indumentum on the lower leaf surface is always correlated

with laminae about 14 times as long as broad. Narrower leaves are always giabrous

except for scurfy brown scales along the midrib. Narrow leaved adults are always

glabrous and always have the inflorescence and fruit characters of Jt. lanceolata.

These facts suggest that hybridization does not occur.

Johannesteijsmannia lanceolata J. Dransfield sp. nov.

DIAGNOSIS

Differt a speciebus ceteris foliis angustis, inflorescentia ramis 3— 6,

crassis, petalis papillatis.

DESCRIPTION

Palma solitaria acaulescens, caule subterraneo procumbente ad 15 cm dia-

metro. Folia erecta, plus minusve lanceolata ad 3.5 m alta. Lamina ad 2.4 m

longa et 30 cm lata, squamis furfuraceis fuscis infra secus costam et nervos

primarios tecta. Plicae numerosae ad 25 utroque costae latere, inter plicas nervulis

lateralis anastomosantibus. Margo inferior folii spinis brevibus (ad 1 mm) armatus

atque petiolus. Petiolus duobus striis luteis conspicuis notatus. Vagina folii fibrosa

ad 25 cm longa marcescens, reticulum fuscum formans.

Inflorescentia axillaris inter petiolos infra cumulum foliorum mortuorum et

fragmentorum plantorum aliorum infossa. Spathae primo cremeae dein fuscentes,

caducae, 3 — 4 indumento fusco sparsim tectae, ad 20 cm longae et 2 cm latae,

basi tubulosae, non inflatae. apicibus acutis, uno latere fissae. Pedunculus

tomentosus ad 25 cm longus, 10 mm diametro, ordine uno ramosus. Rami

floriferi 3 — 6, crassi ad 10 mm diametro et 16 cm longa, pubescentes, arcuati

rective. Flores sessiles albi papillosique, alabastro obtusi, solitarii vel bini vel terni

aggregati, unusquisque bracteola minuta subtentus, spiraliter dispositi. Calyx

brevis, 0.3 mm altus, carnosus glaberque, paullo 3—lobatus. Petala ad 1 mm

longa, late-triangularis, carnosa crassissima, basi in tube breve connata, extus

papillosa, intra apices aliquantum verruculosa. Annulus staminalis minute

epipetalus, in filamenta 6 aequalis superne abrupte subulata. Antherae ovales.

Grana pollinis alba. Ovarium glabrum carpellis 3, basi liberis, apice in stylo

communi 0.3 mm longo connatis. Fructus ad 3.4 cm diametro, plerumque a carpello

uno evolutantes, saepe a 2 — 3 carpellis, c. 90 verrucis humilibus suberosis fusco-

rufescentibus, ad 2.5 mm altis, tectus. Endocarpium lignosum ad 1 mm crassum.

The Genus Johannesteijsmannia H. E. Moore Jr.

Fig. 5. JOHANNESTEIJSMANNIA LANCEOLATA

inflorescence, B: flower

80 Gardens’ Bulletin, Singapore — X XVI (1972)

Endospermium osseum ad 2.5 cm diametro, contextu integumentorum suberoso

basi penetratus. Embryo lateralis. Malay Peninsula: Selangor, in valle humido

prope flumen, Virgin Jungle Reserve, Sungei Lalang Forest Reserve, Semenyih,

Kajang, in Hill Dipterocarp Forest. J. Dransfield 861, Holotypus, May 1968;

(K) et Pahang, in Ulu Sungei Tekal Besar, Temerloh, Henderson March 1923

(SING).

Differs from the other species by the narrow leaves, the inflorescence with

3 —6 thick branches and the flowers with papillate petals.

Solitary acaulescent palm, with subterranean procumbent stem to 15 cm

diameter. Leaves erect, + lanceolate, to 3.5 m tall. Lamina to 2.4 m long and

30 cm wide covered with brown scurfy scales on the underside, along the midrib

and the primary nerves. Plicae numerous to 25 along each side of the costa,

with small anastomosing veins between the plicae. Lower leaf margin with short

spines (to 1 mm) as on the petiole. Petiole with two conspicuous yellow stripes.

Leaf sheath fibrous to 25 cm long, withering to form a brown network of fibres.

Inflorescence axillary buried between the petioles under a pile of dead leaves

and other plant debris. Spathes at first cream, then turning brown, caducous,

3 — 4 in number, sparsely clothed in brown indumentum, to 20 cm long and 2

cm wide, tubular at the base, not inflated, acute at the apex, split down one side.

Peduncle tomentose to 25 cm long, 10 mm in diameter, with one order of

branching. Floriferous branches 3 — 6 in number, thick, to 16 cm long and 10 mm

in diameter, pubescent, curved or straight. Flowers sessile, white and papillate,

obtuse in bud, solitary or grouped in 2’s and 3’s, each subtended by a minute

bracteole, arranged in a spiral on the branch. Calyx short 0.3 mm high, glabrous

and fleshy, shallowly three-lobed. Petals to 1 mm long, broadly triangular, fleshy,

and very thick, connate into a short tube at the base, papillose without, somewhat

verrucose within towards the apex. Staminal ring minutely epipetalous, abruptly

contracted into six equal filaments. Anthers oval. Pollen grains white. Ovary

glabrous, carpels three, free at the base, connate at the apex to form a common

style 0.3 mm long. Fruit to 3.4 cm in diameter, usually developing from one

carpel, often developing from two or three, covered with c.90 low corky warts,

reddish-brown in colour, to 2.5 mm high. Endocarp woody to 1 mm thick. Endo-

sperm bony to 2.5 cm in diameter, penetrated at the base by corky integumental

tissue. Embryo lateral. Malay Peninsula: Selangor in humid valley by the river,

Virgin Jungle Reserve, Sungei Lalang Forest Reserve, Semenyih, Kajang, in

Hill Dipterocarp Forest. J. Dransfield 861, Holotype, May 1968: (K) and Pahang

in Ulu Sungei Tekal Besar, Temerloh; Henderson, March 1923 (SING).

COLLECTIONS EXAMINED

1 — Henderson (1923) Ulu Sungei Tekal Besar, Temerloh, Pahang (SING).

2 — J. Dransfield 861 (1968) Virgin Jungle Reserve, Sungei Lalang Forest

Reserve, Selangor (K).

HABITAT

In Sungei Lalang Virgin Jungle Reserve, Jt. lanceolata is known from the

steep banks of a forest river in a deep humid valley; growing with Jt. magnifica.

Despite the proximity of the river, Jt. lanceolata grows on well drained soils and

not in the low swampy ground at the bends in the river. The population of

Jt. lanceolata was probably once greater—further down the valley scattered

individuals occur growing in logged forest near the river.

The Genus Johannesteijsmannia H. E. Moore Jr. 81

The Geography of Johannesteijsmannia

Johannesteijsmannia is confined to North Sumatra, the Malay Peninsula,

and West Borneo, with one widespread species Jt. altifrons, and three species

endemic to the Malay Peninsula. The concentration of species in the Malay

Peninsula may be a reflection of the more intensive fieldwork carried out in

- Malaya; there are, for example, as far as in known, no post 1940 specimens of

Jt. altifrons from Sumatra, and the possibility of new species of the genus occurring

in Sumatra cannot be ruled out. Jt. altifrons, however, in Sarawak appears to be

genuinely rare, and Forest Staff and Ibans in Brunei did not know of the existence

of this very conspicuous and useful palm.

It seems most reasonable, however, to suppose that the centre of diversity

of the genus coincides with present day centre of distribution —1i.e. the uplands

of Selangor, Negri Sembilan and Pahang, where three of the four species grow

together. In late Mesozoic time this area was somewhere near the centre of the

Sundaland mass (van Bemmelen 1949, Umbgrove 1949), and this area together

with West Sarawak, East Sumatra, Banka and Billiton, has remained above sea

level until the Pleistocene marine incursions (Umbgrove l.c.) when it seems

probable (extrapolation from Malaya) that much of Banka and Billiton and of

lowland East Sumatra would have been submerged during the maximum sea

level of 80 m (Walker 1954). It is the area of Sundaland now represented by

Malaya and Borneo which probably remained as land through the Pleistocene, that

harbours the greatest concentration of species in the palm genera Licuala, Calamus,

Daemonorops, and Korthalsia. (the number of species of Licuala on New Guinea

exceeds the number in Malaya and Borneo individually, but if Malaya/Borneo

is treated as one land mass, i.e. part of Sundaland, then the total exceeds that of

the land mass of New Guinea and its outlaying islands).

The limited distribution of the genus can be explained in two main ways: (1)

the genus is an ancient one and survives as a relatively unsuccessful form in the

area of its diversity as a relic of Coryphoid evolution, (2) the genus has relatively

recently evolved and has not yet spread to areas outside its area of origin. That

Johannesteijsmannia has many characters in common with Licuala but is in many

ways more primitive than Licuala (e.g. leaf and inflorescence structure) suggests

it is probably more ancient than the successful widespread genus Licuala. It

therefore seems more likely that Johannesteijsmannia is an ancient genus surviving

in Sundaland. Speculating further I would suggest an origin during the Mesozoic

from primitive Coryphoid stock.

If a Mesozoic origin of the genus in the Malay Peninsula is assumed then the

spread away from the centre could have occurred any time during the late Mesozoic

and the Tertiary. Any pre-Miocene invasion of Sumatra would probably have been

destroyed by marine incursions, for much of Sumatra was submerged during the

Miocene (Umbgrove 1949). Only one species at the present day, Jt. altifrons, has

spread out of the proposed centre of diversity, and is therefore to be regarded as

the most successful and adaptable species. Jt. altifrons probably reached Sumatra

during the Pliocene, and was then cut off from the Malayan populations by the

appearance of the Straits of Malacca. Jt. altifrons could have reached Sarawak

any time before Borneo was separated from the Malay Peninsula during the

Pleistocene, but the very local occurrence of the species in West Sarawak suggests

82 Gardens’ Bulletin, Singapore — X XVI (1972)

that the invasion of Borneo is relatively recent, (e.g. in the Pliocene corresponding

with the proposed time of spread into Sumatra). Jt. altifrons, although it has reached

Sarawak, is absent from Pulau Tioman (Pahang), Penang and the Anamba and

Natuna Islands; the absence may be due to chance extinctions if the palm

ever colonized these islands or else it was unable for some reason to reach the

islands.

The distribution of Jt. altifrons within the Malay Peninsula is disjunct and it

seems reasonable to suppose that Jt. altifrons was at one time widespread over the

whole area; this may have coincided with the proposed period of active expansion

which resulted in the colonization of Sumatra and Borneo. The disjunction of the

range of distribution has resulted in slight vicarous evolution; the isolated popu-

lations have diverged slightly from each other. In Perak, populations on the Bubu

Range and Gunong Kledang have diverged further and reached a degree of

divergence where it is convenient to regard them as a distinct species, Jt.

perakensis. The evolution and distribution of Jt. lanceolata and Jt. magnifica can-

not be explained by vicariism (as Croizat 1968 would explain the speciation of

Ficus — refuted by Corner 1969); the two species occur alongside Jt. altifrons

in the one Forest Reserve in Selangor, apparently with the same ecological require-

ments. It is most reasonable to regard Jt. lanceolata and Jt. magnifica as rela-

tively unsuccessful early products of form making within the genus which have

survived near the centre of diversity without having spread. (Jt. lanceolata has

reached central Pahang from the proposed centre of diversity).

The dispersal powers of Jt. altifrons appears to be very limited — seedlings

are rare, — and the distribution of the whole genus suggests plants of very limited

dispersal powers at the present day, surviving as relics in the centre of massing

of the Asiatic coryphoid palms, as do Liberbaileya gracilis and Maxburretia

rupicola (Furtado 1941).

Acknowledgements

This study was carried out during the tenure of a Royal Society Leverhulme

Scholarship and a Science Research Council NATO studentship, and represents

part of a thesis submitted in Cambridge for the degree of Doctor of Philosophy.

I should like to thank Professor E. J. H. Corner for supervising my work,

Dr. T. C. Whitmore for an inestimable amount of help in Malaya and England,

and all the people, too numerous to mention, who helped me in my field work.

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van Sumatra. Natuurk. Tijdsch. Ned. Indié 14, 320 — 22.

Tomlinson, P. B. (1960): Essays on the Morphology of Palms. 1. Germination

and the seedling. Principes 4, 56 — 6l.

Umbgrove, J. H. F. (1949): Structural History of the East Indies. Cambridge.

Walker, D. (1954): Studies in the Quaternary of the Malay Peninsula (1) Alluvial

deposits of Perak and Changes in the relative levels of land and sea Fed.

Mus. 1. i +. 2, 19 — 39.

Zollinger, H. (1856) (1858): Uber ein neues Palmengeschlecht von der Insel

Sumatra. Linnaea 28, 657 — 60.

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Soil moisture under forest, Bukit Timah Nature Reserve,

Singapore

By R. D. HILyi

University of Singapore

Abstract

The results of a study of moisture of three forested hill-slope soils are

presented and compared with the results of a similar study in Malaya.

Soil moisture levels were found in general to be rather higher than

under other forms of vegetation. Statistical analysis indicated that change

of moisture content was consistent within each soil but less consistent

between soils. A series of statistical experiments suggested that except

for the A soil horizon, soil moisture (as determined by drying) correlated

reasonably well with number of raindays and that the introduction of

the factor e (evapotranspiration) resulted only in slightly improved

correlations except for the A soil horizon where the use of e gave much

better correlations.

The study of soil moisture in the region is one that is strangely neglected,

a search of available literature failing to reveal more than two studies, both

concerned with soil moisture as related to rubber. Belgrave (1930, 493-4) made

observations from three levels, 2-6 in., 6-12 in. and 12-18 in., of a Serdang

series soil having two types of cover, Centrosema, Mikania, and under bare soil.

Watson, Wong and Narayanan (1963) made eighteen observations at monthly

intervals from two levels, 0-6 in. and 12-18 in. of a Rengam series coarse sandy

clay loam with a vegetation of two and three year old rubber and cover crops

of legumes, grass, adventives and also under bare soil.

Soil Characteristics

The present study reports the results of weekly observations made from

mid-February until mid-June 1970 at three points down a surveyed transect line

off Jungle Fall path, Bukit Timah Nature Reserve which has a somewhat open

lowland Dipterocarp-forest. The upper sampling point, 1, (172 m a.s.l.) was some

25 m from the crest of a gently-rounded interfluve and about 4 m below it with

a local slope of almost 15°. The middle sampling point, 2, (141 m a.s.l.) was about

87 m downslope from the first sampling point, 1, and it had a local slope of 24°.

Both soils belong to the Rengam series (Leamy & Panton 1966, 105). The lowest

sampling point, 3, (126 m a.s.l.) was at the foot of the same slope, 33 m from 2,

and located on a small terrace 1.5 m above the level of a small perennial stream.

The soil materials were derived from colluvium and alluvium with the former

predominating. This soil cannot be assigned to any described series.

86 Gardens’ Bulletin, Singapore — XXVI (1972)

The moisture-holding capacity of soil is significantly influenced by grain-size

and organic material present. These data are presented for four levels in each

soil, 0-10 cm (04 in.), 10-20 cm (4-8 in.), 30-45 cm (12-18 in.) and 60-75 cm

(24-30 in.), in Table 1. Detailed analysis of grain size was made using the

A.S.T.M. standard method (American Society for Testing and Materials, 1963).

Only a summary of the data is presented here.

The data for loss on ignition (at 480°C) do not accurately measure total

organic matter because the initial oven-dry samples still retain tightly-bound

water of hydration. Total organic matter is thus slightly lower than indicated in

Table 1.

Table 1. Grain Size (%) according to International System and Loss on Ignition

Sample Depth Coarse Sand Fine Sand Silt and Clay Loss on

: Textural name Ignition %

Point (cm) —_(2000,-250.) (250-25) (less than 25,,) pecan

1 0-10 44.6 16.7 38.7 Coarse sandy loam 7

10-20 39.3 12.9 47.8 Coarse sandy loam* 8.8

30-45 38.2 13.3 48.5 Coarse sandy loam* is

60-75 34.1 22 DAGL Coarse sandy loam 7.0

2 0-10 40.3 21.9 37.8 Coarse sandy loam 10.3

10—20 Bd nl i338 47.0 Coarse sandy loamt 5.8

30-45 36.2 18.1 45.7 Coarse sandy loam+ 4.5

60-75 33.5 20.6 45.9 Sandy clay loam a7

3 0-10 60.0 5.9 34.1 Not namedt 52

10-20 57.6 20.8 21 & Coarse sandy loam 5.1

30-45 56.7 16.1 26.3 Coarse sandy loam 4.4

60-75 38.5 17.5 440 Coarse sandy loamt fe

* Almost sandy clay

+ Almost sandy clay loam

t Suggested name: coarse sandy clay

Collecting Methods

Prior observation established that the maximum variability amongst samples

taken from the same level at the same point on the slope was + one per cent

provided that augered samples were taken at points where there were no large

roots and which were some distance away from the stems of nearby trees.

Successive weekly samplings thus required only one sample from each level at

each observation point. Several samples taken at point 3 were notably coarser

than the majority and values for moisture are therefore less reliable.

Samples of about 300 gm were augered from each level and immediately

placed in polythene bags and sealed. On arrival at the laboratory samples were

well-shaken or stirred to incorporate moisture condensing on the inside of the

bag and to break down lumps. Samples of about 100 gm were then spooned into

dishes for drying at 105°C for 24 hours followed by weighing at 40°C. Prior

experiment showed that differences arising from more protracted drying and

from cooling in a desiccator to room temperature were much smaller than the

differences arising from variations in the exact point sampled. It was however,

necessary to weigh dried samples very rapidly taking them one by one from

the oven.

LS _eset eee

Soil Moisture under Forest, BT Nature Reserve, S’ pore 87

Weather

The period of observation began with the dry spell which usually occurs

towards the end of the north-east monsoon, and extended through a wetter

inter-monsoon period to the early part of the south-west monsoon. Rainfall data

were collected daily from a rainfall station some three hundred metres from the

soil sampling points. Open pan evaporimeter data were obtained for Paya Lebar

Airport some 14 km from Bukit Timah. Soil samples were taken shortly after

the daily rainfall measurement was made. Rainfall data are presented at the top

of each diagram in Figures 1-3.

Results

The results are presented in Figures 1-3. The diagrams suggest a fairly clear

relationship between rainfall and soil moisture, the general trend of low values

during dry spells being clear. In the two hill-slope soils points 1 and 2, soil

moisture conditions were fairly stable below 30 cm with increasing variability

in the upper portions of the soil. This was more marked in the 0-10 cm zone

which corresponded with the A, soil horizon. Rather large fluctuations, sometimes

even during fairly dry periods reflect large but short-lived rises of soil moisture

following showers.

The mid-slope soil was slightly drier than the upper-slope soil during the

main dry spell but slightly wetter during the remainder of the period. This

suggests that greater slope angle and hence more rapid soil drainage becomes

influential only during dry spells whereas at other times receipt of upslope

water results in slightly moister conditions. Differences of soil texture would

seem to be too minor to account for these differences. Furthermore the mid-slope

soil contained rather less organic matter than the upper-slope soil and this should

have resulted in lower moisture values for the former soil had this been an

influential factor.

The slope-foot soil shows an almost complete reversal of the pattern of the

other two soils. In it variation was greatest at depth, undoubtedly because of

fluctuations in the position of the water-table. The upper zones of this soil were

a good deal sandier than those of the hill-slope soils, organic matter was much

less and rapid infiltration of rainfall and overland flow from upslope occurred.

The range of moisture is summarised in Table 2.

Discussion

Only the study by Watson, Wong and Narayanan (1963) presents data

adequate for some comparisons to be drawn. Expectably the soil moisture values

are considerably higher under forest than those under legumes, grass, adventives and

bare soil reported in that study. The lowest values at Bukit Timah are roughly

equivalent to the mean values at Serdang for the upper 20 cm of soil. The data

of Watson et al (1964, 165) also show much greater variation of moisture in

the 0-15 cm zone than in the 30-45 cm zone. They report moisture values for

the 30-45 cm zone which correspond quite well with those of the present study.

oe)

Gardens’ Bulletin, Singapore — XXVI (1972)

0090

15 05 #

30 1-0 2

1-5 z

2-0

60 2

75 =

305

90 2

seltantr isle bhnials oO] 4” (0 -10¢cm.)

30 mi CU iets See 4- 8" (10-20cm.)

: 12-18" (30- 45cm.)

DAILY RAINFALL (MM.)

24-30" (60-—75cm.)

LOSS OF WEIGHT (*%.)

ceca eaieamam) Soammaieien®| ‘nearer Giese Tiers (Alia ||" ieee [ieee en Dees |? ima: aanann aaa aM! (Pitan Eels ep See

4 11 18 25 4 1113 18 24 1 7 15 22 29 6 13 20 27 - 10 “f

FEBRUARY MARCH APRIL MAY JUNE

Fig. 1 Soil moisture (loss of weight on drying) for the soil near the top of the slope (1) and

daily rainfall.

Table 2. Range of Soil Moisture (%)

Sample Depth Date Date

Ban. ie Highest (1970) Lowest (1970) Range

] 0-10 30.2 18.-2 2h3 4-3 8.9

10-20 26.7 22-4 19.9 114 6.8

30-45 24.7 13-5 19.5 11-3 5.2

60-75 23.5 13--5 20.2 24-3 3.3

2 0-10 31.6 aah 20.0 113 11.6

10-20 218 10-6 17.9 11-3 99

30-45 23,5 13-5 18.6 4-3 49

60-75 26.4 22-4 20.1 11-3 6.5

3 0-10 23.4 22-4 18.5 4-3 4.9

10-20 22.9 22-4 20.2 25-2 ay

30-45 PAS Be 22-4 10.1 13-5 6.7

60-75 30.4 22-4 19.7 1-4 10.7

Statistical Analysis

Two sets of statistical analyses of the hill-slope soils (1 and 2) were made.

The first aimed to establish the extent to which the rate of change of soil moisture

between successive observations at each level within the soil were correlated.

The results are summarised in Table 3 which shows that in general there is a

greater degree of association between moisture change at one Jevel within the

soil and moisture change at the level below it than between moisture change at

the same level of different soils. In other words rates of change within a soil were

internally consistent with the notable exception of the soils at 60-75 cm where

poor or no correlations with the soils at 30-45 cm were found.

Soil Moisture under Forest, BT Nature Reserve, S’ pore 89

2 00 9

= 15 0-5 e

e 30 1:0 >

45 re z

z 20 >

é 60 25°

7 75 3-05

& 90 35 2

> 30 .

Oe ee O=- 4" (O0-10cm.) :

eo oqyoccccce 4- 8" (10-20cm.) :

E 12-18" (30-—45cm,) :

Ww ——$——_— 24-30" (60-75¢m.) : TS ee ore

3 25 é a ny i

4 :

re)

}

a ee eae fee a eee ee i . ahit«l: i. Tr 1

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“FEBRUARY MARCH APRIL MAY JUNE

Fig. 2 Soil moisture (loss of weight on drying) for soil on the middle of the slope (2) and

daily rainfall.

DAILY RAINFALL (MM.)

wo ~ oa > w ak

oO o oO oo o oO

Ww WwW RH A _ — o Oo

omovwvoWws#owso

CSNI) WIW4NIv¥ Ativa

2 Ree O- 4” (0-10cm.)

ee ee 2 es 4- 8” (10-20cm.)

12-18" (30-—45cm.)

24 -30" (60-75 cm.)

(*/e)

OBSERVATIONS

DISCONTINUED

LOSS OF WEIGHT

[oF SA ae EEL et el a ee Tis Tepe ea cia iets eae mea area Bees lee

- 11 18 25 & 11:13 18 24 1 7 15 22 29 6 13 20 27 . 10 17

FEBRUARY MARCH APRIL MAY JUNE

Fig. 3 Soil moisture (loss of weight on drying) for the slope-foot soil (3) and daily rainfall.

90 Gardens’ Bulletin, Singapore — X XVI (1972)

The second set of statistical analyses attempted to find a simple parameter

with which soil moisture was correlated.

Since soil moisture is affected by a large number of factors including

endogenous factors such as grain size and structure and exogenous factors such

as rainfall and evapotranspiration, it was hardly to be expected that with only

19 successive observations statistically significant correlations between soil moisture

and rainfall alone would be found. Such proved to be the case. Soil moisture

was compared with the total rainfall over varying numbers of days prior to

each observation. The best, but still poor correlations were first, for moisture in

soils 1 & 2, (10-20 cm), with total rainfall over the fourth, third and second days

before observation; r = + 0.47, and second, for moisture in soils, 1 & 2 (60--75 cm),

with total rainfall over the fifth, fourth and third days before observation;

r = + 0.47, both values for r being significant at the 5 per cent level with 24 degrees

of freedom. Student’s ¢ test was used to test significance in each case.

Table 3. Correlations of Rates of Change of Soil Moisture at Various

Levels in Soils 1 and 2

Variables Degrees of Significance

¥ y freedom / (better than)

1 (0-10 cm) 2 (0-10 cm) 16 +0.76 i%

1 (10-20 cm) 2 (10-20 cm) 18 +(0).49 5%

1 (30-45 cm) 2 (30-45 cm) 18 + 0.57 hy

1 (60-75 cm) 2 (60-75 cm) 18 + 0.56 1%

1 & 2 (0-10 cm) 1 & 2 (10-20 cm) 34 +0.71 0.1%

1&2(10-20cm) 1&2 (30-40 cm) 36 +0.70 0.1%

1&2(3040cm) 1&2 (60-75 cm) 36 + 0.46 19,

1 (0-10 cm) 1 (10-20 cm) 18 +(0.71 0.1%

1 (10-20 cm) 1 (30-45 cm) 18 +0.76 0.195

1 (30-45 cm) 1 (60-75 cm) 18 + 0.68 1%

2 (0-10 cm) 2 (10-20 cm) 16 + (0.73 1%

2 (10-20 cm) 2 (30-45 cm) 18 + 0.60 1%

2 (30-45 cm) 2 (60-75 cm) 18 + 0.38 None

A similar comparison of rate of soil moisture change with rainfall was even

less satisfactory

A comparison between soil moisture and number of raindays (> 0.1 in. or

2.5 mm per day) in the week preceding measurement gave rather better results

and these are shown in Table 4 together with correlations of rate of moisture

change with number of rain-days in the preceding week.

Soil Moisture under Forest, BT Nature Reserve, S’ pore 9]

Table 4. Correlation of Soil Moisture and Rates of Soil Moisture at Various

Levels with Number of Rain-days in Preceding Week

Soil Degrees of Significance

x y freedom is (better than)

Soil m, 1 & 2, (0-10 cm) t/days 38 + 0.44 jibes

Soil m, 1 & 2,(10-20cm) ___r/days 38 + 0.56 0:1:%

Soil m, 1 & 2, (30-45 cm) r/days 38 +0.53 0.1%

Soil m, 1 & 2, (60-75 cm) r/days 38 +0.55 Oke,

Rate of s.m. change,

1 & 2, (0-10 cm) t/days 34 + 0.56 0.1%

Rate of s.m. change,

1 & 2, (10-20 cm) r/days 36 +().49 1%

Rate of s.m. change,

1 & 2, (30-45 cm) r/days 36 +(0).47 oe

Rate of s.m. change,

1 & 2, (60-75 cm) r/days 36 + 0.47 1%

In an attempt to gain rather better correlations the factor for evapotranspira-

tion (e) was introduced. In this, e¢ is evaporation, multiplied by the empirical

constant 0.9. The actual formula used was R — e, where R = daily rainfall and

e the daily evapotranspiration. Values of R—e were summed for varying numbers

of days prior to each determination of soil moisture. The selected best results are

given in Table S. ;

Table 5. Selected Correlations of Rates of Change of Soil Moisture with R — e

over Various Days before Moisture Determination

si Bnei wcleses tine tro > teen in

1 & 2, (0-10 cm) 3,4204 34 +0.57 Pes

1 & 2, (0-10 cm) I, say 34 + 0.60 1%,

1 & 2, (0-10 cm) dei44 3 34 +0.45 3%

1 & 2, (10-20 cm) 4, 3,2 36 + 0.44 3%

1 & 2, (10-20 cm) 5, 4, 3 36 +0.31 Nene

1 & 2, (30-45 cm) 3, 4.3 36 +-0.37 None

1 & 2, (60-75 cm) >, 4, 3 36 + 0.42 5%

It will be seen from Table 5 that the introduction of the factor e which might

have been expected to have added precision to the correlations failed to do so

except in the case of the uppermost layer of soil from 0-10 cm.

In order that higher values for r might be obtained the empirical formula

Rw — e was used. It was felt desirable to retain the factor e since it represents

a continuous process which results in loss of soil moisture. It was also desirable

to weight the value R since rain falling seven days before a soil moisture deter-

mination would have much less effect upon the soil moisture level than rain falling

92 Gardens’ Bulletin, Singapore — X XVI (1972)

in the day before the measurement. Rainfall was weighted with the following

values, corresponding to 1, 2, 3... 7 days before measurement; 1.0, 0.93, 0.85,

0.74, 0.60, 0.40, 0.10 which approximate to an exponential curve. The values for R

weighted —e were then summed for varying periods prior to each moisture

determination and separately correlated with soil moisture and rate of change

of soil moisture. Selected results, giving highest values for r, are given in

Tables 6 and 7.

Table 6. Selected Correlations of Soil Moisture with R weighted—e, over

Various Days before Moisture Determination

Degrees of Significance

Soil Days freedom r (better than)

1, (0-10 cm) : ee 19 + 0.46 5%

1, (10-20 cm) S. «ett 19 + (0.67 1s

1, (30-45 cm) | 19 + 0.66 1%

1, (60-75 cm) Dieneue 19 +0.60 1%

2, (0-10 cm) SS¢ 1 19 + 0.67 hy 3

2, (10-20 cm) ys l 19 +0.57 1%

2, (30-45 cm) fF 3 2 19 +0.57 1%

2, (60-75 cm) (lke = 19 + 0.62 1%

1 & 2, (0-10 cm) a 37 + 0.56 =

1 & 2, (10-20 cm) a as 38 +0.59 1%

1 & 2, (30-45 cm) ( pre 38 +0.59 1%

1 & 2, (60-75 cm) A eucasd 38 +0.57 1%

Table 7. Selected Correlations of Rate of Soil Moisture Change with R weighted

— e over Various Days before Moisture Determination

Degrees of . Significance

Soil ae freedom : (better than)

1, (0-10 cm) ie l 18 + 0.65 1%

1, (10-20 cm) aie | 18 +0.61 1%

1, (30-45 cm) Ry 2 18 + 0.63 1%

1, (60-75 cm) yee 2 18 + 0.40 None

2, (0-10 cm) Ds | 18 + 0.67 1%

2, (10-20 cm) a l 19 +-0.57 hy 4

2, (30-45 cm) aa 2. 19 +0.57 1%

2, (60-75 cm) Pi 2 19 +0.62 1%

1 & 2, (0-10 cm) ~ Sy l i +0.56 r%

1 & 2, (10-20 cm) Deak l 38 +0.59 0.1%

1 & 2, (30-45 cm) 7? 3 2 38 +0.59 0.1%

1 & 2, (60-75 cm) mu, 2 38 + 0.57 0.1%

Soil Moisture under Forest, BT Nature Reserve, S’ pore 93

These data suggest that while correlation with this empirical formula gives

slightly higher values for r than correlations with number of rain days, especially

for the upper layer of the soil, the introduction of the factor e is hardly necessary.

The weighting of R would also seem to be barely worth the labour of computa-

tion involved. The conclusion is therefore, that with the exception of the upper

layer of the soil where the introduction of e is desirable, the number of rain days

in the period (here a week) before the measurement of soil moisture provides a

simple parameter from amongst the meteorological factors affecting soil moisture.

Since the method of determining soil moisture is by no means highly accurate,

the coupling with it of such a ‘rough-and-ready’ measure as raindays seems

justifiable, whereas the use of more sophisticated meteorological parameters is

scarcely worth the labour involved.

Acknowledgments

I am grateful to the Director of the Botanic Gardens (Chairman, Nature

Reserves Board), Singapore for permission to use the study area and for

providing rainfall data. Evaporation data were kindly supplied by the Director,

Singapore Meteorological Service.

This paper is one of a projected series of studies of forest ecology in part

of the Bukit Timah Nature Reserve. It is my pleasure to acknowledge fruitful

discussions and the assistance of Mr. D. H. Murphy, Department of Zoology,

University of Singapore who also commented upon a draft of this paper.

Literature

American Society for Testing and Materials, 1963, Standard Method for Grain-size

Analysis of Soils, D422-63, Washington D.C.

BELGRAVE, W. N. C., 1930, ‘The effect of cover crops on soil moisture’, Malay.

agric. J., 18, 492-496.

LeaMy, M. L. and Panton, W. P., 1966, Soil Survey Manual for Malayan Condi-

tions, Ministry of Agriculture and Cooperatives, Bull. 119, Kuala Lumpur.

Watson, G. A., Wong Phui Weng and Narayanan, R., 1964, ‘Effects of cover

plants on soil nutrient status and on growth of Hevea V. Loss of nitrate —

nitrogen and of cations under bare soil conditions. A progress report on

results from a small-scale trial’, J. Rubb. Res. Inst. Malaya, 18, 161-174.

le Ux WO: abi rie > inthcitipeh- Abid .

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‘m4

Algae from Gunong Jerai (Kedah Peak), Malaysia

M. RATNASABAPATHY

School of Biological Sciences, University of Malaya, Kuala Lumpur

Abstract

The author’s collection of algae on Gunong Jerai is listed here with brief notes on

habitat and existing records in The Malay Peninsula. Structural features of several species

are also described. The total collection consists of 32 numbers, the 102 species and varieties

are listed under 57 genera.

Introduction

Gunong Jerai or Kedah Peak is a mountain largely composed of sandstone,

quartzite, quartz and some haematite (Robinson and Boden Kloss, 1916), and

acidic intrusive igneous rocks (Alexander 1962). It rises to nearly 1219 metres

(4000 feet) above mean sea level and is a prominent landmark of Kedah state. The

vast flat lowlands more or less surrounding the foothills are mainly planted with

rice. The sea to the west is only about 8 km (5 miles) away and in the mornings

or when the moisture- and possibly salt-laden westerly winds blow, one could

notice a thick mist moving across the peak towards the east. Ridley (1916),

amongst other workers, has suggested that Gunong Jerai was once an island

separated from the Malay Peninsula as in the case of Mt. Ophir, Johore and

showed that there is much resemblance between the floras of these two mountains.

The physical isolation of Gunong Jerai from the main range of the Peninsula and

the absence of neighbouring mountains of any comparable altitude is phyto-

geographically significant, and the lack of a true Thai element in the flora (see

Ridley, 1916) is noteworthy. Though the angiosperms, gymnosperms and especially

ferns have been abundantly collected and well documented, no data on algae

have been published.

Material and Method

This paper contains the results based on the study of 32 numbers of collections.

The algae were collected by the writer at altitudes of about 823-1189 metres

(2700-3900 feet) above mean sea level between May 12 and 20, 1969 (see figure 1).

The two main sites of collection were (1) a stream near the Government bungalow

and (2) the only concrete reservoir situated on the left side of Peak Road one-third

the distance from bungalow to peak. On a clear day, temperature readings of the

stream between 10.30 a.m. and 12.30 p.m. were found to vary between 21.8 and

22.2°C and pH from 8.0 to 8.3: at the reservoir the pH was 6.2 at 22.5°C. The

latter were determined in the field with a Lovibond comparator.

Specimens were first examined fresh, then preserved dry or in solutions of

formol acetic alcohol (Smith, 1950, p.34) or 5% formaldehyde. Of the nineteen

species of diatoms collected eighteen were obtained as empty frustules from muds,

bottom sediments and detrital material. The author consulted Johansen’s ‘‘Botanical

microtechnique” for staining methods and simple microchemical tests.

96 Gardens’ Bulletin, Singapore — X XVI (1972)

Species determination is incomplete due to inadequate reference material

and absence of reproductive stages in some samples. For the nomenclature and

sequence of major taxa (Divisions and Families), the system of Silva (1962) is

largely adopted and to a lesser extent those of Papenfuss (1955) and Smith (1950).

Genera are arranged according to systems of the following authors: Desikachary

(1959, Cyanophyta), Prowse (1959, Bacillariophyta), Huber-Pestolozzi (1955,

Euglenophyta), Randhawa (1959, Zygnemataceae), Scott and Prescott (1961,

Desmidiaceae).

Numbers in the list refer to the author’s collection. A complete set (101-133)

is deposited at the Botany Division, School of Biological Sciences, University of

Malaya, Kuala Lumpur; and 102, 104, 106, 107, 109-130, 132 and 133 are also

at the Tropical Fish Culture Research Institute, Malacca.

List of Species of Algae

CYANOPHYTA

CHROOCOCCACEAE

Chroococcus turgidus Nag. Nos 100 & 111.

Collected from washings of Sphagnum and sedge growing in reservoir and of

moss scraped from submerged concrete sides of reservoir.

Chroococcus ? minutus (Kiitz.) Nag. No 109.

Collected from bottom muds, reservoir.

Gloeocapsa sanguinea (Ag.) Kitz. Nos 112, 113, 115.

Collected from surface of moist rock.

Gloeotheca ? samoensis Wille No 109.

Colonies of 2 to 8 ovoid cells with rounded ends, daughter colonies with

individual sheaths, embedded in clear mucilage of parent envelope.

From bottom muds of reservoir.

Synechococcus ? aeruginosus Kitz. No 114.

Planktonic in reservoir and stream.

Dactylococcopsis acicularis Lemm. Nos 109, 110.

Collected from submerged moss-covered sides of reservoir.

OSCILLATORIACEAE

Phormidium inundatum Kitz. ex Gomont Nos 109, 116, 117.

Bright blue-green sheets; hormogonia abundant.

From surface of concrete roadside drain with water trickling over it.

NOSTOCACEAE

Anabaena sp. Nos 116, 118.

Filaments short, some up to about 500 » long; heterocysts rectangular, about

14 times longer than vegetative cells, intercalary, with two polar nodules; akinetes

absent.

Planktonic in pool, roadside drain.

Algae from Gunong Jerai 97

SCY TONEMATACEAE

Scytonema sp. Nos 115, 124.

In mass appears as black somewhat shiny patches on wet or dripping rock;

abundant.

Tolypothrix sp. No 128.

On submerged rock or mixed in with other algae and vegetation in stream.

STIGONEMATACEAE

Hapalosiphon hibernicus W. et G.S. West Nos 124, 125.

Main axes usually uniseriate, occasionally biseriate, cells shorter or a little

longer than broad, sheaths yellowish; branches uniseriate, mostly unilateral, sheaths

non-lamellated, colourless, cells 2-6 times longer than broad, apex of terminal

cells broadly rounded.

Collected from torrential stream amongst other algae.

Stigonema ocellatum (Dillw.) Thuret No 131.

Scraped from surface of rock in stream.

Reported from Templer Park (Hirano, 1967).

Stigonema panniforme (Ag.) Bornet et Flahault No 129.

Collected from torrential stream.

RHODOPHYTA

BATRACHOSPERMACEAE

Batrachospermum ? moniliforme Ralfs Nos 101-107, 108, 124. — Plates 1,

2A-B

Thalli macroscopic, 2-10 (-20) cm, moniliform, dark or pale blue-green or

purplish brown, mucilaginous, corticated, freely branched. Nodes prominent.

Internodes clear; corticating cells 2-6 times longer than broad, with short or long

uniseriate filaments at the base; internodal filaments bear globose to obovate

antheridial cells. Nodal filaments longer, branched; terminal cell ends in a long

hair with dilated base, when fertile may bear clusters of antheridial cells at distal

end. Cells of nodal branch citriform to oval, distinctly moniliform; apical cell

dome shaped, obscured by upgrowing branches; branches may develop from the

fourth axial cell and on the sixth, branches save as many as eight cells; young

carpogonia clavate, stalked, develop at end of short but many-celled nodal branch;

cystocarp rounded, more deeply pigmented, 1-3, more or less at periphery of

nodal branch.

The plants collected showed considerable variation in such features as colour,

size, length of internodes, degree of branching, structure of cortical and other

cells, and, it is possible that two forms or may be even species are involved here.

“Chantransia’’-like thalli were not uncommon; these were bluish green, uniseriate,

branched, the cells many times longer than broad, without terminal hairs, and

vegetative. Some of these thalli resemble Audouinella.

Found attached to rocks or gravel in stream; few specimens found on wet

soil of stream bank where seepage water overflowed, very few small plants in

the reservoir. Distribution patchy, sometimes gregarious.

98 Gardens Bulletin, Singapore — XXVI (1972)

PYRRHOPHYTA

— PHYTODINIACEAE

Dinococcus bicornis (Woloszynska) Fott Nos 123-125 — Plates 2C,

3A-B

Cells solitary, dorsal margin in front view convex, sometimes slightly depressed

in the middle, showing at each end a short pointed spine; ventral view somewhat

angular; vertical view ellipsoid; end view ovate. Stalk short, stout, colourless.

Chromatophores obscured by dense accumulation of food reserve including a more

or less central area of oil.

Epiphytic on Oedogonium, other filamentous algae, and colonial diatoms

e.g. Eunotia in stream; frequently gregarious.

BACILLARIOPHYTA

EUNOTIACEAE

Eunotia ? camelus Ehrenb. Nos 122, 124.

Collected from stream muds.

Eunotia diodon Ehrenb. var. minor Grun. Nos 122, 124.

In stream muds and epiphytic on other algae.

Recorded in Cameron Highlands and Malacca River (Prowse, 1962a).

Eunotia lunaris (Ehrenb.) Grun. No 123.

In stream muds and epiphytic on other algae.

Recorded in Cameron Highlands (Prowse, 1962a).

Eunotia lunaris (Ehrenb.) Grun. var. capitata Grun. No 122.

Collected from stream muds.

Recorded in Malacca River (Prowse, 1962a).

Eunotia monodon Ehrenb. var. tropica Hustedt Nos 122, 123.

Among bryophytes and other vegetation in stream.

Three varieties were recorded from stream, river and fish pond elsewhere in

West Malaysia (Prowse, 1962a).

Eunotia pectinalis (Kiitz.) Rabenh. var ventralis (Ehrenb.) Hustedt No 120.

Collected from stream muds.

The species was recorded at Cameron Highlands (Prowse, 1962a). Var. minus

(Kiitz.) Rabenhorst has been recorded from Perak (Patrick, 1936).

Eunotia ? pseudoflexuosa Hustedt No 123.

Collected from stream muds.

Eunotia robusta Ralfs Nos 118, 120.

From roadside pool with flowing water and stream sediments.

Recorded in Malacca river (Prowse, 1962a).

Desmogonium rabenhorstianum Grun. Nos 120, 122, 123.

Epiphytic on other algae, bryophytes and in sediments of stream. Prowse

(1962a) reported this species from various habitats elsewhere in Malaysia, including

swamps and coastal streams, at Cameron Highlands in West Malaysia.

Algae from Gunong Jerai 99

ACHNANTHACEAE

Achnanthes sp. Nos 118, 123.

Collected from roadside pool with flowing water; epiphytic on liverwort in

reservoir.

NAVICULACEAE

Frustulia rhomboides (Ehrenb.) De Toni Nos 106, 115, 120-123, 126.

Distributed widely and abundantly in various habitats; its common occurrence

suggests ecological significance and further studies are warranted.

Recorded from most states of West Malaysia, including Cameron Highlands

and Fraser’s Hill (Prowse, 1962a). Very abundant in algal samples collected by

the writer in July 1970 at Tasek Bera, Pahang.

Navicula spp. Nos 104, 118, 120.

Many small species, especially common in bottom muds of streams and

roadside pools.

Pinnularia biceps Gregory Nos 112, 113.

Collected from bottom sediments and submerged concrete sides of reservoir.

Recorded from padi-fields, Beranang (Johnson, 1970). Var minor (Boye

Petersen) A. Cleve has been recorded in Malacca river and fish ponds (Prowse,

1962a).

Pinnularia legumen Ehrenb. No 111.

Collected from stream and reservoir sediments.

Recorded from Perak (Patrick, 1936); Taiping lakes (Prowse, 1962a) and from

Templer Park (Hirano, 1967).

Pinnularia maior (Kiitz.) Cleve Nos 122, 123.

Collected from stream muds and pools.

Reported from Perak (Patrick, 1936). Var. linearis was recorded in Templer

Park (Hirano, 1967).

CYMBELLACEAE

Cymbella ? ventricosa Kitz. Nos 112, 113.

Collected from bottom sediments of reservoir.

Recorded at Cameron Highlands (Prowse, 1962a).

NITZSCHIACEAE

Hantzschia amphioxys (Ehrenb.) Grun. ? var. capitata O. Muller No 126.

Collected from stream mud.

Recorded from stream, Klang Gates (Prowse, 1962a).

100 Gardens’ Bulletin, Singapore — XXVI (1972)

SURIRELLACEAE

Surirella linearis W. Smith Nos 123, 126.

Collected from stream sediments.

Reported from Perak (Patrick, 1936); from fish ponds at Penang and Malacca

river (Prowse, 1962a).

Surirella spiralis Kutz. Nos 101, 114, 120.

Collected from stream sediments. Rare.

Reported from various habitats in Malacca (Prowse, 1962a). The writer has

collected the species at various times during 1968 to 1970 in the Gombak river,

Selangor and in July, 1970 at Tasek Bera, Pahang.

Surirella ? tenuissima Hustedt Nos 123-125.

From sediments on bed-rock and liverwort in stream.

Recorded in Cameron Highlands (Prowse 1962a).

CHRYSOPHYTA

MALLOMONADACEAE

Several species of Mallomonas were encountered in the stream sediments,

reservoir and amongst other algae and bryophytes. Cells were mostly empty and

spines usually lacking. The following species appear to be present:

Mallomonas ? elliptica (Kisselew) Conrad Nos 110, 111.

Mallomonas producta (Zacharias) Iwanoff Nos 112, 113.

Mallomonas ? teilingioides Prowse Nos 112, 113.

DINOBRYACEAE

Dinobryon sertularia Ehrenb. No 111.

Mostly only empty envelopes were found; solitary or in colonies of up to

six cells.

Colonies few-celled epiphytic on filamentous algae, rare in stream; mostly

empty envelopes, common in sediments and waters of reservoir.

Recorded from many acid swamps and fish ponds in many states of West

Malaysia (Prowse, 1962b).

RHIZOCHRYSIDACEAE

Lagynion scherffeltii Pascher Nos. 108, 109.

Epiphytic on filamentous algae in stream, sometimes gregarious, fairly common.

Lagynion ? macrotrachelum (Stokes) Pascher No 124.

Epiphytic on filamentous algae in stream.

XANTHOPHYTA

STIPITOCOCCACEAE

Stipitococcus crassistipitatus Prescott Nos 122, 123.

Epiphytic on filamentous algae in stream.

Algae from Gunong Jerai 101

CHARACIOPSIDACEAE

Characiopsis sp. No 128.

Epiphytic on liverwort in stream.

TRIBONEMATACEAE

Tribonema sp. No 124.

Cells 14-3 times as long as broad, slightly swollen in middle and narrowing

at cross walls, ‘H’-pieces distinct but rare, chloroplasts many. This species is

related to T. viride Pascher.

Tychoplanktonic and in bottom muds of stream; often entangled with other

filamentous algae.

Neonema sp. No 124.

Similar in habitat to that of Tribonema; rare.

EUGLENOPHYTA

EUGLENACEAE

Euglena 2? mutabilis Schmitz No 120.

Collected from stream muds.

Recorded in Kedah padi fields (Sands, 1934), in carp ponds and stagnant

water elsewhere in Malaysia (Prowse, 1957b, 1958) and in Templer Park (Hirano,

1967). The writer has collected it several times between 1968 and 1970 in the

Gombak river, Selangor.

Var. minuta Fritsch has been reported from many stagnant waters in West

Malaysia (Prowse, 1958).

Trachelomonas volvocinopsis Swirenko No 120.

Collected from stream muds.

Recorded from standing water and fish-ponds in various states of West

Malaysia (Prowse, 1962b).

Lepocinclis ? marsonni Lemm. emend Conrad No 120.

Collected from stream muds. Common.

Recorded from Taiping Lakes (Prowse and Ratnasabapathy, 1970).

CHARACIACEAE

Characium sp. Nos 128, 129.

Cells solitary, fusiform, straight or slightly arcuate, poles acute and produced

as long fine spines, each spine about half or more cell length; one spine terminating

in a small rounded colourless attachment disc; chloroplast single, parietal, almost

filling cell.

The specimens were growing attached to the leaves of aquatic liverworts in

a torrential stream. This species of Characium closely resembles Schroederia

setigera, which however is reported to be always planktonic (see Smith, 1950).

The presence of an eyespot and the disc-like ending of the spine in some species

of Schroederia was reported. These species may well be attached forms that

have become subsequently detached by accident and incorporated into the

plankton. Further work on the ecology and taxonomic status of Schroederia with

its described species is needed.

102 Gardens’ Bulletin, Singapore — X XVI (1972)

ULOTRICHACEAE

Stichococcus ? scopulinus Hazen Nos 128, 131.

Filaments uniseriate, short, 1 to 4 or rarely 5 cells; cells cylindrical, sometimes

up to 9 times longer than broad but usually shorter, ends rounded; chloroplast

single, parietal, laminate, encircling about half the cell and usually not extending

to the ends, apple-green coloured; pyrenoids lacking.

Epiphytic on leaves of aquatic bryophyte in torrential stream; tends to be

gregarious.

Ulothrix sp. Nos 120, 132.

Collected from a small pool of Government bungalow; in slow-flowing stretches

of stream near bungalow. Rare.

TRENTEPOHLIACEAE

Trentepohlia iolithus (L.) Wallr. Nos 114, 133.

Orange coloured, forming felted patches; filaments much-branched, unilateral

at times, cells usually 2-5 times longer than broad, length up to about 90 42,

chloroplast indistinct, reticulate; haematochrome abundant; cell wall fairly thick,

lamellated in older cells; sporangia borne terminally or laterally on short or

many-celled branch, usually single; spherical, diameter up to 25 yp, stalk cell

inflated at base and bent distally.

Epilithic, aerial.

OEDOGONIACEAE

Bulbochaete sp. Nos 111, 120.

Thallus mostly uniseriately branched, cells 3-7 times as long as broad, length

of cells 25-45 »; cell apex slightly inflated, with laterally placed seta; setae

swollen at the base and very long, about 250 ». No reproductive stages found.

Attached to other vegetation and twigs in stream and reservoir.

Oedogonium sp. Nos 112, 113, 120.

Only vegetative filaments were encountered, and probably more than one species

are involved.

Attached to aquatic vegetation, twigs, etc. in reservoir and in streams.

ZYGNEMATACEAE

Mougeotia ? elegantula Wittr. No 120.

Collected from quieter stretches and pools of streams. Common.

Several specimens resembling Mougeotia and Debarya (No. 115-119) could

not be assigned accurately as only the vegetative stages were encountered.

Zygnema sterile Transeau No 118.

Cells up to two times longer than broad; cell wall thick, lamellated; chloroplast

two, more or less stellate, each with one pyrenoid.

Collected from stream, roadside pools and drains, often mixed in with other

filamentous algae. Common.

Several unidentified species of Zygnema or Zygnemopsis (Nos 115, 117, 120)

were collected from the stream; some of these had cells many times as long as

broad; reproductive stages were absent.

Algae from Gunong Jerai 103

Spirogyra spp. Nos 120, 127.

At last 3 species of Spirogyra were collected from the streams and one

species from the reservoir; they had one to four chloroplasts. One of these (No. 127)

was found in abundance in almost a pure stand in small cracks of stream bed-rock

with static or gently flowing water, showing recently completed conjugation and

young zygospores. Specific determination was not possible because the sculpture

was not clear.

Sirogonium sp. No 120.

A species with four chloroplasts, each making half to three quarters of a

turn; all filaments were in vegetative condition.

Collected from quieter edges of stream.

MESOTAENIACEAE

Mesotaenium caldariorum (Lagerh.) Hansgirg No 133.

Cells cylindrical, ends rounded, sides slightly to more or less parallel, length

up to two times longer than broad; chloroplast single, axile, laminate, with or

without a pyrenoid; mucilage surrounding cells clear, enveloping several cells.

Collected from moist rock of roadside associated with Trentepohlia.

Cylindrocystis brebissonii Menegh. No 115.

Amongst Stigonema, Scytonema and Cosmarium on moist rock. The writer

has collected the species in April 1970 in Tasek Bera, Pahang.

Netrium digitus (Ehrenb.) Itzigs. et Rothe Nos 120, 122, 124 — Plate 3C

Collected from slow-flowing stretches of stream. Common.

Recorded in Taiping Lakes (Prowse and Ratnasabapathy, 1970).

DESMIDIACEAE

Actinotaenium cucurbita (Bréb.) Teil. No 118, 119.

Collected from pool with flowing water, dripping wall of small iron storage

tank, roadside of hill.

Actinotaenium cucurbitinum (Biss.) Teil. No 119.

Collected from dripping wall of small iron storage tank, roadside of hill.

Actinotaenium elongatum (Racib.) Teil. No 119.

Collected from dripping wall of small iron storage tank, roadside of hill.

Actinotaenium grande (Delp.) Teil. No 112.

Collected from small impoundment to reservoir.

Actinotaenium subglobosum (Nordst.) Teil. No 119.

Collected from dripping wall of small iron storage tank, roadside of hill.

Actinotaenium truncatum (Bréb.) Teil. Nos 112, 118.

Collected from pool with flowing water, roadside of hill and small impound-

ment to reservoir.

Actinotaenium wollei (Gronbl.) Teil. No 119.

Collected from dripping side of small iron storage tank, roadside of hill.

Closterium cynthia de Not Nos 112, 113, 122.

Collected from bottom muds, reservoir.

Recorded in pond, Ampang (Biswas, 1929).

104 Gardens’ Bulletin, Singapore — XXVI (1972)

Closterium libellula Focke var. intermedium (Roy et Biss.) G. S. West Nos

111, 112 — Plate 3D

Collected from bottom muds of reservoir.

The species was recorded from the Taiping Lakes (Prowse and Ratnasaba-

pathy, 1970).

Closterium libellula Focke var. interruptum (West et West) Donat Nos 120,

126.

Collected from stream mud.

Closterium parvulum Nag. No 120.

Collected from stream muds.

Reported from Johore (Bernard, 1909); Setapak Lake (Biswas, 1929); padi-

fields, Beranang (Johnson, 1970); elsewhere in Malaysia (Prowse, 1957a).

Closterium navicula (Bréb.) Liitkem. No 115.

Scraped from moist rock, grows mixed in with Cosmarium spp., Cylindro-

cystis and Cyanophyceae. Recorded from Taiping Lakes (Prowse and Ratnasa-

bapathy, 1970).

Closterium striolatum Ehbrenb. Nos 109-111.

Collected from muds and vegetation in reservoir.

Recorded in Templer Park (Hirano, 1967).

Closterium tumidum Johnson Nos 104, 123.

Collected from muds overlying bottom rock of stream.

Recorded in pond, Setapak (Biswas, 1929).

Species of Closterium were found in practically all the samples of streams,

rock pools, roadside drains and the reservoir. As an adventitious element in the

plankton of both flowing and still waters they were not conspicuous. Multiplication

by transverse division of cells was evident and appears to be the general method

of maintaining the populations. Zygospore formation was lacking. The wide-

spread occurrence, the ability to colonize a variety of ecological niches and fair

abundance suggest ecological and economic importance.

Pleurotaenium minutum (Ralfs) Delp. var. minus (Racib.) Krieg. Nos 104,

119, 125:

From rock scraping and mud of stream; dripping side of small iron storage

tank, roadside of hill.

Var. excavatum was recorded from Taiping Lakes (Prowse, and Ratnasa-

bapathy, 1970).

Pleurotaenium trabecula (Ehrenb.) Nag. No 120, 122-124.

Collected from stream muds, mixed in with other algae and occasionally in

plankton. Fairly common.

Tetmemorus laevis (Kiitz.) Ralfs Nos 120, 124, 126 — Plate 4C

Cells up to 130 » long and eight times the breadth, sinus open and con-

spicuous; semi cell slightly inflated just above base, apex deeply incised with the

lobes somewhat rounded and asymmetrical; cell wall finely punctate, four punctae

radially arranged at base of apical incision; chloroplast ridged with about six

pyrenoids arranged longitudinally.

Collected from stream muds. Fairly common.

Recorded elsewhere in Malaysia (Prowse, 1957a).

Algae from Gunong Jerai 105

Euastrum binale (Turp.) Ehrenb. var. brevius (Bernard) Hirano No 125.

A small species fairly frequently found in waters of streams.

Reported from Templer Park (Hirano, 1967).

Euastrum dubium Wolle No 113.

Collected from small impoundment to reservoir.

Euastrum sinuosum Lenorm. var. dideltoides Krieg. Nos 109-111 — Plate 4A

A larger species, up to 75 p long.

| Amongst vegetation and bottom muds, reservoir.

The species was recorded from Taiping Lakes (Prowse and Ratnasabapathy,

1970).

Euastrum gnathophorum W. et G.S. West Nos 106, 107, 120, 124.

Collected from stream muds.

Recorded from Taiping Lakes (Prowse and Ratnasabapathy, 1970).

Euastrum turgidum Wolle No 119.

Collected from dripping side of small iron storage tank, roadside of hill.

Cosmarium contractum Kirchn. Nos 120, 124.

Collected from stream.

Var. Jacobsenii was recorded in a pond, Ampang (Biswas, 1929).

Cosmarium decoratum W. et G.S. West No 18.

Collected from stream muds.

Cosmarium hians Borge forma No 113.

Collected from small impoundment to reservoir.

Cosmarium lunatum Turn. Nos 111, 118, 120.

Collected from pool with flowing water, hill roadside; bottom muds, stream;

bottom muds, reservoir. Common.

Cosmarium quadrifarium Lund. No 118.

Collected from bottom muds, reservoir.

Cosmarium taxichondrum Lund. No 118.

Collected from pool, roadside drain with flowing waters. Rare.

Cosmarium sp. No 106 — Plate 4B

Several other species of Cosmarium specifically unidentifiable are not listed.

Staurastrum sp. No 114.

Planktonic in reservoir.

Spondylosium pulchellum Arch. No 127.

Short chains of cells epiphytic on aquatic vegetation in stream.

Hyalotheca dissiliens (J.E. Smith) Bréb. Nos 107, 120, 124, 127 — Plate 4D

Collected from stream, common in samples of other filamentous algae.

Recorded in pond, Setapak (Biswas, 1929); from elsewhere in Malaysia

(Prowse, 1957a).

106 Gardens’ Bulletin, Singapore — X XVI (1972)

Analysis of Results

Among the algae collected, 57 genera and 102 species including varieties

were identified. About 25 species were attached forms, and 72 species occurred

in the bottom muds or as tychoplankton. The true planktonic element appeared

to be lacking and the few encountered such as Frustulia rhomboides and Hyalo-

theca dissiliens were largely adventitious, stirred up from the bottom and

suspended by the flowing waters. This was inferred from their abundant presence

in bottom samples, apart from their features of habit and morphology. There was

an abundant population and often a rich variety of species, especially desmids and

diatoms, in the shallower, slow-flowing stretches of streams with aquatic phanero-

phytes, bryophytes and decaying vegetation and where the canopy of trees overhead

was less dense. The reservoir, which is a comparatively small habitat, being only

about twenty metres square, had some 25 species, though few of these also

occurred in the stream.

The number of species in the Divisions of algae represented in these

collections were as follows: Cyanophyta 13, Rhodophyta 1, Pyrrophyta 1,

Bacillariophya 19, Chrysophyta 6, Xanthophyta 4, Euglenophyta 5, and Chlorophyta

51. The Cyanophyta, Bacillariophyta and Chlorophyta were also numerous in

number of individuals and may be considered as dominant. Among the Chlorophyta,

the Zygnematales formed about 80% of which the Desmidiaceae alone comprised

two-thirds. Several of the species of algae are known to occur outside Malaysia,

some have been recorded from elsewhere in Malaysia, and some are being recorded

for the first time here. The occurrence of the temperate species Dinococcus

bicornis Fott here extends its known range of geographical distribution.

The presence of Dinobryon sertularia and species of the related flagellate

Mallomonas in the reservoir may be undesirable with respect to the quality of a

water source. Population outbursts of Dinobryon and Mallomonas have been

reported to impart an obnoxious odour and a fishy taste to the waters of Hoover

Reservoir, Ohio and Lake Michigan thus necessitating the treatment of the water

supplies (Palmer, 1964).

There are many problems concerning the freshwater algae of Gunong Jerai

worthy of further investigation. Species of Closterium, Cosmarium, and Frustulia

rhomboides were common and widely distributed; Batrachospermum, though

rarer, showed an interesting sporadic occurrence with Audouinella-like thalli which

may turn out to be the juvenile stages of the species. Studies on such aspects as

their distribution, morphological variation, taxonomy, reproduction and life history

should be rewarding. A comparative study of the Gunong Jerai algal flora with

similar areas in other parts of Malaysia is not possible owing to the lack of

published records. It may be mentioned, however, that about seven of the

nineteen species of diatoms listed have been previously recorded at Cameron

Highlands (see Prowse, 1962a). There is a wide and exciting field for future

investigation of the Malaysian mountain algal floras.

Acknowledgements

I am indebted to Dr. Gerald Prowse, lately Director of The Tropical Fish

Culture Research Institute in Malacca for advice and for checking identifications.

I also wish to thank Dr. Ralph A. Lewin, Professor, Scripps Institution of Ocean-

ography, University of California, La Jolla for critical comments on the manu-

script and Dr. B. C. Stone, Reader at The School of Biological Sciences, University

of Malaya, for introducing me to the background literature on Gunong Jerai.

Plate 1. Batrachospermum ? moniliforme Ralfs

A: Apex of thallus stained with Phloxine B

B: Whorls of lateral branches of limited growth

C: Cystocarps at periphery of whor! of lateral branches

& ]

Sys

YY ee

Plate 2.

A: Batrachospermum ? moniliforme Ralfs, cystocarp terminal on short lateral branch,

stained with Phloxine B

B: Idem, spermatia on lateral branches of limited growth

C: Dinococcus bicornis Fott, epiphytic on colonial Desmogonium, front view

) S0u >

Dinococcus bicornis Fott, epiphytic on filamentous Xanthophyceae, front view

Idem, end view

Netrium digitus (Ehrenb.) Itzigs. et Rothe

- Closterium libellula Focke var. intermedium (Roy et Biss.) G. S. West

Euastrum sinuosum Lenorm. var. dideltoides Krieg.

Cosmarium sp.

Tetemorus laevis (Kiitz.) Ralfs

Hyalotheca dissiliens (Smith) Bréb. stained with methylene blue (oil immersion)

ad Iwuif SONONNS JO dV¥W L 38NSIS

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108

Alexander, J. B. 1962

Bernard, Ch. 1909

Biswas, K. 1929

Desikachary, T.V.

1959

Fott, "B. 1959

Fritsch, F. E. 1935

Gronblad, R., Prowse,

G. A. and Scott,

A.M 1959

Hirano, M. 1967

Huber-Pestalozzi, G.

1955

Johansen, D. A. 1940

Johnson, A. 1970

Migula, W. 1907

Palmer, C.M. 1964

Papenfuss, G.F. 1955

Gardens’ Bulletin, Singapore — XXVI (1972)

Bibliography

A brief summary of the geology of Malaya. Malayan Nat.

J. le ¢ 30-35.

Algues unicellulaires d’eau douce récoltées dans le dom-

aine Malais. Dept. de TAgric. aux Indes Néerl.,

Buitenzorg. 44 pp., 6 plates.

Papers on Malayan aquatic biology IV. Freshwater algae

(with addendum). J. Fed. Malay States Mus. 14 (3-4) :

404-435, 479-481, plates 9-13.

Cyanophyta. I.C.A.R. Monogr., New Delhi.

Algenkunde. Veb Gustav Fischer Verlag, Jena.

The structure and reproduction of the algae.

Vol. 1. Camb. Univ. Press.

Sudanese desmids. Acta Bot. Fenn. 58 : 1-82.

Freshwater algae collected by the joint Thai-Japanese

biological expedition to Southeast Asia 1961-2. Nature

and Life in Southeast Asia. Vol. V, pp. 1-71, 16 plates.

Das Phytoplankton des Susswassers. Teil 4. Euglenophy-

ceen; E. Schweizerbart’sche Verlagsbuchhandlung,

Stuttgart.

Botanical microtechnique. McGraw Hill Book Co., Inc.,

N. Y.

Blue-green algae in Malaysian rice-fields. J. Sing. natn.

Acad. Sci. 1 (3) : 30-36

Kryptogamen-Flora von Deutschland, Deutsch-Osterreich

und der Schweiz 6, Kryptogamen 2, Algen 1 : Cyanophy-

ceae, Diatomaceae, Chlorophyceae. Text pp. 1-918, plates

1-43.

Algae in water supplies of the United States. IN Algae

and Man, ed. by Jackson, D. F. Plenum Press, N. Y., pp.

239-261.

Classification of the algae. IN A century of progress in the

natural sciences. Calif. Acad. Sci., San Francisco.

Algae from Gunong Jerai 109

Pascher, A. 1925

————. 1937-39

Patrick, R. 1936

Patrick, R. and

Reimer, C. W. 1966

Prescott, G. W. 1962

Prowse, G. A. 1957a

——. 1957b

———. 1958

———. 1960

. 1962a

. 1962b

Prowse, G. A. and

Ratnasabapathy,

M. 1970

Randhawa, M. S.

1959

Ridley, H. N. 1916

Robinson, H. C. and

Boden Kloss, C.

1916

Sands, W. N. 1934

Scott, A. M.,

Gronblad, R. and

Croasdale, H. 1965

Die Susswasserflora Deutschlands, Osterreichs und der

Schweiz. Vol. 11, pp. 1-118.

Heterokonten. In L. Rabenhorst, Kryptogamen — Flora

Deutschland, Osterreich und der Schweiz. Vol. 11, 1092

Pp.

A taxonomic and distributional study of some diatoms

from Siam and the Federated Malay States. Proc. Acad.

Nat. Sci. Philad. 88 : 367-470.

The diatoms of the United States. A. N. S. P. Monogr. 13,

Vol. I. Philad.

Algae of the Great Lakes area. Cranbrook Inst. Sci. Bull.

31 : xiii + 1-977.

An introduction to the desmids of Malaya. Malayan Nat.

J. 11 (3): 42-58.

Fish and food chains. Malayan Nat. J. 12 (2) : 66-71.

The Eugleninae of Malaya. Gard. Bull. Sing. 16

136-204.

Vegetation changes in experimental ponds. In Proc.

Centenary and Bicentenary Congr. Biol. Sing. pp. 288-291.

Diatoms of Malayan freshwaters. Gard. Bull. Sing. 19

(1) : 1-104.

Further Malayan freshwater Flagellata. Gard. Bull. Sing.

19 (1) : 105-145.

A species list of freshwater algae from the Taiping Lakes,

Perak. Gard. Bull. Sing. 25 (2) : 179-187.

Zygnemaceae. 1. C. A. R. Monogr., New Delhi.

The natural history of Kedah Peak V. Botany. J. Fed.

Malay States Mus. 7 (2) : 39-58.

The natural history of Kedah Peak I.

Introduction. J. Fed. Malay States Mus. 6 (4) : 253-256.

The coloured scums of padi fields. Malay. Agri. J. 22

(10) : 484.

Desmids from the Amazon Basin, Brazil. Acta Bot. Fenn.

69 : 3-94.

110 Gardens’ Bulletin, Singapore — XXVI (1972)

Scott, A. M. and Indonesian desmids. Hydrobiologia 17 (1-2) : 1-132, with

Prescott, G. W. 63 plates.

1961

Silva, P. C. 1962 Classification of algae. In Physiology and biochemistry of

algae. Edited by Ralph A. Lewin. Academic Press, N. Y.

Skuja, H. 1949 Zur Siisswasseralgen — Flora Burmas. Nova Acta R. Soc.

Scient. upsal. Series IV, 14 (5) : 1-188, 37 plates. Uppsala.

Smith, G. M. 1950 The freshwater algae of the United States. 2nd ed., N. Y.

Teiling, E. 1954 Actinotaenium genus Desmidiacearum resuscitatum. Bot.

Notiser. 4 : 376-426.

Tiffany, L. H. and The algae of Illinois. Chicago.

Britton, M. E. 1952

Wildeman, E. 1900 Flore de Buitenzorg III. Algues. Leiden.

Review

E. J. H. CORNER AND K. WATANABE, //lustrated Guide to Tropical

Plants.

Hirokawa, Tokyo, 1969. 1147 pp., 29 colour pl., c.2000 line drawings.

The title of this large and ambitious book is deceptive: It is based on a

collection of line drawings of phanerogams, made by Professor Watanabe during

a 3-year stay in Singapore and Malaya; the species selected are therefore largely

unrepresentative of the tropics as a whole, both on taxonomic, ecological and

utilitarian grounds. Besides most of the commoner cultivated plants of Malaya,

many of which are exotics and indeed widely grown in the tropics, they include

a somewhat arbitrary selection of native species, weeds, and many plants existing

in that region only as individuals in botanic gardens, of which some are by no

means common in their countries of origin. The drawings are arranged by

families according to Engler’s system; each is captioned by its family name.

Beneath the illustration is given the scientific, Malayan (if known), Japanese,

and sometimes Indian and English names; a short but useful descriptive note

indicating distribution, habit, leaf arrangement and flower and fruit colour by

Professor Corner; and a note on uses in Japanese. The usefulness of each plant

is then summarised in a curious annotated point-grid system, which appears to

have been intended as a ready-reference abbreviation but is in fact the very

reverse, being difficult to follow and space-demanding; finally, the position in

Hutchinson’s classification is indicated. Some 700 of the more than 2,000 line

drawings were originally published in two volumes by the Japanese administration

of the Singapore Botanic Gardens during the second world war, entitled

“Illustrations of Useful Plants of the Southern Regions: Section 1: Medicinal,”

and “Section II: Edible Plants.” These books are now rare, but many of the

same plates were used again by Professor H. B. Gilliland for his book “Common

Malaysian Plants” (University of Malaya Press, 1958).

This book cannot be used for identifying plants, as the illustrations and

descriptions make no attempt to be diagnostic, and no keys are given. Its purpose

would appear to be a reference book for laymen in S.E. Asia, especially foreign

agriculturists, foresters and businessmen who, on being shown a named plant

to which certain useful properties have been ascribed, wish to check the veracity

of this information. The book is'a monument to Professor Watanabe’s energy,

but it is a pity that the illustrations, the originals of which must generally have

been of high quality, have had to be reduced to a scale where the details, especially

of floral morphology, have often been lost.

P. S. ASHTON

Institute of South-East Asia Botany

University of Aberdeen

111

REVIEW

TSENG-CHIENG HUANG. Pollen Flora of Taiwan

National Taiwan University, Botany Dept. Press, Taipeh. 1972;

pp. v+ 276, 177 photographic plates and index. Price U.S. $27.00 (postage and

handling included).

This is one of the very few local pollen floras that have appeared in the past

decades. The author was probably inspired by M. Ikuse’s “‘Pollen Grains of Japan”

published in 1956.

Part one of this book is an introduction, including an illustrated account on

pollen morphology. Part two constitutes the main body of the book — the systema-

tic treatment. It contains keys to the families (divided into 25 classes) and

descriptions of 197 families of seed plants (Gymnosperms-9, Dicotyledons-157,

and Monocotyledons-31). The families are arranged alphabetically within these

three taxa. Each family consists of a general description of the pollens and a

bracketed key to the genera (if more than 2). The descriptions of genera and

species are in general very concise. Numbers of voucher herbarium specimens are

also cited.

Two most valuable features of this book are probably the comprehensive and

useful keys based on pollen characters for identifying families and genera, and the

excellent quality of photographic plates. The latter is attributable to the facilities

of fine equipment in the Palynological Laboratory at the National Taiwan University

and the skilled technique in preparing slides and photographs.

This book will be very useful to plant taxonomists and palaeobotanists and I

have no hesitation in strongly recommending it.

H. KENG

Department of Botany

University of Singapore

112

Athrodactylis and Pandanus: A brief comment on the

‘*Characteres Generum Plantarum’ of J. R. and G. Forster

BENJAMIN C. STONE

School of Biological Sciences, University of Malaya, Kuala Lumpur

Although it is well known that, in the work “Characteres Generum Plantarum”

of 1776 by Johan Reinhold and Georg Forster, these authors boldly and improperly

utilized information they had acquired through their relationship with Sir Joseph

Banks in working on the materials which Banks and Solander had amassed during

the first of Captain Cook’s voyages. in somewhat the same way that had been

earlier done by the brother (Parkinson) of the expedition artist, it is interesting

to discover that they did not confine their borrowings to any one source.

While working briefly in the Herbarium of St. Xavier’s College in Bombay

I was able to use originals of the Forsters’ work, as well as a number of other

ancient and precious volumes. One of these was the ‘‘Horti Indici Malabarici” of

_ Henricum van Reede tot Draakestein (published between 1678 and 1703). Studying

in particular the references to Pandanus I then turned to the Forsters’ work, and

was immediately struck by the resemblance virtually to the point of identity between

the drawing of a fruit (phalange) in fig. 6 of Reede’s ‘‘pars secunda” with the

name “‘Kaida Taddi” and the illustration (fig. 75) of the Forsters’ ““Athrodactylis

spinosa” (i.e. Pandanus). The Forsters’ illustration (titled “‘Appendix Palmae’’)

shows (a) one stamen, (4) one phalange, and (c) one seed. The last two are almost

_ exact copies from Reede’s illustration of ““Kaida Taddi”’.

This copying, for such I am sure it was, tallies logically with the Forsters’

_ descriptive text, where (p. 150) they give the diagnosis of ‘‘spinosa. 1. ATHRO-

_ DACTYLIS.” in the Linnaean manner, with the synonyms “Bromelia Sylvestris.

Lin. Flor. Zeyl. Burm. Ind. 79.; Keura Forskal Arab.; Kaida Rheed. Hort. Malab.

_p. IL. t. 1-8. Pandanus Rumph.” This is somewhat confusing, but it seems to

_ suggest that the Forsters considered all these names synonymous with their species

_ “Athrodactylis spinosa,” although Reede’s figures show clearly (to the modern

_ reader) about four distinct species of Pandanus. The Forsters’ list of synonyms

_ beneath the ‘‘species” rather than beneath the “‘genus” diagnosis (although, strictly

_ speaking, there is no separate botanical diagnosis of the species; only an explanation

of the generic name, followed by the ‘“‘Explicatio” for Tab. LXXV.) seems to a

modern reader a mixture of generic and specific synonyms. It is difficult to come

to a conclusion as to whether the Forsters thought that all the synonyms they

cite really referred to a single species or to a single genus. The only post-Linnaean

- name cited is Keura Forskal. This was monotypic, and thus by citing it in synonymy

_ we would expect the epithet of the species to be taken up if we apply the current

- nomenclatural Code. The full name of Forskal’s plant was Keura odorifera. This

_ name is based on material which Forskal purchased in a small village in south-

western Arabia, where the staminate inflorescences were on sale as a perfume.

113

114 Gardens’ Bulletin, Singapore — X XVI (1972)

To this day in both India and in adjacent regions the practice persists, and in 1968

in the villages on Madh Islet in Bombay (for example) one can still see these

inflorescences for sale; children and women carry and wear them, “harvesting”

them from the trees. The plants are as common in cultivation as wild, especially

along the coast. Forskal’s genus Keura is certainly synonymous with Pandanus

Stickm. of 1754; the publication of Forskal’s ‘Flora Aegypto-Arabica” was in

1775. The generic name Athrodactylis J. R. & G. Forst. is thus clearly a synonym

since Forskal’s publication was cited. Unfortunately the type material of Keura

no longer exists. It is, however, very likely to be the plant which is still used in the

way mentioned — as perfume — along the coasts of India and is traded for such in

adjacent areas. This is clearly identical with Pandanus odoratissimus L. f.

(the type of which is from Ceylon). At any rate, it is clear that: (1) Athrodactylis,

as a genus, is equivalent to Pandanus; (2) Athrodactylis as a name is not nomencla-

turally usable, being published with a valid name in synonymy (i.e. Keura);

‘3) the epithet “‘spinosa”’ is also illegitimate since the epithet “‘odorifera’’ of Forskal

should have been taken up; (4) except for the reference to Pandanus Rumph. the

remaining synonyms all refer to Indian or Ceylon plants; thus it is plain that

A. spinosa is none other than Pandanus odoratissimus Lf. This is borne out

convincingly by the fact that to illustrate A. spinosa, the Forsters copied Reede

tot Draakestein’s illustration of “‘Kaida Taddi” which is unquestionably P. odoratis-

simus Lf.

As a final point it is interesting to note that G. Forster in his later work on

Pacific Island Plants (Prodr. 69. 1786) used the binomial Pandanus odoratissima

(sic). This certainly means that he then considered Pandanus as the correct generic

name, and by taking up Linnaeus filius’ specific epithet also (though altering it to

feminine gender) he apparently also meant that the Tahitian plants which he was

discussing were to be equated with the Indian and Ceylon species. This interpreta-

tion can perhaps therefore be viewed as a confirmation of the conclusions given

above.

It is interesting to note further that although much of the ‘Characteres Generum

Plantarum’ was based on Pacific Islands plant collections, the “‘Appendix’’ with

Athrodactylis does not specifically mention any Pacific materials. The reference to

Rumphius suggests Forsters’ belief that the species of India and Ceylon extended

into the East Indies. But there is no mention of Pandanus-tectorius (sic) of

Parkinson (attributed by many authors to Solander, but never published by him; |

Parkinson wrote it as a monomial hence it is illegitimate) which appeared in 1773 |

(Journ. Voy. South Seas ... H.M.S. ‘Endeavour’ p. 46). There is no mention

of any collection by either of the Forsters, although certainly they must have

seen many Pandanus plants while they were in Tahiti.

BIBLIOGRAPHY

Forster, J. R. and G. 1776. Characteres Generum Plantarum, Londoni.

Reede tot Draakestein. Henricum van. 1678-1703. Horti Indici Malabarici.

Cytogenetics and Taxonomy of the Genus Globba L.

(Zingiberaceae) in Malaya

IV Distribution in Relation to Polyploidy

Lim Srew-Nco*

Genetics Division, School of Biological Sciences

University of Malaya**, Kuala Lumpur, Malaysia

Abstract

Of the twelve species, five subspecies and eight varieties of Globba in Malaya, some are

exclusively allotetraploid (2n = 32, AABB), some exclusively hexaploid (2n = 48, AAABBB),

one decaploid (2n = 80) and some with both allotetraploid as well as hexaploid chromosome

traces. Globba species with only 2n = 32 occur in lowland forests and at moderate elevations,

while those with only 2n = 48 generally in montane forests. Both are of rather restricted and

localised distribution when compared to taxa with 2n = 32, 48. In the latter, the hexaploid

infraspecific taxa are usually but not strictly montane in distribution. Where both the

tetraploid and hexaploid taxa are montane in distribution, the hexaploids are of greater

abundance. The reverse is true where tetraploid taxa occur in lowland forests and hexaploid

in montane regions. Globba species in Malaya thrive in moist, partially shaded niches. An

exception is G. marantina L. which colonises open habitats and has a wide distribution from

India and Philippines to the Solomon Islands.

The genus Globba L. (Zingiberaceae) is confined in distribution to the eastern

Himalayas and southern China southwards to Malaysia and the Solomon Islands

(Holttum 1950, Pendleton 1949). Of the 128 species recorded (in Index Kewensis,

compiled up to 1966), twelve species, five subspecies and eight varieties occur in

Malaya. These include eight new taxa which are described in Lim (1972 a). Of

these Malayan taxa, three species, two subspecies and five varieties are exclusively

allotetraploid (2n = 32, AABB), three species and a subspecies exclusively

hexaploid (2n = 48, AAABBB), one species decaploid (2n = 80) and three species

and a subspecies with both allotetraploid as well as hexaploid chromosomes races.

(see Table. 1; Lim 1972 b). This paper presents the results of distributional studies

of these Malayan taxa with special reference to their chromosome numbers.

Materials and Method

Sources of information for distribution mapping were the 363 herbarium

specimens on loan from the Botanic Gardens, Singapore, and personal field

collection data. Cases in which identification was doubtful and handwritten

data dubious were rejected.

In the mapping of each taxon, a distinction was made between herbarium

and field collections. Herbarium collections are represented by solid circles,

semi-circles and squares and field collections by outlined symbols. A single

dot represents one to three collections from the same station, an enlarged

symbol four or more. A large number of collections need not necessarily

* Now Dr (Mrs) Chong Siew-Ngo nee Lim

** Current address: Department of Biology, Nanyang University, Singapore.

115

116 Gardens’ Bulletin, Singapore — XXVI (1972)

represent increased density of occurrence of the taxon, as it might well represent

over-collection. Absence of dots might merely mean under-collecting. With the

available information, the maps (figs. 1, 2 and 3) represent what is known of

Globba distribution in Malaya to date.

No attempt was made to distinguish between the polyploids in the herbarium

specimens. Taxa with 2n = 48 were distinguished from those with 2n = 32

in the mapping of field collections.

Root tips as well as pollen mother cells were investigated for chromosome

counts. Details are as described in Lim 1972 b.

Information on the sources and chromosome numbers of fresh specimens

are in Table 2 on p. 124 Herbarium specimens of these are deposited in the

Herbarium, Botany Division, School of Biological Science, University of Malaya,

Kuala Lumpur.

Table 1. Intraspecific polyploidy in the Malayan species of Globba.

Species ange neyo Species with 2n = 32, 48

(a) Species with 2n = 32 on ="S7 an = 9

1 G. fragilis Lim 1 G. pendula Roxb. G. pendula Roxb.

2 G. albiflora Ridl. G. pendula var. —

G. albiflora var. elegans (Ridl.)

aurea Holtt. Holtt.

3 G leucantha var. G. pendula spp. —-

peninsularis Holtt. montana (Ridl.)

4 G. unifolia Ridl. Lim.

2 G. patens Miq. G. patens Mig.

(b) Species with 2n = 48 G. patens var. -——

1 G. curtisii Holtt. costulata

2 G. holttumii Lim Lim

G. holttumii ssp.

aurea Lim 3 G. cernua Bak. G. cernua Bak.

G. cernua ssp. G. cernua =s

crocea Lim crocea Lim

1 G. marantina L. porphyria Lim

4 — G. variabilis Ridl.

(d) Species with unknown G. variabilis se

chromosome number ssp. pusilla

1 G. fasciata Ridl. Lim

Distribution

The distribution of Globba species with two anther appendages is recorded

in figure 1 while that of Globba species with four anther appendages is recorded

in figures 2 and 3.

Globba species with only 2n = 32 or 2n = 48 show a rather restricted

distribution compared to those with 2n = 32, 48.

G. fragilis Lim (2n = 32) appears to be confined to Pulau Langkawi, Kedah

(see fig. 3), G. albiflora Ridl. to Penang, G. albiflora var. aurea Holtt. to Gua

Lambok and Sungei Betis in Kelantan, and G. unifolia Ridl. with its subspecific

taxon to Trengganu and Kelantan. G. unifolia also occurs in Thailand. Trengganu

might be its southern limit of distribution. G. fasciata Ridl. of unknown

chromosome number, also has a very restricted distribution (see fig. 1).

Cytogenetics and Taxonomy of the Genus Globba L. 117

G. curtisii Holtt. (2n = 48) has so far been found only in a restricted part

of the Main Range, between Bukit Kutu, the Gap and Fraser’s Hill (see fig. 3).

Further investigation might show that its distribution extends to other parts of

the Main Range. G. holttumii Lim grows in large clumps in restricted localities

in the Gombak Forest Reserve, Selangor (c.2500’), Genting Simpah, Selangor

(2700’), Genting Highlands, Selangor (c.4000’) and Fraser’s Hill, Pahang (c.4000”).

Both G. curtisii and G. holttumii are markedly montane in their distribution.

G. holttumii ssp. aurea Lim, however, was collected from lowland Dryobalanops

Forest, Lenggor Forest Reserve, Johore (c.300’).

Four of the twelve species of Globba in Malaya show 2n = 32 and 48 and these

are relatively widespread in distribution.

G. pendula Roxb., “the commonest Globba in Malaya, occurring in all

parts of the country.” (Holttum, 1950) was collected from all along the west coast

of Malaya from Pulau Langkawi in the North to Singapore in the South (see fig.

1). Distribution seems densest in Perak, Selangor, Negri Sembilan and Malacca.

On the eastern side of the Main Range, collections are from the interior of

Pahang and from Kelantan. None of the 118 herbarium collections came from

Trengganu and coastal Pahang. One single collection came from Pulau Tioman,

off the Pahang coast. The species has a predominantly lowland distribution.

Field specimens with 2n = 48 are from Fraser’s Hill, Pahang (c.4000’). Chromo-

some races of 2n = 32 occur alongside those with 2n = 48 by the waterfall,

near the old dairy farm on Fraser’s Hill. Field collections of specimens solely

with 2n = 32 were made along the west coast from the foot of Gunong Jerai,

Kedah to around Kuala Lumpur. The chromosome race with 2n = 48 appears

to be confined to montane areas. Where G. pendula Roxb. occurs, plants are

found in abundance. This is also true of the subspecific taxa: G. pendula var.

elegans (Ridl.) Holtt. in the Fraser’s Hill region of Pahang and G. pendula ssp.

montana (Ridl.) Lim in Penang. Outside Malaya, the species extends to India,

Sumatra, and the Banka Islands.

Common on the west coast from Penang to Malacca, in both lowland and

montane forest is G. patens Miq. (2n = 32, 48) a species hitherto misidentified

as G. aurantiaca Mig. see Lim (1972 a). Chromosome races with 2n = 32 seem

confined to lowland forest up to c.2000’, while those with 2n = 48 are from

Frasers Hill, Pahang (c.4000’), Bujang Melaka, Perak (c.2200’), Genting

Highlands, Pahang (c.4000’) and Bukit Lagong Forest Reserve, Selangor

(c.200-800’). Excepting the last, G. patens with 2n = 48 appears mainly montane

in distribution. Single herbarium collections have been made from Johore and

Kelantan. A field collection of a narrow-leaved and nearly glabrous form was

made from Lenggor Forest Reserve, Johore (c.200’). This appears similar to

the form collected from Ulu Kahang (Holttum, S.F.N., 10927), and has 2n = 32.

At Fraser’s Hill, Pahang (c.4000’), Cameron Highlands, Pahang (c.4750’),

and Sungei Lallang Forest Reserve, Selangor (c.250’), G. patens var. costulata

Lim was found. It resembles the specimens from Bukit Kutu in Selangor by

Ridley (mentioned in Holttum, 1950, p.37). This variety (2n = 32) apparently

occurs in both lowland and montane forests.

Overlapping in distribution with G. patens Miq. is G. cernua Bak, which

also consists of chromosomal races with 2n = 32 and 2n = 48. Globba cernua

Bak., however, is confined to montane forests between 2000’ and 5000’, within

the region between Temangor, Perak and Gunong Angsi, Negri Sembilan. South

118 Gardens’ Bulletin, Singapore — X XVI (1972)

of this, one single collection was made from Singapore. Both chromosomal races

occur in abundance in Fraser’s Hill, Pahang and Maxwell’s Hill, Perak. More

common are plants with 2n = 48 than those with 2n = 32. On Maxwell’s Hill,

the two chromosomal races occur together in mixed stands. In Cameron High-

lands, only plants with 2n -- 48 have so far been found. At Reid’s Third Quartz

Ridge, 12th mi, Gombak Road, Selangor all plants collected had 2n = 32

chromosomes. Fruiting specimens in the herbarium collections from Larut Hills

and Maxwell’s Hill in Perak, and Gunong Angsi in Negri Sembilan suggest that

plants with 2n = 32 are also present.

Two new subspecific taxa, G. cernua ssp. crocea Lim (2n = 32, 48) and

G. cernua ssp. porphyria Lim (2n = 32), were collected from Jenka Forest

Reserve, Pahang (lowland Dipterocarp forest, c.200’) and Bujang Melaka, Perak

(c.1370") respectively. The two chromosomal races of G. cernua ssp. crocea were

found side by side in Jenka Forest Reserve. Abundant occurrence along the

jungle path from Kuala Tahan to Kuala Trengganu in the National Park in

Pahang is reported by G. Smith in 1968 (personal communication).

G. variabilis Ridl. (2n = 48), shows a rather widespread distribution from

Gunong Korbu (Perak) on the western side of the Main Range and Kuala Klah

(Kelantan) on the eastern side of the Main Range to Singapore (see fig. 3). It

occurs mainly in lowland forest but also in montane areas. All living specimens

collected from four main localities: Fraser's Hill, Pahang (c.4000’), Ulu Gombak

Forest Reserve, Selangor (c.2500’), Sungei Lallang Forest Reserve, Selangor

(c.250’) and Bukit Timah Nature Reserve, Singapore (c.500’) had somatic

chromosome numbers of 48.

The relatively high occurrence of fruiting materials in 9.8 per cent of the

121 herbarium sheets in the Singapore Botanic Garden collection seems to

suggest the possibility of the existence of a chromosome race with 2n = 32.

The possibility of these having 2n = 48, however, could not be completely ruled

out.

A new subspecies, G. variabilis ssp. pusilla Lim (2n = 32) was collected from

Gunong Panti (1500’), Sungei Kayu and Sungei Sedili in Johore. So far,

the distribution appears limited to the southern tip of the Malay Peninsula.

An apparent exception to the general finding (that species with 2n = 32

show a rather restricted distribution compared to those with 2n = 32 and 48)

is G. leucantha Miq. Brief cytological investigations of G. leucantha var. peninsularis

Holtt. from two localities in southern Johore disclosed somatic numbers of 32.

Globba leucantha shows a widespread but discontinuous distribution in the north

and extreme south of Malaya (see fig. 1). The relative paucity of both herbarium

and field collections may be related to the seasonal dormancy of the species.

Globba leucantha var. peninsularis occurs in southern Johore and Singapore,

reportedly “common in forest” (Holttum 1950). A few collections were from

southern Trengganu, northern Pahang and Perak and two field collections

were from Gunong Panti, Johore and Bukit Timah Nature Reserve, Singapore.

G. leucantha var. bicolor Holtt. appears to be confined to the eastern part of the

Main Range, in northern Pahang, southern Kelantan and Trengganu. G. leucantha

var. violacea (Ridl.) Holtt. has been collected from Perak; and G. leucantha var.

flavidula (Ridl.) Holtt. from Gunong Panti, Johore (100’). Although a direct

cytological investigation of G. leucantha var. bicolor and var. violacea has not been

conducted, the rather high incidence of fruiting materials in the two varieties seems

to suggest that they have somatic numbers of 32.

0 f= ei ae ie

—— cl ss eta iterate

—

ee eet

———

res

i (ine ee

‘PERLIS (Rs.

Miles

0 10 #20 30 40 50

eee ee ee ee ee

6°

00"

SOUTH CHINA

SEA

4°

00' |

BON

g PULAU e

LANGKAWI

THAILAND

tien Ha

fs)

Land between 3,000ft.

and 5,000 ft.

eee Land between 1,000 ft

and 3,000 ft.

Globba albiflora Ridl.

Globba albiflora var. aurea

60)

Globba fasciata Ridl.

Globba leucantha var. peninsularis Holtt. (fresh

Globba leucantha var. violaceae (Ridl.) Holtt.

Globba leucantha var. bicolor Holtt.

Globba leucantha var. flavidula (Ridl.) Holtt.

Globba pendula Roxb.(> 4 herb. coll.)

Globba pendula Roxb. (1-3 herb, coll.)

Globba pendula Roxb. ( 2n=32, fresh coll.)

Globba pendula Roxb. (2n=48, fresh coll.)

pendula var. elegans (Ridl.) Holtt.

Globba pendula var. montana (Ridl.) Lim

Globba unifolia Ridl(herb. coll.)

Globba unifolia Ridl. (fresh coll.) " :

> SINGAPORE

Globba unifolia var. sessiliflorg Holtt. -

Fig. 1 Distribution map of Globba species with two anther appendages in Malaya.

@QOe2®22°* @ae Ko *

Miles

OR 1020) 300740 50

_ aS ee ee

SOUTH CHINA

SEA

Land: above 5,000 ft.

Land between 3,000 ft.

and 5,000 ft.

Land between 1,000 ft.

and 3,000 ft.

cernua Bak. s.s.(>4 herb.coll.)

ex ioe

cernua Bak. s.s.(1-3 herb. coll.) Pee NE SE

| @. - Aes

e Fs

ole)

cernua Bak. ss. (2n=32, fresh coll.)

i@ 1o

cernua Bak. s.s. (2n=48, fresh coll.)

. cernua . porphyria Lim (2n= 32)

. cernua . erocea Lim (2n=32)

. cernua . erocea Lim (2n=48)

. patens Mig. (>4 herb. coll.)

i JOHORE

ig. (1-3 herb. coll.)

iq. (2n= 32, fresh coll.)

iq. (2n=48, fresh coll.) ite).

_ costulata Lim (2n=32, fresh coll.) ¢

SINGAPORE

2008 @ ¥H+O -°*ol7 Ill

Fig. 2 Distribution map of Globba species with four anther appendages (G. cernua Bak. and G. patens Miq.) in Malaya.

Ny | Po

IPERLIS “Gp...

we ‘i THAILAND Miles

PULAU Rea 0 10 20 30 40 50

ea ; anne:

LANGKAWI

PENANG

ones

M.S

SOUTH CHINA

i] Land above 5,000 ft.

Land between 3,0001tO.

and 5,000 ft.

Land between 1,000 ft. 4

and 3,000ft.

Curtisii Holtt.(herb. Coll.) We

. Curtisii Holtt. (fresh colt.) Se

fragilis Lim - A EP

¢,

Holttumii Lim 2S 8 JOHORE

Holttumii ssp “aurea Lim

Marantina L.

variabilis Ridl. (herb. Coll.)

Globba variabilis Ridl. (fresh colt.)

Globba yariabilis ssp. pusilla Lim oo

SINGAPORE

Fig. 3. Distribution map of Globba species with four anther appendages (G. curtisii, Holtt., G. fragilis Lim, G. holttumii Lim. G. marantina

Cytogenetics and Taxonomy of the Genus Globba L. 123

A comparison of the distribution of Globba species with only 2n = 32

and those with only 2n = 48 shows that, with the exception of G. leucantha,

both are rather restricted and localised in occurrence. Species with 2n = 32

have been found on lowland forests and at moderate elevations, while species

with 2n = 48 usually occur in montane forests. The exception was G. holttumii

ssp. aurea (2n = 48) found so far in lowland Dryobalanops forests in north-

eastern Johore.

G. marantina L. (2n = 80) was found in Ayer Itam, Penang and Jason

Bay, Johore. Living materials for investigation were collected from the waterfall

Botanic Garden, Penang. The paucity of collections may be attributable to its

seasonal dormancy for about four months annually when it is nowhere to be

seen. G. marantina is apparently localised in distribution in Malaya, but has a

very widespread distribution in the Indo-Malaysian region — from India (G.

strobilifera Zoll. & Mor. = G. marantina L., G. bulbifera Roxb. = G. marantina

L., fide Index Kewensis, 1895, herbarium sheet examination of G. bulbifera Roxb.

in comparison to living kpecimens of G. marantina L.) to Sumatra (G. bracteata

Heyne = G. marantina L., fide Index Kewensis, 1895) to the Philippines, New

Guinea, the Moluccas Islands and the Solomon Islands where G. marantina is

the only species of the genus. Pendleton (1949) surmised that since the Pulau

Islands, west of Solomons, were very similar to the Solomons, G. marantina might

be there too. The ecological success of G. marantina may be attributed to its pro-

pagation by rhizome and bulbils, both being very resistant to adverse conditions,

and its ability to colonise efficiently open habitats where competition for survival

would be less keen. The other Malayan Globba species thrive well only in moist,

partially shaded niches.

A study of intraspecific distribution in the four Globba species with 2n =

32, 48 shows that the subspecific taxa with 2n = 48 are generally but not strictly

montane in distribution. Almost all the chromosomal races with 2n = 32 in G.

pendula Roxb. s.s. are lowland in distribution, the one with 2n = 48 is solely

montane. Those of G. patens Miq. s.s. show a similar distributional distinction,

with the exception of the single collection of specimens with 2n = 48 from Bukit

Lagong Forest Reserve, Selangor (c.200-800’). Both the chromosomal races of

G. cernua Bak. s.s. are restricted to the highlands; but those of G. cernua ssp.

crocea are found in lowland forests. In contrast, G. variabilis Ridl. s.s. (2n = 48)

occurs both in the lowlands as well as in montane area, though predominantly

in the latter. The subspecies pusilla has been collected from localities of moderate

elevation.

The ecological spread of chromosomal races with 2n = 32 of G. pendula

Roxb. s.s. and G. patens Mig. s.s. was found to be more extensive than that of

chromosomal races with 2n = 48. In G. cernua Bak. s.s., where the two chromosomal

races occur together in montane areas, plants with 2n = 48 are more abundant

than plants with 2n = 32.

124 Gardens’ Bulletin, Singapore — X XVI (1972)

Acknowledgements

This paper contains material extracted from a thesis accepted for the degree

of Ph.D. in the University of Malaya. I wish to record my gratitute to Professor

R. E. Holttum (Royal Botanic Gardens, Kew) and Dr. B. C. Stone (School of

Biological Sciences, University of Malaya) for their kind assistance and advice.

I wish to acknowledge the generous loan of herbarium specimens from the Botanic

Gardens, Singapore, the Royal Botanic Gardens, Kew and the Central National

Herbarium, Calcutta. I would also like to thank all those who have helped in the

collection of live specimens and Miss Kuan Lai Wah for her efficient typing of

this manuscript.

Table 2. Source and Chromosome Numbers of Living Specimens of

Malayan Globba Species.

G. marantina L.: PENANG, Waterfall Gardens. lowland, LIM Siew-Ngo, KLU

4831, 2n = 80.

G. cernua Bak. PAHANG, Western Hill, Fraser’s Hill, 4000’, LIM Siew-Ngo,

KLU 4844, 2n = 32.

———., idem, KLU 4843, 2n = 48.

—_—., idem, KLU 4832, 2n = 48.

—, Bukit Peninjau, Fraser’s Hill, 4000’, LIM Siew-Ngo,

KLU. $205. 20 =..42.

———., idem, M.£.D. POORE, KLU 4796, 2n = 48.

—_—, Parit Falls, Cameron Highlands, 4700’, LIM Siew-Ngo,

KLU 8233, 2n = 48.

—., Robinson Falls, Cameron Highlands, 4400’, LIM Siew-Ngo,

KLU 8235, 2n =-48.

—, Gunong Jasar, Cameron Highlands, 4500’, LIM Siew-Ngo,

KLU 8222, 2n = 48.

, idem, *, 2a. 48:

PERAK, Maxwell’s Hill, 3750’, LIM Siew-Ngo, KLU 4833, 2n = 32.

—_—., idem, KLU 4834, 2n = 48.

SELANGOR, Reid’s 3rd quartz ridge, Gombak, 12th mi., 1800’, A.

LETHBRIDGE, KLU 8228, 2n = 32.

G. cernua ssp. PAHANG, Jenka Forest Reserve, 200’, M.E.D. POORE, KLU 4791,

crocea Lim an FZ.

—_——., idem, I. CLEAR & C. C. HO, KLU 4817, 2n = 48.

G. cernua ssp. PERAK, Bujang Melaka, 1370’, K. JONG, KLU 8240, 2n = 32.

prophyria Lim

G. unifolia var. KELANTAN, Sungei Lebir, 300’, B. C. STONE, KLU 7367, 2n = 32.

sessiliflora Holtt.

G. fragilis Lim KEDAH, Pulau Langkawi, lowland, K. C. CHEANG, KLU 4847,

Ut = "S72.

G. curtisii Holtt. PAHANG, Waterfall by old dairy farm, Fraser’s Hill, 4000’, K.

JONG, KLU 4793, 2n = 48.

Cytogenetics and Taxonomy of the Genus Globba L. 125

G. holttumii Lim

G. holttumii ssp.

aurea Lim

G. X intermedia =

G. patens xX

cernua Lim

G. patens Miq.

G. patents var.

costulata Lim

G. variabilis Ridl.

G. variabilis ssp.

pusilla Lim

, idem, KLU 4822, 2n = 48.

SELANGOR, Gombak Forest Reserve, 2500’, LIM Siew-Ngo,

“2 =<40,

—_—-, Genting Simpah, 2700’, K. JONG, *, 2n = 48.

——., Genting Highlands, 4000’, LIM Siew-Ngo, *, 2n = 48.

JOHORE;. Lenggor Forest Reserve, 300’, P. C. LEE, KLU 8206,

2n = 48.

PAHANG, Parit Falls, Cameron Highlands, 4700’, LIM Siew-Ngo,

KLU 4840, 2n = 48.

——., Robinson Falls, Cameron Highlands, 4400’, LIM Siew-Ngo,

KLU 4841, 2n = 48.

—-——, Gunong Jasar, Cameron Highlands, 4800’, LIM Siew-Ngo,

KLU 4842, 2n = 48.

———., idem, KLU 4848, 2n = 48.

SELANGOR, Gombak 22nd mi., 1500’, LIM Siew-Ngo, KLU

BALL. 2 = 32.

———, Genting Simpah, 1800’, LIM Siew-Ngo, KLU 8219,

20y = 92;

———., Kanching Dryobalanops Forest Reserve, lowland, Honours

students, KLU 4827, 2n = 32.

— ———, Ulu Langat Forest Reserve, 700’, T. WHITMORE, *,

2 = 32.

——, Gombak Forest Reserve, (12th mi.), 1500’, A. LETH-

BRIDGE, KLU 4849, 2n = 32.

JOHORE,, Eengcor Forest: Reserve. 2007, P. C.. Lee, *, 2n = 32.

PAHANG, by Golf Course, Fraser’s Hill, 4000’, K. C. CHEANG,

KLU 6232, 2a = 46.

—, Genting Highlands, 4000’, B. C. STONE, KLU 6588,

2n ‘= 48.

PERAK, Bujang Melaka, 2200’, K. JONG, *, 2n = 48.

SELANGOR, Bukit Lagong Forest Reserve, 200-800’, M. E. D.

POORE, *; 2n'= 48.

PAHANG, Waterfall by old dairy farm, Fraser’s Hill, 4000’, A.

LETHBRIDGE, KLU, $209, 2n = 32.

—-—---, Cameron Highlands, 4750’, W. L. CHEW, KLU 8231,

Zhe. 32.

SELANGOR, Sungei Lallang Forest Reserve, Kajang, 250’, J.

DRANSFIELD, *, 2n = 32.

SELANGOR, Gombak Forest Reserve (12th mi.), 1500’, LIM Siew-

Ngo, KLU 8208, 2n = 48.

- Sungei Lallang Forest Reserve, Kajang, 200’, J. DRANS-

FIBLD, :*,.2n°= 48.

PAHANG, Fraser’s Hill, 4000’, LIM Siew-Ngo, *, 2n = 48.

SINGAPORE, Bukit Timah Nature Reserve, 500’, LIM Siew-

Ngo, *, 2n = 48.

JOHORE, Gunong Panti, 1500’, W. L. CHEW, KLU 8246, 2n = 32.

126

G. albiflora Ridl.

Gardens’ Bulletin, Singapore — XXVI (1972)

PENANG, Penang Hill, 1300’, LIM Siew-Ngo, KLU 8221, 2n = 372.

G. pendula Roxb. SELANGOR, University Campus, Kuala Lumpur, lowland, LIM

Siew-Ngo, KLU 4799, 2n = 32.

——-—-—-, Kanching Dryobalanops Forest Reserve, lowland, Honours

students, KLU 4813, 2n = 32.

-, Genting Simpah, 1800’, K. Jong, KLU 8213, 2n = 32.

———, Ayer Hitam Forest Reserve, 350, P. C. LEE, KLU

5201) 2 =" 32:

PERAK, Ulu Kinta, off Ipoh, lowland, P. C. LEE, *, 2n = 32.

PENANG, Waterfall Gardens, lowland, LIM Siew-Ngo, KLU

S210, 20 =" 32.

KEDAH, Foot of Gunong Jerai, lowland, LIM Siew-Ngo, KLU

8225, 2n = 32,

, Jungle south of Gunong Jerai, lowland, LIM Siew-

Ngo, KLU 8226, 2n = 32.

PAHANG, Waterfall by old dairy farm, Fraser’s Hill, 4000’, LIM

Siew-Ngo, KLU 8203, 2n = 48.

—— —.,, idem KLU 4836, 2n = 48.

G. pendula var.

elegans (Ridl.)

Holtt.

PAHANG, idem, LIM Siew Ngo, KLU 4830, 2n = 32.

G. pendula ssp. PENANG, Penang Hill, 1200’, LIM Siew-Ngo, KLU 8219, 2n = 32.

montana (Ridl,) ———., Waterfall Gardens, lowland, LIM Siew-Ngo, KLU 8216,

Lim Zu. = 32:

——., Ayer Itam Dam, 1090’, LIM Siew-Ngo, KLU 8229,

2n =="SZ,

G. leucantha var.

JOHORE, Gunong Panti, 1500’, J. SINCLAIR, KLU 8200, 2n = 32.

peninsularis Holtt.

SINGAPORE, Bukit Timah Nature Reserve, 500’, LIM Siew-

Ngo, KLU 4839, 2n = 32.

Bukit = Bill

Gunong = Mountain

Sungei = River

Pulau = Island

5 = Sine specimen exsiccaitum

LITERATURE CITED

Holttum, R. E. (1950) The Zingiberaccae of the Malay Peninsula. Gard. Bull. Sing.

XIII (1): 1-249.

Lim Siew-Ngo (1972 a) Cytogenetics and Taxonomy of the genus Globba L.

(Zingiberaceae) in Malaya I. Taxonomy. Notes Roy. Bot. Gard. Edinburgh

31(2) : 243-269.

(1972 b) Cytogenetics and Taxonomy of the genus Globba L.

(Zingiberaceae) in Malaya II. Cytogenetics. Notes Roy. Bot. Gard. Edinburgh

31(2) : 271-285.

Index Kewensis — An Enumeration of the Genera and Species of Flowering Plants

(1895) Vol. I. Oxford.

——— - (1901-1906 to 1966) Supplements I to XIII. Oxford.

Pendleton, R. C. (1949) “The rain shadow effect on the plant formation of

Guadalcanal.” Ecol. Monogr. 19: 75-93.

Two new Theaceous plants from Malaysia and a proposal

to reduce Tutcheria to a synonym of Pyrenaria

HSUAN KENG

Department of Botany

University of Singapore, Singapore

1. Introduction

Late 1960, I examined several specimens, incl. Elmer 21628, 21832, and

Wood SAN 16482 (Plate 1) from North Borneo which is now known as Sabah,

a state of Malaysia. These were in the Herbarium of Singapore Botanic Gardens

and identified as Camellia lanceolata (B1.) Seem. A study of the fruit (a 3-loculate

capsule with thin woody wall, partly dehiscent) and the seed (exalbuminous, the

embryo with large thin, contorted cotyledons) reminded me of those of Pyrenaria

and Tutcheria. Later, through Professor C.G.G.J. van Steenis, I learned from

Mr. J. R. Sealy, that the identification of these specimens as Camellia lanceolata

was erroneous. Dr W. Meijer, the former Forest Botanist from Sandakan, Sabah,

kindly sent all the theaceous specimens belonging to this group on loan to

Singapore. But I failed to find any flowering material of this plant. He also

informed me that the forest in Tawau area, where the collections were made,

had since been felled.

Early 1970, when checking some unidentified theaceous specimens, my

attention was drawn to a specimen from the Malay Peninsula (Corner s.n., on

Sept. 9, 1937, from Sungei Tahan, Pahang) (Plate 2). It had for many years been

preliminarily identified as Euphorbiaceae, but Mr. H. K. Airy-Shaw of the Royal

Botanic Gardens, Kew, wrote on an annotated sheet ‘‘cf. Theaceae’’, when he

worked over the un-named Singapore sheets of Euphorbiaceae in February 1969

in preparation for the chapter in the new Tree Flora of Malaya. The fruit and

seed structures of this plant reminded me again of those of Pyrenaria and Tutcheria.

At my request, this specimen together with another old collection from the same

area (Haniff & Nur 8067) and recognized by Ridley as a new species of Pyrenaria,

was sent to Mr. Airy-Shaw for further confirmation. He agreed that these two

specimens represent a hitherto undescribed species belonging to the Pyrenaria-

Tutcheria complex.

Originally, I planned to collect at least the flowering material of the Malayan

plant. During a recent trip (June, 1971) to Pahang in Malaya with Dr. T. C.

Whitmore, we reached the lower end of Sungei Tahan. Unfortunately, our efforts

to find this plant failed. I decided then that it would be better to describe these

two species of Pyrenaria (one from Sabah, one from Malaya) without further

delay. A discussion of the reason why Tutcheria should be merged with Pyrenaria

with a special note on their fruit, seed and seedling characters follows the

description.

127

128 Gardens’ Bulletin, Singapore — X XVI (1972)

Fig. | Pyrenaria tawauensis H. Keng, sp. nov.

Habit sketch, fruit and seed (based on Wood SAN 16482, Supplemented by Elmer

21628)

Two new Theaceous plants from Malaysia 129

2. Two new species of Pyrenaria from Malaysia

1. Pyrenaria tawauensis H. Keng sp. nov. (Plate 1, figure 1)

Arbor ad 10 m alta. Ramuli puberuli. Folia disticha, subcoriacea. anguste

elliptica vel anguste ellipto-lanceolata, 10-19 cm longa, 44.5 cm lata, apice acuta

vel caudata, basi acuta, supra glabra, subtus verrucosa vel fere glauca, margine

repanda, nervis lateralibus 8-12, bene intra marginem anatomosantibus; petiolo

ca. 1 cm longo. Capsula ovoidea vel subglobosa, 3—3.5 cm longa, 2.5—3.5 cm in

diametro; pericarpio tenuiter lignoso. Semina atropurpurea, 1.2—2.5 cm longa.

A small or medium-sized tree, 10 m tall. Branches slender, puberulous.

Leaves alternate, distichous, thin coriaceous, narrowly elliptic or narrowly elliptic-

lanceolate, 10-19 cm long, 4—4.5 cm wide, acute or shortly to long-caudate at the

apex and gradually or abruptly narrowed to the acute base, shining glabrous above,

verrucous, puberulent or sub-glaucous beneath; lateral veins 8-12 pairs, oblique,

curved and merged into the submarginal vein, slightly depressed above, elevated

beneath; margin remotely serrulate or repandous; petiole about 1 cm long, slender,

puberulent. Flowers not seen. Fruit broadly ovoid to depressed globose, 3-3.5 cm

long, 2.5-3.5 cm across, verrucous, puberulous or glabrescent, usually 3-loculate.

Pericarp thin woody (2-3 mm in thickness in mature specimen) eventually partly

dehiscent along the sutures. Seeds usually 2 (rarely 3) in each locule, dorsally

convex-rounded, the other surfaces variously angulate due to mutual compression,

1.2-2.5 cm long, 1.2—-1.4 cm broad, dark purple, shining.

Specimens examined:

Sabah (North Borneo), Malaysia: Tawau, alt. 100 m, G.H.S. Wood SAN

16482 (Type, Sing.) Oct. 28, 1955; A.D.E. Elmer 21628, 21832 (Sing.), Oct.

1922 to March 1923; C.E. Carr 26387 (Sing.) March 5, 1933.

This species is characterized by the fruit with thin, woody, partly dehiscent

pericarp.

2. Pyrenaria pahangensis H. Keng sp. nov. (plate 2, figure 2)

Arbuscula c. 4 m alta. Ramuli glabrescenti. Folia pseudo-verticillata,

membranacea, elliptica vel anguste oblongo-oblanceolata, 25-38 cm longa, 8-15

cm lata, apice acuta vel breviter caudata, basi cuneata, imo subcordata, supra

glabra, subtus glabrata, margine serrulata vel subintegra, nervis lateralibus 7-8,

bene intra marginem anastomosantibus; petiolo 0.25-0.5 cm longo. Capsula

Ovoidea vel subglobosa, 3-4.5 cm longa, 2.5-4 cm in diametro; pericarpio

cartilagioseo. Semina badia, 1.5-2 cm longa.

A shrub or small tree, 3-4 m tall. Branches stout, glabrescent. Leaves pseudo-

verticillate (i.e., 3-5 spirally congested to form a false whorl), membranaceous,

elliptic to narrowly oblong-oblanceolate, 23-38 cm long, 8-15 cm wide, acute or

shortly caudate at the apex and gradually narrowed to the cuneate and cubcordate

base, shining, glabrous above, puberulent or glabrescent beneath; lateral veins

7-9 pairs, oblique to nearly perpendicular, curved and merged into the submarginal

vein, rather faint above, distinct and slightly elevated beneath; margin remotely

serrulate, nearly entire below the middle; petiole short, pulvinoid, 0.25—-0.5 cm

long. Flowers not seen. Fruit broadly ovoid or subglobose, 3-4.5 cm long and

2.54 cm across, normally 3-loculate. Pericarp very thin, cartilaginous, (1-2 mm

in thickness in the mature specimen), eventually partly dehiscent along the

sutures. Seeds usually 2 in each locule, dorsally convex-rounded, the other surfaces

variously angulate due to mutual compression, 1.5—-2 cm long, 1.2-1.8 cm broad,

chestnut brown, shining.

130 Gardens’ Bulletin, Singapore — X XVI (1972)

Specimens examined:

Pahang, Malaysia: Sungei Tahan, E.J.H. Corner s.n. (Type, Sing.) Sept. 9,

1937 (Common shrub or treelet, in streams, off the main river); Teku, Gunong

Tahan, Mohamad Haniff & Mohamad Nur S.F.N. 8067 (Sing.) June 21, 1922

(Small tree, 10-12 ft. high).

This species is characterized by its sub-verticillate, short-petiolate leaves with

a subcordate base and especially by its fruit with thin cartilaginous and partly

dehiscent pericarp.

3. A proposal to reduce Tutcheria to a synonym of Pyrenaria with

a special note on their fruit, seed and seedling characters

The genus Pyrenaria was established by C. L. Blume (in Bijdr. 1119) in 1827,

based on the Javanese species, P. serrata Bl. It is interesting to note from the

original description, (““Pomum subglobossum, depressum, carnosum, guinqueloculare,

loculis 2 pyrenis. Pyrenae l-spermae...’’) that Blume recognized the fruit as a

pome, with 2 pyrenes in each of the 5 locules. From this character, the generic

name Pyrenaria was derived. Most authors today, however, generally described the

fruit as succulent, baccate or drupaceous. About twenty binomials have since

been described, they were from E. India, Burma, Malay Peninsula, Borneo,

Sumatra, Indo-China, Thailand and S.W. China (Yunnan). Some of them would

probably be merely synonyms after critical revision; for example, out of 4 species

described from Java only Blume’s original species stands (cf. Backer & Bakhuizen,

Fl. Java, 1: 321, 1963).

The genus Tutcheria, on the other hand, was established by S. T. Dunn

(in Jour. Bot. 46: 324) in 1908, based on a Hongkong species, T. championi

Nakai (formerly as T. spectabilis (Champ.) Dunn). Dunn clearly pointed out

that the systematic position of Tutcheria is closely next to Pyrenaria which it

resembles in flower and seed and from which it is distinguished by the dehiscence

of its capsule and by its multiovulate locules. About ten speciecs of Tutcheria

were described, mostly from Southern China, one from Formosa, and one from

the Riukiu Islands.

In Melchior’s monographic study of Theaceae in Engler’s Pflanzenfamilien

(2nd ed. 21: 126, 1925), Pyrenaria and Tutcheria were treated separately as

belonging to two different subtribes. Recently, in Sealy’s Rev. Gen. Camellia

(p. 13, 1958) brief discussion on the classification of the subfamily Camellioideae

(as Tribe Camellieae), Pyrenaria, Tutcheria, Camellia and Yunnania were treated

together under a subtribe, Camelliinae.

Since Yunnania is rather imperfectly known, the following brief discussion

on flower, fruit, seed and seedling characters is concentrated on those of Camellia,

Pyrenaria and Tutcheria.

I. On flowers

In general, the flowers of Pyrenaria and Tutcheria, like those of Camellia,

are variable to a certain extent. Whereas the subdivision of Pyrenaria into two

sections for instance was based on the number and degree of fusion of the styles

(Melchior, 1.c. 138); the classification of Camellia into 12 sections (Sealy 1.c. p. 28)

was primarily based on a pair of characters: styles united or free, and bracteoles

and sepals clearly differentiated or not. A full range of variation in flower parts

of Camellia was also given by Sealy (1.c. pp. 16-18). No attempt has been made

to subdivide Tutcheria.

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13]

Two new Theaceous plants from Malaysia

Fig 2. Pyrenaria pahangensis H. Keng sp. nov.

Habit sketch, fruit and seed (based on Corner s.n. [Sept. 9, 1937], supplemented by

Haniff & Nur SFN 8067).

132 Gardens’ Bulletin, Singapore — X XVI (1972)

The number of locules in an ovary in all these three genera varies from 3-5.

One particular point is the number of ovules per locule. Sealy mentioned (1.c. p.

18) that in Camellia, each locule contains 3-4 (very rarely 8) axile ovules attached

near the apex. Dunn gave the number of ovules in each ovary in Tutcheria as

‘2-5’. Although Blume’s original description states that there are two ovules per

locule in Pyrenaria, the occurrence of three or four ovules in each locule in this

genus is in fact not uncommon. Therefore, there is no real difference between

Pyrenaria and Tutcheria in the number of ovules per ovary-locule as Dunn once

unduly stressed.

2.. On fruit

The fruit in Camellia is a woody capsule, loculicidally dehiscent from the

apex downwards (Sealy, l.c. p. 18). A typical form of the fruit of Pyrenaria (e.g.

P. acuminata, plate 3, c) is baccate, the pericarp fleshy or soft woody, indehiscent,

3-5-loculate. On the other hand, that of Tutcheria, like Camellia, is a woody

capsule, loculicidally dehiscent into 3-6 valves, the latter often deciduous and

falling from the base.

The fruit characters of Pyrenaria and Tutcheria will be further discussed

below.

3. On seeds and seedlings

The seeds of Camellia, Pyrenaria and Tutcheria are uniformly ex-albuminous.

They are generally globose, hemispheric or plano-convex in Camellia, and much

flattened, often irregularly angulate (due to mutual pressure) in Pyrenaria and

Tutcheria. The most noticeable fact is that in Camellia, the embryo possesses a

pair of thick, hemispheric cotyledons, while in Pyrenaria and Tutcheria, it possesses

a pair of very thin and large, but contorted and twisted cotyledons.

Following Martin’s phylogenetic classification of seeds (Amer. Midl. Nat.

36: 513-660, esp. 519-523, 1946), Camellia seed belongs to the ‘investing’ type,

and Pyrenaria and Tutcheria seed, the ‘folded’ type, both types are classified in

his ‘axile’ division, and both represent the zenith of seed evolution (see his Fig. 3

on p. 523). Morphological differences of these two types are obvious, nevertheless,

their selective advantages are probably even more striking. In the ‘investing’ type,

the two thick cotyledons generally remain in the seedcoat and embedded in the

ground, serving as a food storage organ for the initial growth of a seedling. In

the ‘folded’ type, in contrast, the two thin cotyledons, during the process of

germination, gradually emerge from the seedcoat, rapidly unfold and spread out,

thus performing the function of photosynthesis. Previously the present writer

observed the germination of species of Camellia and Tutcheria (Keng in Univ.

Calif. Publ. Bot. 33: 276, Fig. 3 A & B). Photographs of seedlings of two Malayan

species of Pyrenaria are presented in Plate 3 (e to A) of this paper. As expected,

the germination processes of Pyrenaria and Tutcheria so far as it concerns the

few species observed, are identical.

From the above discussion, it can be concluded that Pyrenaria and Tutcheria,

as Dunn (l.c.) pointed out, are in fact, closely related. This can be further

emphasized by their peculiar seed and seedling characters which are unique

among the family Theaceae (cf. Keng, l.c. p. 280). It is also clear that their only

substantial difference lies in their fruit character i.e. baccate, indehiscent, in the

typical forms of Pyrenaria, and capsular in the typical form of Tutcheria. Is this

fruit character so vital that demarcation of these two genera can be based on it

alone? A negative answer could emerge from the following four examples.

Uh,

Why

Wy,

Plate 3. Two species of Pyrenaria

P. acuminata Planch.: a-flowering branch; b-flower and flower-bud: c-fruits; d-seeds:

e & f-seed germination; g-seedling.

P. kunstleri King: h-seedling.

Two new Theaceous plants from Malaysia 133

a. Lobelia-Pratia (Campanulaceae or Lobeliaceae).

Traditionally, these two were treated as two separate genera merely on fruit

characters alone: apically 2-valved capsule in Lobelia Linn. and baccate in

Pratia Gaudich. Moeliono & Tuyn (Fl. Mal. I, 6(1): 122, 1960) recently fused

them into one genus.

b. Buddleja-Nicodemia (Loganiaceae).

Likewise, the fruit of Buddleja Houst. ex Linn. is a septicidal, 2-valved capsule,

and Nicodemia Tenore, a berry. The latter was reduced into a subgenus of the

former by Leenhouts (Fl. Mal. I, 6(2):336-337, 1962).

c. Euphorbia-Elaeophorbia (Euphorbiaceae).

The fruit is drupaceous in Elaeophorbia Stapf., and capsular in Euphorbia

Linn, Recently Webster (Jour. Arnold Arb. 48:397, 1967) reverts Elaeophorbia

to Euphorbia, but makes a separate section of the genus for it.

d. Lomatophyllum-Aloe (Liliaceae).

Fruit of Lomatophyllum Willd. is a large fleshy berry, whereas that of Aloe

Linn. is a dehiscent capsule. Based on ‘‘Phenetic diagrams’? from the use of a

computer, Rowley (in Taxon 18:625—7, 1969) strongly queried the validity of

keeping the status of Lomatophyllum as a genus merely on a single character basis.

Undeniably, in a number of taxa (such as Leguminosae, Umbelliferae,

Ulmaceae, Juglandaceae, to mention only a few), the fruit is a paramount

diagnostic character. However, in the four examples cited above, and also in

case of Pyrenaria and Tutcheria, the present writer tends to believe that the

fruit character, in these instances, is probably merely of ecological or physiological,

rather than of phylogenetic importance.

To strengthen the argument that Pyrenaria and Tutcheria should be merged,

the present writer would like to supply the description of two types of fruits from

Pyrenaria species which appear to bridge the gap between baccate and capsulate

fruits.

The first type of fruit can be found in the two new species of Pyrenaria

described in this paper, both from the Malaysian region. Their fruit walls are

thin woody or crustaceous, partly dehiscent along one or few sutures in a

rather irregular way when fully matured. The second can be found in a Chinese

species, Pyrenaria camellioides Hu from Yunnan which is characterised by the

fruit with spongy pericarp partly dehiscing septicidally from above, and partly

dehiscing loculicidally from below. The apex of this fruit is slightly concave,

with 5 bosses (which represent the style bases) around the depression, a situation

extremely similar to the Thai species, P. garrettiana Craib. The fruit of the latter,

however, is succulent and not dehiscent.

Incidentally, Pyrenaria garrettiana Craib, together with two other species

from Yunnan, has been segregated into a new genus, Sinopyrenaria Hu (in Act.

Phytotax. Sin. 5:281, 1956) and Pyrenaria camellioides Hu, into another new

genus, Glyptocarpa Hu (in Act. Phytotax. Sin. 10:25, 1965). Meanwhile, a third

new genus, Parapyrenaria H. T. Chang (in Act. Phytotax. Sin 8:287, 1963), was

proposed based on P. hainanensis H. T. Chang, from Hainan, China. All these

three are probably superfluous, if we accept a reasonably broader generic concept ot

Pyrenaria, comparable to that of Camellia as recently defined by Sealy (I. c. pp.

14-372). :

For the reasons given above, the present writer therefore formally proposes

to merge Tutcheria Dunn with Pyrenaria Blume, and to reduce the following known

species of Tutcheria to Pyrenaria.

134 Gardens’ Bulletin, Singapore — X XVI (1972)

Pyrenaria Blume, Bijdr. 1119, 1827; Mig. FI. Ind. Bat. 1:493, 1855, et Suppl.

1:484, 1861; Hook. f. Fl. Brit. Ind. 1:289, 1872; Melch. in Engl. & Prant.

Nat. Pflanzenfam. ed. 2, 21:138, 1925; Back. & Bakh. f. Fl. Java 1: 321,

1963.

Synonym: Tutcheria Dunn in Jour. Bot. 46: 324, 1908, 47: 197, 1909; Nakai

in Jour. Jap. Bot. 16: 708, 1940; Li, Wood. Fl. Taiwan, 597, 1963; Hu

& Chang in Chun, Chang & Chen, FI. Hainan. 1:495, 1964. Syn. nov.

A list of new combinations follows.

1. Pyrenaria championi (Nakai) H. Keng comb. nov.

Basionym: Tutcheria championi Nakai in Jour. Jap. Bot. 16:708, 1940.

Synonyms: Camellia spectabilis Champion ex Benth. in Jour. Bot. 3:310,

1851 (non Berlese 1843).

Tutcheria spectabilis (Champ.) Dunn in Jour. Bot. 46:324, 1908.

Distribution: Hongkong.

2. Pyrenaria greeniae (Chun) H. Keng comb. nov.

Basionym: Tutcheria greeniae Chun in Jour. Arnold Arb. 9:129, 1928.

Distribution S. China (Kwangtung).

3. Pyrenaria hirta (Hand.-Mazz.) H. Keng comb. nov.

Basionym: Gordonia hirta Hand.-Mazz. Anz. Akad. Wiss. Wien. 58:180,

1920.

Synonyms: Tutcheria hirta (Hand.-Mazz.) Li in Jour. Arnold Arb. 26:64,

1945.

Tutcheria villosa Wu in Engler, Jahrb. 71:192, 1940.

Distribution: S.W. China (Hupeh, Kweichow, Kwangsi).

4. Pyrenaria microcarpa (Dunn) H. Keng comb. nov.

Basionym: Tutcheria microcarpa Dunn in Jour. Bot. 47:197, 1909.

Distribution: S. China.

5. Pyrenaria multisepala (Merr. & Chun) H. Keng comb. nov.

Basionym: Tutcheria multisepala Merr. & Chun in Sunyetsenia 2:41, 1934.

Distribution: S. China (Hainan).

6. Pyrenaria ovalifolia (Li) H. Keng comb. nov.

Basionym: Tutcheria ovalifolia Li in Jour. Arnold Arb. 25:209, 1944.

Distribution: §. China (Hainan).

Two new Theaceous plants from Malaysia 135

7. Pyrenaria shinkoensis (Hayata) H. Keng comb. nov.

Basionym: Thea shinkoensis Hayata in Jour. Coll. Sci. Tokyo 30 (1):45,

1911,

Synonyms: Camellia shinkoensis (Hayata) Cohen-Stuart in Meded. Proefst.

Thee 40:68, 1916.

Tutcheria shinkoensis (Hayata) Nakai in Jour. Jap. Bot. 16:708, 1940.

Distribution: China (Taiwan).

8. Pyrenaria symplocifolia (Merr. & Metcalf) H. Keng comb. nov.

Basionym: Tutcheria symplocifolia Merr. & Metcalf in Lingnan Sci. Jour.

L672 P93:

Distribution: S. China (Kwangtung).

9. Pyrenaria virgata (Koidz.) H. Keng comb. nov.

Basionym: Thea virgata Koidz. in Bot. Mag. Tokyo 32: 252, 1918.

Synonyms: Camellia virgata (Koidz.) Makino & Nemoto, Fl. Jap. (ed. 2):

740, 1931.

Tutcheria virgata (Koidz.) Nakai in Jour. Jap. Bot. 16:708. 1940.

Distribution: The Riukiu Islands.

The geographic range of Pyrenaria, after the merging of Tutcheria species

into it, thus covers from E. India, Burma, Thailand, Indo-China to S. China, the

Riukiu Islands, Taiwan and to Malesia (Malaya, Borneo, Sumatra, Java), and

conforms closely to those of Camellia (Sealy, 1. c.), Coptosapelta (Steenis, in Amer.

Jour. Bot. 56: 806, 1969) and a host of other genera.

ACKNOWLEDGEMENTS

I would like to express my sincere gratitude to. Mr. H. K. Airy-Shaw,

Dr. R. C. Bakhuizen van den Brink, Jr., Dr. P. W. Leenhouts and Professor

C. G. G. J. van Steenis for useful discussions, responsibility for the views

expressed in this paper, however, is solely mine. I also like to thank the directors

of the Singapore Botanic Gardens and the curators of the herbarium for the

facilities provided, to Dr. C. X. Furtado for going through the Latin descriptions,

to Dr. T. C. Whitmore for arranging a field trip to Pahang, Malaysia, to my wife,

Mrs. Ro-siu Ling Keng for preparing two habit sketches, and to Mr. D. Teow

for taking the photographs which are reproduced in the plates of this paper.

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Mount Maguiling Bryoflora (Luzon)

By P. T1xIER*

Résumé L’auteur analyse ses récoltes personnelles faites au Mont Maquiling en 1965. Cette

liste comprend 34 espéces de Mousses et 42 espéces et variétés d’Hépatiques. Il y a

16 Mousses nouvelles pour la station et une pour la Science. Les chiffres concernant les

Hépatiques s’élévent 4 35 et 5 espéces et variétés nouvelles. Les espéces et variétés nouvelles

sont: Endotrichella maquilinensis; Cololejeunea pseudoschmidtii, CC. haskarliana var.

luzonensis et var. thermarum, C. selaginellicola, et C. armata.

Summary The author analyses in this paper his collections from Mt. Maquiling of 1965. This list

contains 34 species of Mosses and 42 species and varieties of the Liverworts. Of the mosses 16

are new for this hill station and one for science. Liverworts number 35 as novel records and

new species and varieties 5. The species and varieties described here as new are:—

Endotrichella maquilinensis; Cololejeunea pseudoschmidtii, C. haskarliana var. luzonensis

and var. thermarum, C. selaginellicola, and C. armata.

This paper contains the list of the author’s collections on Mt. Maquiling of

10th July 1965, consisting of numbers 1358 to 1457, a complete set of which is

deposited at the “Laboratoire de Cryptogamie, Muséum National d’Histoire

Naturelle” in Paris. The reader is reminded that Mt. Maquiling is a dead volcano,

rising up to 1100 m, on the side of Laguna de Bay — the inland lake of central

Luzon — at sixty kilometers from Manila. The annual rainfall at the College of

Agriculture in Los Banos at the foot of the hills reaches 1930 mm annually for

179 rainy days (figure I). The botanical aspect of this hill station is renowned by

the work of the great north American botanists early in the century: Merrill,

Copeland and Elmer.

I acknowledge with thanks The South East Asia Treaty Organisation which

permitted me to stay in The Philippines, and Dr. D. Umali, Dean of The College

of Agriculture, who kindly gave the material for my collecting trip. In addition,

I express my gratitude to Professor J. V. Pancho of the Applied Botany

Department. From his experience and knowledge of the mountain I profited

much when he guided us in the forest of the volcano. To Dr. C. X. Furtado,

Botanist in Singapore, I am grateful for his corrections of the Latin diagnosis in

this paper.

I MUSCI

1. Fissidens sylvaticus Griff. — Corticolous, no. 1435.

Thailand, Cambodia, Vietnam, Sumatra, Java, Celebes, Borneo, Hongkong,

Bataan, Luzon (M.Q.).

2. Leucoloma molle (C. Muell.) Mitt. — On twigs and trunks, nos. 1410, 1443,

1444, 1436.

Widely distributed from Indo Malaya to Pacific Islands, Hawaii, China,

Japan, Luzon (Maquiling and Mountain Province), Mindoro, Palawan, Mindanao.

3. Dicranodontium uncinatum (Harv.) Jaeg. — On twigs, no. 1403.

India, Thailand, Japan, Moluccas, Luzon (Benguet and Mt. Banahao).

* Laboratoire de Cryptogamie, Muséum National d’Histoire

Naturelle (Paris), and, Faculté des Sciences,

Phnom Penh (République Khmére)

137

138 Gardens’ Bulletin, Singapore — X XVI (1972)

‘5

MANILA Laguna de Bay

Tagaytay amt Maquiling , \.Mt Banahao

Taal

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ern)

MANILA and surroundings

Rain/month mm

Summer monsoon

Autumn monsoon

Temperature

YZ Dry season

OT EM AM J DOR 3 eo hep

CLIMATOGRAM of LOS BANOS

Fig. I. Southern part of Luzon. Climatology of Los Banos.

Mount Maquiling Bryoflora 139

4. Leucobryum bowringii Mitt. — Corticolous, nos. 1420, 1436.

Ceylon, Malaya, China, Japan, Formosa, Vietnam, Cambodia, Thailand,

Bataan, Luzon, Sibuyan, Negros.

5. Calymperes serratum A. Braun — Corticolous, no. 1435.

Malaya, Eastern China, Vietnam, Cambodia, Thailand, Fiji, Samoa, New

Caledonia, Luzon (MQ).

6. Syrrhopodon tristichus Nees ex Schwaegr. — On twigs, nos. 1416, 1420,

1424, 1425.

Ceylon, Sumatra, Java, Ambon, Borneo, Luzon, Negros.

7. Macromitrium cuspidatum Hampe — On trunks, nos. 1400 (c. fr.), 1410.

Sumatra, Cambodia, Java, Borneo, Luzon, Mindoro, Palawan.

8. Macromitrium semipellucidum Doz. & Molk. — On branches, nos. 1432

(c. fr.), 1433.

It is nearly impossible to separate M. semipellucidum from M. pungens.

According to Bryologica javanica only one character is different i.e. the base

of the leaf in M. semipellucidum is bordered with small cells.

Luzon, Panay, Mindanao, Vietnam, Cambodia, Sumatra, Borneo, New Guinea.

9. Rhizogonium spiniforme (Hedw.) Bruch — On trunks in low positions, nos.

1436, 1437.

Pantropical.

10. Hypnodendron vitiense Mitt. — Very common on trunks, in ridge forest,

nos. 1430, 1443, 1446, 1447.

Fiji, Solomon Islands, Luzon (MQ), Negros, Mindanao.

11. Mniodendron fusco-mucronatum (C. Muell.) Broth. — On rocks, circa

600 m; nos. 1416, 1455.

Luzon, Negros, Biliran, Mindanao.

12. Endotrichella maquilinensis P. Tixier sp. nov. — On twigs, nos. 1400 (c. fr.),

1414, 1418 (c. fr.), 1424, 1425.

13. Endotrichella compressa (Mitt.) Broth. — On twigs, nos. 1400 (c. fr.), 1436.

Borneo, Luzon.

14. Symphysodon subneckeroides Broth. — On trunk, nos. 1424, 1400.

Sumatra, Java, Borneo, Panay, Luzon, Negros, Mindanao, New Caledonia.

15. Aerobryopsis longissima (Doz. & Molk.) Fleisch. var. prostratula (C. Muell.)

Fleisch. — On twigs, nos. 1418, 1424, (c. fr.), 1399.

Luzon.

16. Aerobryidium cf. filamentosum (Hook.) Fleisch. — On twigs, no. 1403.

Near A. filamentosum by length and colour of leaves, differing only by shape

of the cells. I also found this taxon in South Vietnam.

Nilghiries, Sikkim, Ceylon, Yunnan, Szechwan, Sumatra, Vietnam, Laos.

17. Barbella cf clemensiae Broth. — On trunks, nos. 1401, 1437.

Luzon,

140 Gardens’ Bulletin, Singapore — XXVI (1972)

18. Himantocladium cyclophyllum (C. Muell.) Fleisch. — On trunks, circa

700 m, no. 1455.

Java, Celebes, Borneo, New Caledonia, Tahiti, Luzon (MQ), Mindoro, Negros,

Biliran, Mindanao, Basilan.

19. Homaliodendron flabellatum (Sm.) Fleisch. — On trunk, no. 1457.

India, Ceylon, Japan, Malaya to Hawaii Islands, Philippines, New Caledonia,

Indochina.

20. Pinnatella cf. ambigua (Bosch. & Lac.) Fleisch. — no. 1435.

This small plant, 1 cm high, may be allied to P. nana (Williams) Fleisch.

Bhotan, Burma, Sumatra, Java, Thailand, Luzon, Palawan, Negros, Mindanao.

21. Eriopus microblastus Broth

Very small specimen, epiphytic on Clastobryum cuculligerum. The drawing

of Bartram is correct but it seems better to follow Brotherus and to separate

E. microblastus from E. parviretus. The denticulation of the edge is, in the former

species, denser, no. 1409.

Luzon, Negros.

22. Chaetomitrium perarmatum Broth. — On twigs, nos. 1408, 1411, 1414, 1415.

Mountain Province, Apayo district (Luzon).

23. Thuidium glaucinum (Mitt.) Bosch. & Lac. — On twigs, no. 1400.

Himalaya, Southern India, Ceylon, Thailand, Cambodia, Vietnam, Japan,

Luzon (MQ), Negros.

24. Clastobryum cuculligerum (Lac.) P. Tixier — Whitish mat on twigs, nos.

1409, 1412 (c. fr.), 1416 (c. fr.).

Java, Thailand, Cambodia, Vietnam, Luzon (Benguet), New Guinea, New

Caledonia.

25. Trismegistia calderensis (Sull.) Broth. — nos. 1441, 1444.

Sumatra, Malaya, Luzon (MQ).

26. Acanthocladium piliferum (Broth.) P. Tixier comb. nov.

Syn.: Acanthocladium longipilum Broth. Beih. Bot. Centralbl. 28, 2, (1909),

361.

Raphidostichum eberhardtii (Thér. & Varde) Broth. Nat. Pfl. 2, 11,

(1925), 434.

Raphidostichum luzonense Broth. Nat. Pfl., 2, 11, (1925), 435.

Raphidostichum piliferum (Broth.) Broth. Nat. Pfl., 2, 11, (1925), 434.

On twigs and trunk, nos. 1426, 1445.

Species of variable size, distributed from Thailand to Philippines.

27. Warburgiella cupressinoides C. Muell. — On branches, nos. 1413, 1421.

Luzon, Panay, Mindanao, New Caledonia.

28. Acroporium diminutum (Brid.) Fleisch. — On twigs. nos. 1416, 1420.

Sumatra, Java, Malaya, Luzon (MQ), Mindanao, Moluccas.

Mount Maquiling Bryoflora 141

29. Acroporium hermaphroditum (C. Muell.) Fleisch. — On trunk, no. 1455.

Thailand, Sumatra, Java, Celebes, Moluccas, New Guinea, New Caledonia,

Luzon (MQ), Mindanao.

30. Acroporium secundum (Reinw. & Hornsch.) Fleisch. — On trunks and twigs,

nos. 1402, 1425, 1426, 1440, 1446, 1454.

Java, Sumatra, Borneo, Celebes, Ceram, Luzon (MQ); Negros, Mindanao.

31. Ectropothecium buitenzorgii (Bel.) Jaeg. — On twigs and trunks, nos. 1413,

1438.

Widely distributed in South East Asia.

32. Ectropothecium falciforme (Doz. & Molk.) Jaeg. — On trunk and twigs,

nos. 1413, 1426, 1442, 1444, 1448.

Luzon, Panay, Negros, Sumatra, Java, Lombok, Borneo, Celebes.

33. Ectropothecium ichnotocladum (C. Muell.) Jaeg. — On trunk, no. 1398.

Luzon (MQ), Bohol, Mindanao, Sumatra, Java, Borneo, Celebes, Ambon,

Vietnam, New Caledonia.

34. Isopterygium albescens (Hook.) Jaeg. — Twigs, nos. 1415, 1419, 1436.

India and Indochina, Malesia, Japan, Pacific Islands.

Ii HEPATICAE

1. Metzgeria furcata (L.) Dum. — Epiphyllous, no. 1370.

Cosmopolitan.

2. Chiloscyphus communis St. — On trunk in low position, no 1436.

Assam, Vietnam, Cambodia, Sikkim, Japan, Java, Borneo, Queensland, Luzon

(MQ).

3. Schistochila aligera Nees & Bl. — On trunk, nos. 1443, 1447.

Java, Sumatra, Moluccas, New Hebrides, Nicobar, Borneo, Ambon, Philippines,

Samoa, Vietnam, Thailand.

4. Lepidozia cf. subintegra Ludg. — no. 1439.

South East Asia.

5. Herberta cf. minima Horik. — On trunk, very poor specimen, no. 1445.

Formosa.

6. Trichocolea pluma Mont. — no. 1446.

Cosmopolitan.

7. Radula anceps Lac. — Epiphyllous, no. 1378.

Java, Luzon, New Caledonia.

8. Radula borneensis St. — Epiphyllous on Selaginella sp. no. 1359.

Southern India, Borneo, Vietnam, Cambodia.

9. Archilejeunea mariana (G.) St. — Epiphyllous, no. 1445.

Formosa, Botel Tobago, Thailand, Sumatra, Java, Sarawak, Philippines,

Celebes, New Guinea, Mariannas, New Caledonia, Solomon Islands, Tahiti.

142 Gardens’ Bulletin, Singapore — X XVI (1972)

10. Ceratolejeunea maritima St. — On twigs, corticolous, no. 1409.

Carribean region.

11. Cheilolejeunea (Xenolejeunea) ceylanica (G.) Schust. & Kachr.

Epiphyllous, no. 1405.

Tropical Asia, Ceylon, Pulau Penang, Ambon, New Guinea, Formosa, Vietnam,

Cambodia.

12. Cololejeunea floccosa (Taeniolejeunea) (L. & L.) St.

Epiphyllous, no. 1395.

Japan, Ryukyu, Formosa, China, Philippines, Borneo, Java, Sumatra, Vietnam,

Chittagong, Thailand, Cambodia, Laos, Tanzania.

13. Cololejeunea gynophthalma (Yaeniolejeunea) Bx. — Epiphyllous, no. 1396.

Sumatra, Java, Vietnam, Cambodia, Malaya.

14. Cololejeunea maquilinensis (Taeniolejeunea) P. Tixier — Epiphyllous, no.

1394. Endemic.

15. Cololejeunea pseudoschmidtii (Globigerae) P. Tixier sp. nov.

Epiphyllous, no. 1394 (TYPE).

16. Cololejeunea hispidissima (St.) P. Tixier comb. nov. (Syn.: Leptocolea

hispidissima St. Spec. Hep., VI, 243, 1923). Epiphyllous, no. 1394.

17. Cololejeunea haskarliana (G.) Bx. var. luzonensis var. nov. Epiphyllous,

no. 1375:

18. Cololejeunea haskarliana (G.) Bx. var. thermarum var. nov. Epiphyllous,

no..1377-

19. Cololejeunea selaginellicola (Venustae) P. Tixier sp. nov.

Very common, epiphyllous, no. 1383.

20. Cololejeunea armata (Venustae) P. Tixier sp. nov. — Epiphyllous no. 1383.

21. Cololejeunea flavicans (Lasiolejeunea, latebrosae) (St.) Mizutani. Epiphyllous,

nos. 1360, 1362.

Endemic of Luzon.

22. Cololejeunea aff. nymanii (Lasiolejeunea, latebrosae) (St.) Bx. — Epiphyllous,

nos. 1364, 1397.

Sumatra, Java, Borneo, New Guinea, Thailand. Vietnam, Cambodia.

23. Cololejeunea panchoana (Lasiolejeunea, latebrosae) P. Tixier — Epiphyllous,

no. 1358.

24. Cololejeunea vulcania (Lasiolejeunea, latebrosea) P. Tixier — Epiphyllous, no.

1392.

25. Cololejeunea aff. yulensis (Lasiolejeunea, latebrosae) (St.) Bx. Epiphyllous,

nos. 1361, 1363, 1368, 1384, 1388.

Malaya, Sumatra, Java, Borneo, New Guinea, Cambodia, Vietnam.

26. Colura acutifolia $.J.A. — Epiphyllous, no. 1397.

Borneo, Sumatra, Chittagong, Thailand, Vietnam.

Mount Maquiling Bryoflora 143

27. Drepanolejeunea bakeri Herz. — Very common on the leaves. Endemic of

Philippines, known also of Mt. Polis in Mountain Province.

28. Drepanolejeunea dactylophora (Nees) Spruce ex Herz. — Epiphyllous, no.

1383.

Luzon, Mindanao, Malaya, Java, Ambon.

29. Lejeunea flava (Sw.) Nees. — Corticolous.

Tropical Asia and Oceania.

30. Leptolejeunea foliicola St. — Epiphyllous no. 1360.

Malaya, Sumatra, Java, Vietnam, Cambodia.

31. Leptolejeunea schiffneri Herz. — Epiphyllous, no. 1446.

Andaman, Malaya, Sumatra, Java, Borneo, Celebes, Moluccas, Philippines.

32. Lopholejeunea applanata (R.B.N.) St. — Epiphyllous, no. 1446.

Java, Ryukyu, China, Japan, Borneo, Sumatra.

33. Lopholejeunea subfusca (Nees) St. — Corticolous, no. 1401.

Japan, Ryukyu, India, Ceylon, Malaya, Sumatra, Java, Borneo, Philippines,

New Guinea, New Caledonia, Northern and Southern America, Africa.

34. Microlejeunea cucullata (Nees) St. — Epiphyllous, no. 1339.

Tropical Asia and Oceania.

35. Ptychocoleus cumingianus (Mont.) Trev. — Corticolous, no. 1435.

Andaman, Nicobar, Sumatra, Singapore, Malaya, Java, Borneo, Philippines,

Indochina, Celebes, Ambon, New Guinea, Queensland, Marquesas, Samoa, Tahiti,

Caroline Islands.

36. Pycnolejeunea bidentula St. — Epiphyllous, s.n.

Vietnam, Formosa, Ryukyu, Java, Borneo, Philippines, New Guinea, Fiji.

37. Pycnolejeunea eximia S.J.A. & Tixier — Corticolous, no. 1432.

Close to P. fitzgeraldii St.

Vietnam, China.

38. Stenolejeunea apiculata (G.) Schuster — Epiphyllous, no. 1360, 1388.

Formosa, Central Vietnam, Java, Cambodia, New Caledonia.

39. Frullania apiculata Verd. — Epiphyllous, no. 1435.

Ceylon, Indo-Malaya to Philippines, Moluccas, New Guinea.

40. Frullania integristipula (Nees) Nees. — On trunk, 530 m. alt. no. 1433.

Java, Malaya, Philippines.

41. Frullania squarrosa (R.B.N.) Dum. — Corticolous, no. 1435.

Pantropical.

42. Frullania ternatensis G. — Corticolous, no. 1378, 1424.

Malaya, Ceylon, Sumatra, Java, Vietnam, Borneo, Celebes, Ternate, Bataan,

Philippines.

144 Gardens’ Bulletin, Singapore — XXVI (1972)

lif DESCRIPTION OF NEW SPECIES

Endotrichella maquilinensis sp. nov. (Fig. II).

Plantae caespitosae, caules flavi, usque ad 4 cm alti, apice divisi. Folia

caulina enervia, elongata, acuminata, 4-5 mm longa, 1.2 cm lata, in parte basilari

leviter plicata, apice paucidentata, auricula relative parva. Cellulae apicales pariete

incrassatae, 100 » longae, 10u latae; marginales 80 x 10 yu; centrales 90-100

vp X 10-15 p; alares scutulatae, quadrataeve, 20-30 » x 10-15 yu. Folia apicalia

cum propagulis, pariete aurantiacea, 1-1.6 mm longa, 20 p» lata. Fructus proprius

generis; seta 0.8 mm longa; theca ovalis, elongata, 2 mm longa, 0.8 mm Jata.

Bracteae perichaetiales usque ad 2.5 mm.

i TAN

ENDOTRICHELLA 1400

Fig. II. Endotrichella maquilinensis sp. nov.

1: Habit; 2-3: Leaves; 4: Leaf with propagules; 5: Auricle; 6: Leaf cells; 7: Edge

cells; 8: Denticulate apex; 9: Female inflorescence; 1/0: Capsule.

Plant epiphytic up to 4 cm tall, branched at the top, light yellow. Stem leaves

without veins, elongated, acuminate, with some folds in the basal part of the

leaf. Apex with some teeth. Auricle relatively small. Cells in apex with thick walls,

100 » long, 10 » wide; cells of the edge 80 X 10 u; cells of the central part

90-100 » X 10-15 pw. Auricle cells lozenge-shaped to rectangular 20-30 yp long,

10-15 p» wide. Leaves of the top of the stem bearing propagules with orange

coloured cell walls, multicellular, 1-1.6 mm long 20 wide. Fructification charac-

teristic of the genus, seta 0.8 mm long. Ripe capsule horizontal, 2 mm long and

0.8 mm wide, oval, elongated. Perichaetial bracts up to 2.5 mm.

Species very different from all those reported by Bartram in Mosses of

Philippines and in his subsequent papers. Characterized by the elongated shape

of leaves and the edge entire almost up to the apex. Resembles EF. dimorphophylla

Tx. from South Vietnam.

Mount Maquiling Bryoflora 145

THE COLOLEJEUNEA COMPLEX

We now arrive at a group of 6 taxa. The first is Cololejeunea hispidissima

(St.) Tx. conformable to the type of Stephani. Benedix, in 1953, rendered intricate

the question by putting together in synonymy, under the name Cololejeunea

haskarliana (G.) Bx., taxa which are evidently different. Mizutani, in 1961, reported

that C. venusta Lac. was unlike C. haskarliana (G.) Bx. sensu lato because

the Sande Lacoste’s species has papillous lobule cells. In Maquiling forest the

author observed the sympatry of the taxa allied to Cololejeunea haskarliana in

the locality and sometimes on the same leaf. In Tjibodas (Java) up to 4 different

taxa on one leaf were collected. The author intends to return to this problem.

The following three taxa are allied to Cololejeunea haskarliana: Cololejeunea

pseudoschmidtii is related to the Section Globigerae of Benedix, Cololejeunea

selaginellicola and Cololejeunea armata to Section Venustae of Mizutani.

Cololejeunea pseudoschmidtii sp. nov. (Fig. II).

Planta dioica, parva, pallide virens, sicca, ad substratum appressa. Caules

usque 0.5 cm longi, 0.10 mm crassi, cum foliis 1.4 mm lati. Folia inter se 0.2 mm

distantia. Lobus obovalis, 0.9 mm longus, 0.5 mm latus. Cellulae basalis intermedio

leviter incrassatae 20-40 pm longae, 15-20 » latae haud papillosae; marginales

quadratae hexagonales, 10-20 » longae, 10-15 wu latae, cum papillis humilibus.

Lobulus elongatus, inflatus, 0.25 mm longus, 0.15 mm latus, papillosus ad carenam.

ae Shi

Fig. II. Cololejeunea pseudoschmidtii sp. nov.

1: Stem; 2: Leaf; 3: Basal part of the lobe; 4: Edge of the lobe; 5: Young lobule;

6-8: Mature lobule.

146 Gardens’ Bulletin, Singapore — X XV] (1972)

Dens apicalis parvus, triangulari-unicellulatus. Dens medianus cum cellules in

3 series dispositis, una serie, duabus alteris unicellulatus. Papilla hyaline sphaerica,

sub dente media 20 » in diametro. Stylus haud visus. Androecia lateralia, 0.3 mm

alta, bracteis 2-jugis.

Plant dioic, small, light green, applied to the substrate. Stem 0.5 cm long,

0.10 mm thick, with leaves 1.4 mm wide. Rhizoids hyaline. Leaves spaced on stem at

0.2 mm. Lobe oboval, 0.9 mm long, 0.5 mm wide. Basal cells with light intermediate

thickenings 20-40 p long, 15-20 » wide, apapillose. Edge cells rectangular to

hexagonal, 10-20 » long, 15-20 » wide with a low papilla thickened at the top

similar to those of C. schmidtii, Lobule elongated, inflated on mature leaves,

0.25 mm long, 0.15 mm wide, sometimes papillose near the keel. Apical tooth

weak, reduced to a triangular cell. Median tooth with 3 tiers of cells, one with

3 cells, the others with one cell only. Papilla hyaline, spheric, below the median

tooth, 20 » in diameter. Male inflorescence lateral with 2 tiers of fertile bracts,

0.3 mm high.

Cololejeunea hispidissima (St.) P. Tixier comb. nov. (Fig. IV).

Small plant, light green, applied on substrate. Stem to 0.5 cm or more,

0.07 mm thick, with leaves 1.2 mm wide. Rhizoids hyaline. Leaves spaced at

0.25 mm. Lobe oval, trapeziform, 0.5 mm long, 0.4 mm wide. Basal cells making

a short pseudo-vitta, cells 20-70 » long, 10-35 » wide with a slight papilla. Cells

of the edge more or less isodiametric, with thin walls, 10-20 » x 10-16 m with

cylinder-shaped papillae.

Fig. IV. Cololejeunea hispidissima (St.) Tx.

1: Stem; 2: Leaf; 3: Basal part of the lobe; 4: Edge of the lobe; 5: Young lobule;

6 & 7: Mature lobule; 8: Propagule.

Mount Maquiling Bryoflora 147

Lobule inflated, rounded, 0.25 mm long, 0.25 mm wide, keel without papillae.

Apical tooth present on the young leaf, missing on the mature ones. Median

tooth with 2 cells, young erect, repelled on the inferior edge.

Hyaline papillae spheric, 10 » in diameter, under the first cell of median

tooth. Fugaceous stylus 16 » high and 10 » wide. Propagules discoidal with 20

cells and 65 » in diameter. Fructification unknown.

Cololejeunea haskarliana (G.) Bx. var. luzonensis var. nov. (Fig. V).

Planta dioica, parva, pallida, ad substratum appressa. Caules 0.5 cm longi,

0.07 mm crassi, cum foliis 1.1 mm lati. Rhizoidea hyalina. Folio inter se 0.2 mm

distantia. Lobus subtriangularis, 0.5 mm longus, 0.4 mm latus. Cellulae basales in

pseudo-vitta 30-40 » longae, 30-15 p» latae, cum papilla crassa usque ad 25 p

altae, marginales parvae, 10-15 » in diametro, papillosae. Lobulus ovalis 0.2 mm

longus, 0.15 mm latus, carina papillosa. Dens apicalis unicellularis, dens medinus

cellula arcuata, latere externo lobuli locata. Una cellula e dente medio usque ad

marginem posticum. Papilla hyalina subsphaerica, 10 m in diametro. Stylus

bicellularis, 40 » altus, 10 » latus. Propaguli disciformes, in statu 12 cellulares,

50 » in diam. Capitula feminea 0.4 mm alta. Bracteae florales cum lobo 0.4 mm

longo, 0.15 mm lato, et lobulo 0.3 mm longo, 0.10 mm lato. Perianthia 0.3 mm

alta, 0.2 mm lata.

Dioic plant, small, light green, applied on substrate. Stem 0.5 cm _ long,

0.07 mm thick, with leaves of 1.1 mm wide. Rhizoids hyaline. Leaves spaced on

stem at 0.2 mm. Lobe subtriangular, 0.5 mm long, 0.4 mm wide. Basal cells

making a pseudo vitta, cells 30-40 p» long and 30-15 » wide with a strong papilla

Fig. V. Cololejeunea haskarliana Bx var. luzonensis var. nov.

I: Stem; 2: Leaf; 3: Basal part of the lobe; 4: Edge of the lobe; 5: Young lobule;

6-8: Lobule; 9: Propagule; 10: Perianth; 77: Lobule of the perianthal bract.

148 Gardens’ Bulletin, Singapore — XXVI (1972)

up to 25 » tall. Cells of the edge, small, more or less isodiametric, 10-15 » in

diameter, papillose on the edge of leaf. Lobule oval, 0.2 mm long, 0.15 mm wide

papillose on the keel. Apical tooth unicellular, median tooth of one, arc shaped

cell lying on the external face of lobule. One cell between the median tooth

and the keel. Papillae spheric, hyaline, 10 » in diameter, stylus bicellular, 40 p»

high, 10 » wide.

Young (?) propagules, 50 » in diameter and with 12 cells. Female inflorescence

0.4 mm high. Perianthal bracts with a lobe 0.4 mm long, 0.15 mm wide and the

lobule 0.3 mm long and 0.10 mm wide. Perianth shorter than perianthal bracts,

piriform, 0.3 mm high, 0.2 mm wide.

Cololejeunea haskarliana (G.) Bx. var. thermarum var. nov. (Fig. VI).

Planta dioica, parva, pallide virens, ad substratum appressa. Caules usque

0.5 cm longi, 0.07 mm crassi, cum foliis 1 mm lati. Rhizoidea hyalina. Folia inter

se 0.2 mm distantia. Lobus subtriangularis, 0.4 mm longus, 0.4 mm latus. Cellulae

basales in pseudo-vitta, 30-60 m» longae, 15-30 yw latae cum papilla humili

(10 p» alta): marginales parvae, 10-20 » X 10-15 yw metientes. Papillae per

tutam marginem. Dens apicalis unicellularis, triangularis, pariete hyalinis.

Dens medinus unicellularis, rotundatus cum dente medinus decussatus: 1 cellula

unter dentem medianum et marginem posticum. Papilla hyalina subsphaerica,

25 » in diametro. Stylus unicellularis, 30 p» altus, 10 » latus. Capitula feminea

0.5 mm alta. Bracteae florales cum lobo 0.5 mm longo, 0.25 mm lato, et lobulo

0.3 mm longo, 0.15 mm lato. Perianthia piriformia, inflata, 4 plicata, 0.5 mm alta,

0.3 mm lata.

Fig. VI. Cololejeunea haskarliana Bx var. thermarum vat. nov.

1: Stem; 2: Leaf; 3: Basal part of the lobe; 4: Edge of the lobe; 5: Young lobule;

6 & 7: Lobule; 8: Perianth; 9; Lobule of the perianthal bract.

Mount Maquiling Bryoflora 149

Small dioic plant, light green, applied on substrate. Stem to 0.5 mm _ long,

0.07 mm thick, with leaves 1 mm wide. Rhizoids hyaline. Leaves spaced on stem

at 0.2 mm. Lobe subtriangular, 0.4 mm long and 0.4 mm wide. Basal cells making a

pseudo-vitta, cells 30-60 » long and 15-30 » wide with a low papilla 10 » high.

Edge cells small, more or less isodiametric, 10-20 » long, 10-15 » wide. Edge

papillose on almost all the periphery. Oval lobule, keel lightly papillose, 0.2 mm

long, 0.15 mm wide. Apical tooth triangular with hyaline walls. Median tooth

unicellular, crossed with apical tooth. One cell between the median tooth and the

edge. Papillae hyaline, spheric, 25 » in diameter. Stylus unicellular, 30 » high,

10 » wide. Female inflorescence 0.5 mm long, perianthal bracts with a lobe 0.5 mm

long, 0.25 mm wide and a lobule 0.3 mm long, 0.15 mm wide. Perianth piriform,

inflated, with 4 folds, 0.5 mm high, 0.3 mm wide.

Near to the preceding species, differing by the ornamentation of leaf cells,

the disposition of the lobule teeth, the shape of the papillae and the perianthal

bracts.

Cclolejeunea selaginellicola sp. nov. (Fig. VII).

Planta dioica, parva, pallide virens, ad substratum appressa. Caules usque

ad 0.6 cm longi, 0.10 mm crassi, cum foliis 1 mm lati. Rhizoidea hyalina. Folia

0.2 mm inter se distantia. Lobus ovalis, 0.6 mm longus, 0.25 mm latus. Cellulae

basales 30-60 » longae, 15-20 » latae, papillosae; marginales, pariete incrassatae

rotundatae 10-15 mw in diametro, cum papillis altis conicis. Lobulus magnus

0.25 mm longus, 0.15 mm latus, papillosus usque ad + latitudinis. Margo superior

lobuli cum uniseriatis cellulis hyalinais dens apicalis magnus, unicellularis, trian-

gularis, parietibus hyalinis. Dens medianus unicellularis, sphaericus, 2 cellulis

usque ad marginem posticum. Papilla hyalina parva sphaerica 7-10 p» in diametro.

Stylus tricellularis, 40 » longus, 10 » latus. Capitula feminea 0.8 mm alta. Bracteae

cum lobo 0.6 mm alto, 0.25 mm lato, et lobulo 0.45 mm longo, 0.08 mm lato,

apice acuminato. Perianthia piriformia, 4 plicata, rostro notato, 0.8 mm_ alta,

0.5 mm lata.

Dioic, small species, light green, applied on substrate. Stem to 0.5 cm long

and more, 0.10 mm thick, with leaves 1 mm wide. Rhizoids hyaline. Leaves

spaced on stem at 0.2 mm. Lobe oval, 0.6 mm long, 0.25 mm wide. Basal cells

30-60 » long, 15-20 » wide, papillose. Lobule large, 0.25 mm long, 0.15 mm wide

with an angle between the keel and the edge of the leaf. Lobule papillose, up to

4 its width. Superior edge of lobule with a row of hyaline cells finishing with an

apical, unicellular tooth, triangular, large, thin-walled. Median tooth reduced to

a spheric cell; 2 cells between the median tooth and the edge. Papillae hyaline,

small, spheric, 7-10 » in diameter. Stylus tricellular, 40 » long 10 p» wide.

Female inflorescence 0.8 mm high. Perianthal bracts with a lobe 0.6 mm long,

0.25 mm wide and a lobule 0.45 mm long, 0.08 mm wide with an acute top.

Perianth piriform with 4 folds and a strong beak, 0.8 mm high, 0.5 mm wide.

Cololejeunea armata sp. nov. (Fig. VIII).

Planta dioica, parva, pallide virens, ad substratum appressa. Caules 0.5 cm,

0.06 mm crassi, cum foliis 0.6 mm lati. Folia 0.15 mm inter se distantia. Lobus

Ovalis, magis aut minus arcuatus, 0.4 mm longus, 0.25 mm latus. Cellulae basales,

pariete pauciter incrassatae, 20-30 p» longae, 10-17 » latae, papillosae; marginales

rotundatae 10 » in diametro. Lobulus rotundatus 0.15 mm longus, 0.10 mm latus,

papillosus praeter ad basin. Margo superior lobuli cum cellulis hyalinis uniseriatim

dispositis usque ad dentem apicalem ipsam hyalinam. Dens medianus, bicellularis

150

Gardens’ Bulletin, Singapore — XXVI (1972)

Fig. VII. Cololejeunea selaginellicola sp. nov.

1: Stem; 2: Edge of the lobe; 3: Basal cells of the lobe; 4: Lobule; 5: Young lobule;

6: Perianth; 7: Lobule of perianthal bract.

Mount Maquiling Bryoflora

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Fig. VIII. Cololejeunea

armata sp. nov

I: Stem; 2: Edge of the lobe; 3: Basal cells of the lo

be; 4: Lobule; 5: Propagules;

6: Perianth; 7: Perianthal bract; 8: Lobule of the perianthal bract.

$52 Gardens’ Bulletin, Singapore — XXVI (1972)

decussatus cum dente apicali. Papilla hyalina, parva, sphaerica, 10 » in diametro.

Stylus cylindratus, 30 p» altus, 10 » latus. Propaguli disciformes, in statu 12

cellulares, 40 » diam. Capitula feminea magis aut minus lateralis, 0.4 mm alta.

Bracteae cum lobo, 0.4 mm longo, 0.18 mm lato, et lobulo 0.3 mm longo, 0.10 mm

lato, margine crenulato, Perianthia piriformia, rotundata, 4 plicata, 0.4 mm alta,

0.25 mm lata, rostro notato.

Dioic, small plant, light green, applied on substrate. Stem to 0.5 cm,

0.06 mm thick, width with leaves 0.6 mm. Rhizoids hyaline. Leaves spaced on

stem at 0.15 mm. Lobe oval, more or less arc shaped, 0.4 mm long and 0.25 wide.

Basal cells with thick walls, 20-30 » long and 10-17 p» wide; papillose edge with

isodiametric cells about 10 » in diameter, with a papilla low and cylindric. Lobule

rounded, 0.15 mm long, 0.10 mm wide, papillose except in the basal part. Superior

edge with a row of hyaline cells finishing in apical tooth cell, hyaline, thickwalled.

Median tooth of 2 cells, crossed with the apical one. The apical border of lobule

from apical tooth to keel arc shaped. Papillae hyaline, small, spheric 10 p» in

diameter. Stylus cylindric, 30 » high, 10 » wide. Propagules, 40 » in diameter

and with 12 cells. Female inflorescence on the lateral side, 0.4 mm high. Perianthal

bracts with a lobe 0.4 mm long, 0.18 mm wide and a lobule 0.3 mm long and

0.10 mm wide, the superior edge of the lobule crenulated. Piriform perianth,

rounded with 4 folds 0.4 mm high, 0.25 mm wide with apparent beak.

IV CONCLUSIONS

This study devoted to few species shows the necessity of serious reflexion

because of the complexity of Cololejeunea bryoflorula.

1. Interest of these collections

The object of this work is to find out whether these gatherings bring an

interesting contribution to the knowledge for Bryology for Mt. Maquiling. Accord-

ing to Bartram (1939) sixty species of Mosses were found at the station. My list

contains 34 species, 16 of which are new for Mt. Maquiling and one for science.

This contribution shows an increase of known mosses by 20 per cent. There arose

a few inquiries on the different gatherings of liverworts. The Herzog’s list

concerning Baker’s collections had 28 species. My own has 42, 34 of which are

new and 5 new for science. For liverworts a 50 per cent increase is found. The

two figures based on a one-day collection show that further collecting trips to

Mt. Maquiling would be worthwhile.

2. Biogeographical connections

For mosses of my list, the community floristic coefficients between Mt.

Maquiling and the different countries of South East Asia is established as

follows:—

40 Java, Sumatra, Thailand

30-40 Vietnam, Borneo, Cambodia, Celebes

30-20 Ceylon, Japan, China

20-10 Himalaya, Malaya, Hawaii, New Caledonia.

The main axis of affinities goes from Japan to Sunda Straits. Some relations

with Hawaii and New Caledonia still remain puzzling.

Mount Maquiling Bryoflora 153

LITERATURE

HeErRzoG, T. (1931): — Hepaticae philippinenses a Cl. C. Baker lectae. Ann. Bryol.,

IX, 79-94.

BARTRAM, E. B. (1939): — Mosses of the Philippines. Philip, J. Sc., 68, 1-424.

TIxiER, P. (1966): — La végétation orophile de I’Asie du Sud Est. Le Mt.

Maquiling et ses Epiphytes. — Science & Nature, no. 78, 3-11.

(1969): — De Sematophyllaceis. I-Essai de revision des Clastobryaceae

de Luzon, J. Hattori Bot. Lab., no. 32, 21-64.

(1970): — Contribution a la connaissance du genre Cololejeunea en

Asie due Sud Est. II/La section Radulae du sous genre Lasiolejeunea. Ann.

Fac. Sci. Phnom Penh, no. 3, 174-190.

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Philippine Species of Nepenthes

SHIGEO KURATA* and Masami TOYOSHIMA#*

This paper lists the results of two expeditions to the Philippines in search for

Nepenthes. It records seven species and describes one of them and three hybrids

as new taxa. All but one were collected during the three-week expedition made by

both authors to the islands of Mindanao and Luzon in August 1965. N. burkei

was collected by the senior author (S. Kurata) when he participated in the second

expedition from December 1967 to February 1968 to Mindoro Island. Some

Bornean species were also found but are excluded in this paper. The authors are

grateful to Dr. C. X. Furtado for correcting the latin diagnosis.

The new taxa are N. globamphora, N. mirabilis x alata, N. petiolata x alata

and N. truncata x petiolata.

1. Nepenthes alata Blanco, Fl. Filip., ed. 1, p. 805 (1837).

LUZON: Mountain Prov., Baguio, 1470 m. alt., Aug. 28, 1965, S. Kurata

1136-a 1136-b; en route to La Trinidad, 1500 m. alt., Aug. 28, 1965, S. Kurata

1135. MINDANO: Surigao del Sur, eastern slope of Mt. Legaspi, 270 m. alt., Aug.

19, 1965, S. Kurata 1101-a, 1101-b, 1104.

Distribution. Philippines, Malaya, Sumatra, Borneo.

Between the Luzon plants and those from Mindanao there are some differen-

ces. The former are hairy on the pitchers and tendrils. The purple blotches or

stripes are usually distributed on its pitcher wall. On the other hand, the pitcher of

the latter is strikingly narrow but dilate in the lower part, and the hair is absent

from its wall. The colour of the pitcher is mostly yellowish green and at times

becoming brownish green near the mouth.

2. Nepenthes burkei Masters. Gard. Chron. Ser. 3, VI p. 492 (1889).

MINDORO: Occidental Mindoro, Mt. Kidayap, summit, 1550 m. alt., Feb. 12,

1968, S. Kurata 2033, 2034, 2035.

Distribution. Philippines (Mindoro and Panay).

3. Nepenthes (Montanae) globamphora Kurata et Toyoshima, sp. nov.

(Fig. 1, pl. 1). — Nepenthes globamphora (nomen nudum), Kurata et Toyoshima,

Journ. Insectivorous Plants Soc. 36: p. 15-22, 1966.

Planta tenuis, breviter scandens, luteo-viridis. Caulis 30-80 cm altus, 4-5 mm

crassus, cylindricus. Folia 8-12 cm longa, 1-1.8 cm lata, lanceolata, coriacea,

sessilia, basi 4 amplexicaulia, utrinque glabra; nervi longitudinales utrinque 3;

nervi transversales obscure, oblique ascendentes; cirrhus 15-20 cm longus, cylind-

ricus, glaber, luteo-viridis saepe etiam lucido-purpureus. Ascidia 4-9 cm longa, 3-5

cm lata, subdimorphia, inferiora ellipsoidea vel globosa, superiora ampulliformia,

omnia luteo-vel, rubro-viridia, diffuse purpureo-maculata, coriacea, glabra, intus

omnino glandulosa, bialata, alis margine dentatis, dentibus 3-ciliatis; os orbiculare

obliquum: peristomium 5-8 mm latum, sub-cylindricum, aequale, luteo-viride,

margine exteriore revolutum, interiore denticulatum; operculum 2-3.5 cm latum,

* Insectivorous Plants Society, Department of Biology, Nippon Dental College, Fujimi,

Tokyo, Japan.

155

156 Gardens’ Bulletin, Singapore — XXVI (1972)

Fig. 1. Nepenthes globamphora Kurata et Toyoshima: A. male plant with flowers and

pitchers (x 0.3); B. upper portion of female plant with flowers (x 0.3); C. male

flower (x 2); D. capsules (x 1).

her on the lower stem (x 2.5).

a pitc

Nepenthes globamphora

PLATE 1

al al ORNL RN LOR i tac intl Mab tabs

Philippine Species of Nepenthes 157

cordato-orbicularum; calar ca. 1 mm longum, filiforme. Inflorescentia 10-15 cm

longa, racemosa, pedunculus 6-9 cm longus, ferruginio-tomentosus, pedicelli 3-4

mm longi, uniflori; sepala 4, ovata, 2-2.5 mm longa, 1-1.2 mm lata, extus minute

tomentosa; columna staminae sepalis brevior, glaber, antherae 8, uniseriatae; cap-

sula 17-20 mm longa, 4 mm lata, fusiformis, brunneo-nitida, glabra, valvae lanceo-

latae.

MINDANAO: Surigao del Sur, eastern slope of Mt. Legaspi, 270 m alt., Aug.

22, 1965, S. Kurata and M. Toyoshima 1128 holotype in Herb. NDC (Nippon

Dental College) 1129, 1130, 1132.

This new species is easily distinguishable from all others by the peculiar

pitcher which has an unusual roundness (hence the epithet “‘globamphora’’), and

by the trifid ciliae fringing the pair of wide wings — a very unique occurrence in

the genus. The lid has no appendix on its lower surface where nectar glands are

sparingly distributed. The glands are very small and deep. The remarkable feature

of this species is that the inner surface of the pitcher is wholly glandular. When dry

the specimen is red-brown, and the lower surface of leaves and flowers are more

reddish. The lid has a ceramic luster on its lower surface. This plant is comparati-

vely common on the eastern slope of Mt. Legaspi, but its distribution seems limited

to a narrow area.

4. Nepenthes merrilliana Macf., Contrib. Bot. Lab. Univ. Pennsylv., III,

mecu7..t. (1911).

MINDANAO: Surigao del Sur, Camp David (a lumberer’s camp, 10 miles

upper stream of the mouth of Carrascal river). 150 m alt., Aug. 20, 1965, S. Kurata

1117, 1118, 1119, 1120-a, 1120-b, 1124, 1125, 1126; eastern slope of Mt. Legaspi,

mo m alt.. Aug. 19, 1965, S. Kurata 1116, 1121, 1122, 1123, 1127.

Distribution: Philippines (Mindanao), Sulawesi.

5. Nepenthes mirabilis Druce, Rept. Exch. Cl. Br. Isl., 1916, p. 637 (1917).

MINDANAO: Surigao del Sur, on a river side near the Carrascal Bay, 20 m

alt., Aug. 19, 1965, S. Kurata 1107.

Distribution: Southern China, Indochina, Malaya, Philippines (Mindanao),

Borneo, Java, Sumatra, Sulawesi, New Guinea, Parau, Australia (York Pen.).

6. Nepenthes mirabilis x alata, hybr. nov.

Hybrida naturalis inter N. mirabilis et N. alata. Folia lanceolata, herbacea,

glabra, petiolata. Ascidia 10-15 cm longa, 3-4 cm lata, monomorphia, cylindrica,

bialata, alis fimbriatis, luteo-viridia, herbacea, glabra; os orbiculare obliquum:

peristomium 2-3 mm latum, subcylindricum, aequale; operculum 3-5 cm iatum,

orbicularum, facie inferiore prope basin appendiculatum; calcar ca. 1 cm longum,

bifidum. Inflorescentia ignota.

MINDANAO: Surigao del Sur, on a river side near the Carrascal Bay, 20 m

alt., Aug. 9, 1965, S. Kurata 1111-a (type in Herb. NDC), /1/1-b.

This specimen has thin, herbaceous, petiolate leaves and cylindrical pitchers

with orbicular lids. The pitcher is intermediate between its parental species; the

strong folds on the boundary between the waxy zone and the glandular zone are

derived from N. mirabilis whilst the appendix on the lower surface of the lid

agrees with N. alata.

158 Gardens’ Bulletin, Singapore — XXVI (1972)

7. Nepenthes petiolata Danser, Bull. Jard. Bot. Buitzg. ser. III, Vol. IX,

p. 353 (1928).

MINDANAO: Surigao del Sur, eastern slope of Mt. Legaspi, 270 m alt.,

Aug. 19, 1965, S. Kurata 1108.

Distribution. Philippines (Mindanao).

8. Nepenthes petiolata x alata, hybr. nov.

Hybrida naturalis inter N. petiolata et N. alata. Folia elliptico-lanceolata,

coriacea, glabra, petiolata. Ascidia 10-15 cm longa, 3-4 cm lata, monomorphia,

cylindrica, bialata alis fimbriatis, luteo-viridia, diffuse purpureo-maculata, coriacea,

minute tomentosa; os orbiculare obliquum; peristomium ca. 2 mm latum, sub-

cylindricum, aequale; operculum 3 cm latum, ovatum, facie inferiore prope basin

appendiculatum; calcar ca. 1 cm longum, filiforme. Inflorescentia ignota.

MINDANAO: Surigao del Sur, eastern slope of Mt. Legaspi, 270 m alt., Aug.

19, 1965, S. Kurata 1113-a (type in Herb. NDC), J1/3-b.

This specimen has elliptical leaves with petioles and cylindrical pitchers with

ovate lids. The colour of the pitcher is yellow-green with purple blotches and/or

stripes and is densely covered with short hairs which seem to be the same of that

of N. petiolata. On the other hand, the small glands and the appendix on the

lower surface of the lid seem to be derived from N. alata. A lustrous belt on the

boundary between the waxy zone and the glandular zone on the inner surface

of the pitcher is also from N. alata. Both species N. petiolata and N. alata are

found growing in the same area.

9. Nepenthes truncata Macf., Contrib. Bot. Lab. Univ. Pennsylv., II, p. 209,

t. IT (1911).

MINDANAO: Surigao del Sur, Camp David, 150-300 m alt., Aug. 20, 1965,

S. Kurata 1105, 1106, 1115.

Distribution. Philippines (Mindanao).

10. Nepenthes truncata x petiolata, hybr. nov.

Hybrida naturalis inter N. truncata et N. petiolata. Folia oblonga, apice

truncata, coriacea, supra glabra, subtus villosa, petiolata. Ascidia 10-15 cm longa,

3-4 cm lata, monomorphia, inferne ventricosa, superne cylindrica, bialata, alis

fimbriatis, luteo-viridia, diffuse purpureo-maculata, coriacea, tomentosa; os orbi-

culare obliquum; peristomium 5-8 mm latum, subcylindricum, margine exteriore

expansum, margine interiore denticulatum; operculum 4-5 cm latum, cordato-

ovatum, facie inferiore prope basin appendiculatum; calcar ca. 1 cm longum,

bifidum. Inflorescentia ignota.

MINDANAO: Surigao del Sur, eastern slope of Mt. Legaspi, 270 m. alt.,

Aug. 22, 1965, S. Kurata 1109-a (type in Herb. NDC), 1109-5.

This specimen has wide coriaceous leaves with petioles, and the lamina is

cordate at the apex. The petiole and lamina are densely covered with brownish

coarse hairs beneath. The pitcher has a wide and expanded peristome. The lid has

on its lower surface an appendix and two kinds of nectar glands: one small and

deep, surrounded with a wide rim, the other large and shallow, surrounded with

a narrow rim. The above characters show that this specimen is a natural hybrid

between N. truncata and N. petiolata. At the place where this plant was collected,

both parental species and the other i.e. N. alata were growing together.

Studies in the basidium

Spore-spacing and the Boletus spore

FE. J. H. CORNER,

Botany School, University of Cambridge

A geometrical analysis of the 4-spored homobasidium is made on the assumption that

spore-spacing is determined by sterigmatic spacing on the basidium-apex. Given, then, the

values of spore-width and the interval between adjacent spores of the tetrad, other details of

the basidium-unit can be worked out except the lengths of the spores, sterigmata and basidium.

The relations between the parts are summarised as equations; their values in ratios of spore-

width are tabulated in an appendix. Observations on Mycena illuminans with subglobose

spores are given as a check on the theory.

The smooth Boletus-spore is shown to be the unornamented endospore of a subglobose

spore with endosporic ornamentation that has suffered compression and reduced spacing

through the adaxial displacement of the sterigmata. The spores of Strobilomyces and Phyllo-

boletellus appear as the more primitive.

The use of this theory is shown by analysis of the cruciform spores of Marasmius

nigricans.

During the tumultuous years 1942-1945, when Dr Kwan Koriba maintained

the scientific status of the Singapore Botanic Gardens, I was permitted to work

with him in the upstairs laboratory of the office. We studied trees, their seasons

and their reproduction, and I was allowed a more personal investigation, which

had long intrigued me, into the construction of the basidiomycete hymenium.

Some of this work has been published (Corner 1947, 1948). It has shown that the

lengths and widths of basidia, spores and cystidia are bound by specific constants

which have generic trends. The dimensions of these parts are usually given inde-

pendently in taxonomic descriptions, but they are related in consequence of the

characteristic of the hypha that produces them. The relations can be displayed

as sporographs, basidiographs and so on, as explained in those papers, but their

interconnection lies in the dependence of spore-width on that of the basidium.

This is the matter that I now take up, and it becomes the problem of spore-spacing

on the basidium.

With many detailed observations for such genera as Amanita, Lepiota, Hygro-

phorus, Marasmius, Russula, Entoloma, Agaricus and Coprinus, I had intended

an extensive survey but the long hours necessary for such a work have never

recurred. I take it up for the special case of Boletus because in the course of my

account of the rich Boletus-flora of Malaysia (Corner 1972) the problem has loomed

continually and is fundamental to the classification of the species by means of their

distinctive spores. My thesis is the geometrical analysis of the position of the spores

on the basidium. It is not an explanation of their arrangement by physical means

but a guide to understanding how the spore grows to its full size and shape and,

indeed, how it may develop its superficial structure. I have set forth this analysis

in the simplest terms, step by step, so that it may be followed by mycologists who,

like myself, may have lost touch with advanced mathematics.

159

160 Gardens’ Bulletin, Singapore — X XVI (1972)

Most basidiomycetes have four spores on the basidium. There have been

several accounts of the development of individual spores (Corner 1948, 1968;

Malencon 1958; Perreau-Bertrand 1967). I propose to consider the tetrad as a unit.

The four spores are regularly spaced. Their arrangement can be affected by adjacent

basidia. The spacing is affected by the contour of the hymenium. Spore-shape

results from the way in which spore-volume is contained in this spacing. A prime

factor is the width of the basidium. Yet the tetrad of one basidium does not

contact that of another; the spores of a tetrad are not in contact; they are separated

from the basidium by sterigmata; maturing basidia are separated from each other

by immature. To secure this spacing there must be action at a distance. The force

appears to be electrostatic repulsion at free surfaces; the whole mechanism

collapses when wetted. To prove this, I had intended to grow basidia in an induced

electric field, but this has not materialised. I hope, however, that this article will

Open a new line of enquiry from which some real understanding of the precision

of the hymenium will come and better appreciation of the basidiocarp which

houses it. The toadstool is essentially an umbrella opened geotropically.

Method

The sporing basidium is a delicate turgid structure that readily shrinks in

contact with a fixative. As my object was to make fine measurements of spore-size

and the intervals between spores on the basidium, it was necessary to examine

undistorted living basidia under high enough magnification to make reasonably

large camera lucida drawings. I first used dry mounts of thin strips of hymenium

for surface-view and moderately thick sections for side-view, and observed them

with an oil-immersion lens. Condensation on the coverslip, however, in the high

humidity of Singapore brought failure and I had to resort to aqueous mounts.

Thus I learnt that sooner or later, after ten minutes with some species or an hour

or two with others, the apex of the wetted basidium collapsed and the sterigmata

inclined, even jumped, inwards; to be effective the basidium had to be superficially

dry. By dint of perseverance I managed to obtain for subsequent analysis many

accurate camera lucida drawings of a great variety of agarics, boleti, polypores,

stereums and clavarias. The present article merely summarises these results because

names are not available for most of the fungi, and the labour is great; about a

thousand measurements are needed for accuracy in any one species.

The two aspects of the basidium, end-view and side-view, need to be combined.

From this a geometry of the basidium emerges to relate the shape of the basidium-

apex via the sterigmata with the spore-spacing and, thus, with spore-shape. As

the spores are borne at the same level above the basidium (though there are

exceptions), one can obtain in end-view an optical section of the tetrad. Focussing

down, one can see the basidium-outline in its optical section at maximum width

and, on then focussing slightly upwards, even the tips of the sterigmata. However,

it is easier to see these tips on discharged basidia before they have collapsed.

The four spores, as seen in end-view, occupy the corners of a square and one can

imagine them to be confined by two spore-circles, an outer circumscribing circle S’

(Figure 3) and an inner inscribing circle $’”’, to which may be added a third spore-

circle S” passing through the centres of the spores. Then there is the real basidium-

circle as the outline of the basidium, which I call w, and for the sterigmata one

Studies in the basidium 161

can imagine a sterigmatic circle M passing through the tips of the sterigmata.

The circles are concentred on the long axis of the basidium. S’ exceeds the

basidium circle which can often be seen to lie within S$”; the M circle lies

slightly outside or abaxial from $”’. These real and imaginary circles, derived from

end-view of the basidium-unit, can be projected on to the basidium apex, as in

Figure 3.

Since a basidium cannot be examined both in end-view and side-view, an

average must be taken from a set of measures for each. In end-view basidia are

often seen to be compressed and ellipsoid in optical section. Such can be avoided

but they cannot be recognised in side-view, for which the average is necessary.

The figures in the following pages are based on such averages from sets of twenty

to one hundred drawings.

Following the system adopted in my previous papers, I continue with this

notation: —

nis the spore-number per basidium.

D is spore-length.

d is spore-width; d rad, when measured in side-view, and d tan, when

measured tangentially or at right angles to d rad.

is the ratio D/d.

is the length of the basidium.

“Se

w is the maximum width of the basidium.

M is the diameter of the sterigmatic circle.

m is the diameter of the sterigmatic patch circle.

S’ is the diameter of the outer spore-circle.

S” is the diameter of the middle spore-circle.

S’” is the diameter of the inner spore-circle.

(S’), (S”), (S”’) and (m) refer to the inner spore or endospore when needed

to be distinguished.

o is the ratio of spore-interval to d.

8 is the spore-angle.

5 is the angle subtended by the half-spore on the spore-circle.

«: is the angle subtended by the sterigmatic patch at the centre of the

curved apex of the basidium.

Evidence of spore-spacing

The following considerations supply the evidence that the spores of a tetrad

are regularly spaced.

Elongate spores. A tetrad of elongate spores in end-view appears as four

circles equidistant round the basidium-apex (Figure 1; Buller 1924, fig. 84-86, 91;

Corner 1964 p. 234). The long axes of the spores are parallel with that of the

basidium; the spores face this axis and are not inclined. This is the main evidence

for supposing the imaginary circles S’ and S”’ and the lateral spacing factor ‘ o’.

162 Gardens’ Bulletin, Singapore — XXVI (1972)

ZS,

OG Ff GAL

500

Figure 1. Spore-tetrads in end-view, x 800 (a. x 500); a, Oudemansiella; b, f, Amanita; c,

Pluteus; d, Panaeolus; e, Psathyra; g, Boletus ravenelii, with two normal basidia

and iwo pairs of approximated basidia having their eight spores set in an ellipse.

Interfering tetrads. Usually a basidium develops its tetrad freely at some

distance from other sporing basidia. New basidia are intercalated and take the

place of those that have discharged their spores and become ineffective. Occasion-

ally two basidia mature so close together that their tetrads are distorted and, as

seen in end-view, their eight spores are combined into one imaginary ellipse

(Figure 1). This re-arrangement shows that sporing basidia act as spacing units

and influence their neighbours. Where two S’ circles approach or would intersect

the adjacent spores of each tetrad are repulsed outwards. The unit has a boundary

at S’.

The adaxial patch. In many verrucose or echinulate spores, for which Russula

and Thelephora are good examples (Malencon 1958; Corner 1968), the warts or

spines develop all over the spore except in a small area on the adaxial side of the

spore just above the apiculus. I call this smooth area the adaxial patch. It is the

part of the spore impinging on the inner spore-circle S”’ and it shows that this

imaginary circle is a boundary defining a central ‘dead space’ over the basidium-

apex where spore-growth is forbidden. It is the sharpest boundary in the tetrad.

Tangentially compressed spores. The tetrads of Panaeolus (Figure 1) and some

species of Coprinus (Buller 1922, fig. 104; 1931, fig. 29-32), show radial or adaxial

compression of the spores; d rad is less than d tan. In contrast with globose spores

that are strictly confined by the one spacing given by S’ and S”’, they bulge

laterally; a lateral spacing has to be considered as well as a radial. More spectacular

are the trigonous and cruciform spores in various agarics (Figure 12). Evidently

there may be variation in the lateral spacing and in the intensity of inhibition

from S’ (as shown by abaxially echinulate spores), while that of S”’ is rigid. Here,

however, the exception comes with the Boletus-spore.

Studies in the basidium 163

Reversed spores. Tetrads of homobasidiomycetes do not bear reversed spores;

that is, spores do not develop with the apiculus turned abaxially. This may happen

with diads in normally tetrasporous species (Corner 1950, fig. 271). In such cases

the two spores are evidently so far apart that the sterigmatic spacing by the

sterigmatic disc is ineffective; nevertheless, S’ and S”’ operate from the basidium-

apex to maintain the spore-shape, even if o is also ineffective.

6-8-spored basidia, When n is 3, the spores in end-view fit the angles of an

equilateral triangle; when n is 4, they fit a square; when n is 5, they fit a regular

pentagon. But, when n is 6 or 8, they do not fit a regular hexagon or octagon;

they are disposed in an ellipse, as with interfering tetrads. This ellipse, however,

seems not to be plane but to have decurved ends. Four spores occupy the central

part of the ellipse and one or two spores at each end lie at a slightly lower level

through the obliqueness of their sterigmata. The same effect may be seen in the

tetrads of Russula and Lactarius. | have not succeeded in examining such hexads

and octads in any detail. The obvious material is to be found in Cantharellus,

Craterellus and Pterygellus (Corner 1966), but the basidia of these fungi are too

long for suitable study in end-view. The regular disposition of the spores proves,

nevertheless, that spacing factors operate through the ring of 6-8 sterigmata placed

peripherally at the basidium-apex with its superimposed dead space.

Towards the geometry of the basidium

Certain features of the basidium-unit need more explanation.

Positioning of the spores. They are positioned by the sterigmata. These out-

growths, at first normal to the curved apex of the basidium, become parallel with

the basidium-axis and develop at their attenuate tips the sterigmatic discs. From

these the spores, inhibited by S$”, develop abaxially to the limit of S’ and then

elongate within these limits, parallel to the basidium-axis, if they are to become

larger. The transverse sterigmatic discs appear to be the initial spacers and the

apiculus of the mature spore is the adaxial feeler which indicates, but seldom

coincides with, S’”. Careful measurement of the spore- and sterigmatic circles

have shown me that, as a general rule, M= S” — 0.7d; for a single spore, that is,

the tip of the sterigma is situated external to S”’ by a distance of 0.15d.

If spore-spacing is determined by sterigmatic spacing, the cause of this cannot

be discovered by simple observation because it must lie within the basidium.

It must be connected with the hyaline cap which forms at the apex of the basidium

shortly before the sterigmata emerge (Corner 1948). The substance of this cap

forms, evidently, the walls of the sterigmata and those of the spore-rudiments;

presumably it becomes electrically charged at the surface. It must be partitioned

into four blocks which underlie the sterigmatic patches, or bases of the sterigmata

(Corner 1948). I have pursued the geometrical consequences of the projection of

this patch on the plane of the spore-circles and its hypothetical function in spore-

Spacing (p. 169).

Spore-angle. The direction in which the spore grows from the transverse

sterigmatic disc was studied by Buller. He concluded that this angle was at 45°

to the long axis of the basidium. I have measured the angle on camera lucida

drawings and, though it is impossible to be precise with such minute structures,

I agree with Buller’s conclusion. It is supported by the following argument.

164 Gardens’ Bulletin, Singapore — XXVI (1972)

With reference to Figure 3, ca MTC is the spore-angle 0. If this is 45°, then

MO=7C tin Gr= og sin 45° = 0.35d ~

or, for the whole basidium-unit,

M = S” —2MC = S” — 0.70d

This is the value of M found by direct measurement of S”’, S’ and M. The

longitudinal plane of S”’ lies adaxially to that of M by this small fraction 0.15d.

The ratio d/w. The positioning of the sterigmata and that of the spores about

the longitudinal axis of the basidium so that the basidium circle (in end-view

of the tetrad) lies within S” imply that there is a close relation between spore-

width d and basidium-width w. These are two easy measures and their relation

is an important feature of the basidium-unit. For the smooth ellipsoid spores of

many homobasidiomycetes from Amanita to Clavaria, I find that in general

d = 0.6 to 0.65w. Large globose spores may increase the ratio to 0.8w and narrow

spores may decrease it to 0.5w. These differences may occur within a genus; the

latitude in positioning of S’ may be the explanation, but specific differences in

spore-width generally imply differences in the valve of w. Thus, a basidium 8

wide will bear spores 5u wide; one 10» wide will bear spores 6.34 wide. Despite

records to the contrary, I have not seen a tetrasporous homobasidiomycete in

which -d equals or exceeds w. The exception among the disporous is Clavulina

with d = 1.18w (Corner 1948).

No species has basidia of constant width. There is always some variation.

Hence I have used as values of w and d either the mean of their variation or,

when there have been sufficient measurements, their averages.

Lateral spacing in the tetrad. As a measure of this feature, I take the ratio ~

of the spore-interval to spore-width, as seen on the spore-circle. In Figure 3

KL WAKE

oS DS

2CK d

Measurements of the tetrads of various basidiomycetes with isodiametric

spores (d rad equals d tan) show that in general o = 0.5.

The spacing can also be expressed by the angle 6 (Figure 3), as explained

on p. 169.

The basidium-apex. The projection of the spore-circles and other items of the

basidium-unit on to the basidium-apex in median longitudinal section implies

knowledge of the curvature of the apex, but this is a difficult matter and I can give

only an approximation. The clavate basidium, fully grown but without sterigmata,

has a hemispherical apex. When the hyaline cap forms, the apex protrudes slightly

and in the long basidia of agarics with polymorphic basidia (such as Coprinus)

this projection is lengthened and becomes almost cylindric with hemispheric apex.

The question is whether it is ellipsoid or paraboloid. To test this, I measured the

diameter of the basidia with hyaline caps, yet without sterigmata, at certain levels.

These were the level of maximum width w and those at 4, 4 and ? of the vertical

distance between the apex: of the basidium and the plane of w. I compared these

results with those that can be calculated for similar chords of a sphere (or ellipse) —

and a parabola, as shown in Table 1.

Studies in the basidium 165

Table 1. Diameter of the basidium-apex, as ratios of w, at

successive levels from the apex.

Levels w 0.25w 0.5w 0.75w

ellipse 1 0.661 0.866 0.968

observed 1 0.687 0.854 0.954

parabola l 0.438 0.750 0.938

The apex is clearly ellipsoid but, as rounding off of a clavate apex with slight

protrusion, it is probably ellipsoid with a small dome of spherical curvature.

This is the dome between the sterigmatic bases that struts them and collapses

when wetted.

Globose spores. Longitudinal alignment cannot be discerned directly from

such spores but, from their positioning on the sterigmata, they clearly obey the

dead space defined by S$”. Presumably they have lateral spacing and external

limits, defined at least by the total effect of the basidium-unit.

Spore-length. A globose spore fills the space between the limits of S’ and S”’.

If more protoplasm enters the spore, it elongates within these limits. For a given

spore-volume, the closer these limits the longer the spore will be. But spores vary

in the way that they grow into and fill this space. Some are globose from the

first and then lengthen; others expand gradually as they lengthen and become

pip-shaped, obovoid or clavate. None of these factors, however, has prime signi-

ficance in spore-spacing, though the limits of S’ and S”’ explain why ellipsoid

spores vary much in length though little in width, and conform to a sporograph-

locus.

Sterigmatic length. Since the sterigmata become parallel, their length has no

immediate effect on spore-spacing. They may lengthen abnormally, however, and

remove one or two spores of the tetrad from the spacing effect of the whole, and

this may be usual in Tremellaceae, but it is exceptional in homobasidiomycetes

and leads to the abortion of the spore. Many exceptional states occur in Gastero-

mycetes with their variously disorganised basidium-mechanism.

I have been unable to detect any relation between the length of the sterigmata

and other features of the basidium-unit. The length seems to depend on the

hydrostatic pressure in the basidium and to be related with spore-number. Thus

in species with n =3 or 2 or 1, as variations on n= 4, the sterigmata are

progressively longer. Sterigmatic length can, therefore, be neglected in basidium-

geometry.

Basidium-length. With any one kind of basidium as defined by its equation

(Corner 1947) w depends on /, but 7 does not otherwise determine spore-spacing.

In all species / varies, but the resulting differences in w are so slight that the

shorter basidia bear spores of practically the same shape and size as the longer.

Thus, in tetramorphic species of Coprinus, there are not four sizes and shapes

of spores. Nevertheless, as always, there are exceptions. At least two species of

Hygrophorus have normally basidia of two sizes, very different in length, and

they bear spores of very different size (Corner 1936). For the purpose of spore-

spacing, basidium-length can also be ignored.

166 Gardens’ Bulletin, Singapore — XXVI (1972)

Figure 2. Diagrams of basidium-units, treated as cones, divergent in the convex hymenium

(upper left), convergent in the concave hymenium (upper right), and divergent

or paraboloid in the plane hymenium (lower figures).

The basidium-unit, The whole basidium with its spores can be regarded as

an inverted cone expanding from the base (inverted apex) to the value of S’.

Within the cone the unit develops; outside the cone it restrains other basidia.

The effect is well seen in clavarioid basidia. Their sterigmata often diverge slightly

and their long spores may also diverge and appear in optical section on the spore-

circle as short ellipses. I suppose that this divergence is connected with the

convex curvature of the hymenium, developed on the outside of the cylindric

or clavate fruit-body, and that this curvature makes the basidia slightly divergent

(Figure 2). In the agaric hymenium the basidium-unit becomes paraboloid with

sterigmata and spores parallel to the long axis of the basidium. But the poroid

hymenium is concave, lining tubes; the basidia are convergent and their cone-

effects interfere; the outer spore-circles of adjacent units conflict and the whole

unit is distally compressed. This is the problem in Boletus, many species of which

Studies in the basidium 167

have long and narrow spores with convex adaxial face, known as the boletoid

spore. Superficially this spore may resemble a Clavaria-spore; actually it is developed

in a more complicated manner because the sterigmatic circle is displaced inwards.

The hydnoid hymenium resembles the clavarioid. Its basidia are often small

and their divergence may explain the prevalence of globose or subglobose spores.

The stereoid hymenium with plane extension resembles that of the agaric but the

basidium-units are not strictly aligned because of the thickening hymenium; new

basidia may project beyond the influence of the mature and have slightly divergent

sterigmata.

Asymmetric basidia. Very occasionally a basidium that has grown aslant

develops a typical spore-circle asymmetrically on the exterior side of the apex.

Whether or not this is the effect of light, gravity or some other external stimulus

on the hyaline cap, the configuration proves that the sterigmatic arrangement is a

structural unit.

Measurements. I have relied on my own measurements from living material.

The basidium-unit is an inflated, semi-rigid structure prone to collapse, and it is

not certain that measurements from dried material or that fixed in alcohol-

formalin agree exactly with those from the living. It is tempting to use the

published data in descriptions, but I have avoided these not only because they

often refer to preserved specimens but because it is clear that immature basidia

are often measured and such will give misleading results.

The geometry of the basidium

The geometry, as I have analysed it, is set out in Figure 3. The upper part

shows the spore-tetrad, as seen in optical section from the end-view of the basidium,

together with the spore-circles S’, S$”, S’”, the basidium-circle w, the sterigmatic

circle M, the square joining the spore-centres C, and the tangents from the long

axis of the basidium A to the spores. Then there are shown four small circles m

on the spore-radii; they are the orthogonal projections of the sterigmatic patches

on to this transverse plane. The lower part of the figure shows the longitudinal

geometry of the basidium-apex which is drawn as a hemisphere on the base ww

of maximum width of the basidium. The right half of the basidium-apex shows

the sterigmatic patch (arc Hw) and the angle Haw which it subtends; I call

this angle « . The various circles of the upper figure are projected on to the

base-line ww. The lower right-hand spore represents a globose spore in side-view

as it would fit on to the sterigma which, for clarity, has been omitted. If the

upper figure is rotated through 90°, it represents the spore-circles above the

basidium.

Many relations of the parts can be derived from this figure. I give merely

’ the more useful that can be reduced to the minimum data d and c , when n = 4.

It must be noted that the symbols S’, 8’, S’’, M, w, d and m refer to diameters

and not radii.

168 Gardens’ Bulletin, Singapore — X XVI (1972)

—-— fe =e se ww ee Se SF es ew ewe ee SP se se as 8 =

. §

“sf

’

; :

S' S.» feos Mefab Ai os oS" Mai \Gr S$}

Figure 3. The geometry of the spore-circles in relation to that of the basidium-apex in

median longitudinal section; constructed for ¢ = 0.5, w = 1.65 d. For explana-

tion, see text.

Studies in the basidium 169

err re Te GE GE) sees ttc ies boca iv wba reassess orca accsenecsacvdanscosesevess (1)

because CC = CK + KL + LLC, and by definition KL = do, beetacate

d d

ae ee ge ot to)

en eR SoS os ca oa'y ac nc aan vals gne wndoohewelin apie buyuiewcana ait (2)

because S” = JAC and, as S” (diameter) is hypotenuse to the rt. < triangle with sides CC,

then

(2AC)2 = 2(CC)2

AC = Jae = J id “(1 + oy as from (1)

Ns ES eI MR EI a aie cakes iw tah Wigs On ncenensuansstenassavsedencesevassenedsege (3)

NY ccs cee icon adh sansa dace Mags vos esedl tiandhusddecdessiorionaees (4)

Now it is convenient to turn these relations into functions of § . In Figure 2,

P is the point where the tangent from A meets the spore.

PC d 2 d 1 .

sin § = —— = — x — = SN earas cum CECT ORE E TEE CEP CETT ETT TT SeT rere (5)

eae Seer emcee fC +c + o)

or cosecS = 2 (1 + o)

= SS POSED Sacra treanenndemiinasnnnt Oe cise Ssdvahbiuivedeos satus (6)

sin 6

d d(1 + sin 8)

S’ = 8S” + d = — + d = ————— =_ (4 Cosec§) ciccecccccccccccsecceeccecteceeeceecers (7)

sin § sin 6

o XS =d{ 20 +0) +1]

d d (1 — sin 8)

S’”’ = §S” — d = — — d = — —— = d (cosec§ — 1) ooeceiccceccccccccceccceccceeeeeeees (8)

sin § sin 6

or S”=d[ 2(1+o0)—-1]

ane PO ek a aN Seah in las sawn awianghcdnnae sumqagle ass enepeee s (9)

or M=d{[ /2 (1 +o) -0.7]

cosec §

= oe et eae ee ee eee. 0 a, (10)

Sterigmatic patch hypothesis. I assume that the spacing of the sterigmatic

patches provides that of the spores by transmission through the sterigmata. The

symmetrical partition of the hyaline cap in the basidium-apex into four areas,

set as far as apart as possible on the level of w, may be the physical basis. The

geometrical consequence must be traced by the projection of the sterigmatic patch

on to the transverse plane of the spore-circles. In Figure 3 any circle with centre

on AC’C and inscribing AP will provide the spore-spacing, but the sterigmatic

patch implies certain conditions. Its projection must lie within the basidium-

circle; it must place the sterigmatic tips on the sterigmatic circle; and it must be

related with w (that is the perpendicular ww in Figure 3). For convenience I use

m’ for the diameter of the projected circle of the sterigmatic patch and denote

the circle accordingly.

The most likely position for the centre of m is C’ where AC cuts the S”’

circle (in the upper part of Figure 3). This m circle contacts the basidium-circle,

as can be seen from the following argument with reference to Figure 4.

170 Gardens’ Bulletin, Singapore — XXVI (1972)

A C B c

oe WO) 5"

Figure 4. Construction to prove that the m circle with centre C on the S” circle contacts the

basidium circle w.

For a circle with centre on AC and with AP as tangent, then C’ P’ = C’ B,

as radii of this circle.

C’P’ = AC’ sin 6, or in this case

S’’ sin § d (1 — sin 6) sin § d (1 — sin 6)

from (8)

2 2 sin §

C’B = AB — AC’, or in this case

= 4 (W = ore)

d (1 — sin 4)

Now S’”” = from (8)

sin 6

m = S’’ sin 6 ex hypothesi

w= 5S’ + m= 8S” + S’ smo = SS’ (1 + sino)

Therefore,

d (1 — sind§) (1+ sin§6) d (1 —sin 68)

oO): ae oo Saeed Vee eS

2 sin§ 2 sind

2 sin§ C’B

— = 1 — sin2§ — 1 + sin§ = sin6 — sin2§

d (1 —sin8)

: 2.

Therefore C’B = C’P’

Studies in the basidium 171

The following equations may therefore be added

ee NE PN le 5a Vol sib Si din'n aveca vce GAibsee die ol nen dk Cees Dek cies Sou naseareanas (11)

iS ize

sah ret i ae SCN ee SINR ir sa ca dda ced aernscannevr eves “Weastuondoecdioesteamteths (12)

This curious relation between w and the spore-circles follows from that

between w and d as determined by o and 6. Thus

w= 8” + m and S” = S” —d

m = S’” sinS =(S” — d) x — =

ic” d a d2

Therefore w = (S” — d) + —————

(Sys dts’ di Oe (8”.4.¢d) 6" —d)

S 4” S 4 AY ”

AS S74

= from (3)

ce ih eek eS Ss) Pn | ea (13)

d (1 — sin 8) d (1 — sin2 §)

because S”’ (1 + sin §) = ———————- x (1 + sin §) = ———————_

sin 6 sin §

d cos2 §

= = d cots§ coss

sin 6

The longitudinal relations may now be considered.

Let the sterigmatic patch circle m be projected on to the basidium-apex,

as in the lower part of Figure 3. It occupies arc Hw and subtends 2 HAw or «

Bisect « and the bisecting radius AF will meet the arc Hw where it is intersected

by the projection of the sterigmatic circle M. That is, if the centre of the

longitudinal sterigmatic arc on the basidium is projected parallel with the long

axis of the basidium up to the level at which the spores are developed, it becomes

the sterigmatic tip, whence the spores grow abaxially with the spacing initiated

on the basidium-apex. This is one of the most remarkable agreements between

the geometry of the basidium and the observation of the spore-circles, the sterig-

matic circle and the spore-angle. At first there seems to be no reason for the

position of the sterigmatic tips, other than structural convenience; now it is seen

to be an integral part of the basidium-unit. The tips are eccentric in the sterigmatic

patch circle. It would be interesting to compare this construction with the basidium

of Gasteromycetes with sessile or subsessile spores.

By means of the angle « , other relationships in the basidium can be found

which have value in checking reconstruction from specific data.

172 Gardens’ Bulletin, Singapore — XXVI (1972)

ford

MEW COG vga ksinasnoesvancce'esy ous Geallatineaasmasencie ceuee ene atabadenninte caees yee (14)

1 — sin §

OS oa ssn nnasiee an nen un ote hina mmc oaebanesns ale ae eee (15)

1 + sing

Am 2Am 2Am

because cos“« = ——— = = —_______- from (12)

AH w S’”’ (1 + sin 8)

A ig m Ss” S”’sng% Soer— ea.)

Am = — —- — = — — — = from (11)

z 2 2 2 Z

Therefore

S’”” (1 — sin 6) 2 1 — sin §

COS 0 = —AA > X« =

2 S’” (1+ iné) 1+ sin6g

SY a m

Alternatively, Os OC Se mn isis ige sn cucigeapas canes seine s sean alvs eile cana (16)

Ss” +m

1 — cos «©

sin § = Fronts (1S) oa ccsscsnexcesenucanshumesenadeemes ss hen tes aaenpege eg aden (17)

1+ cos«

S” —m

ST a riiiccdvuscevcns usanachen eon 0 sy aoanne aapheduusan sw ickiens sco entinngean sued: emus aeaaennnnnnnnn (18)

MC S”—M 2 S’—m

because sin 9 = —— = —————- x —- =

Bi, 2 d d

5S” —M

d = ——____- froma (16) auncbincsviincc ee tRD Mewes coneweue cee (19)

sin 6

1 + cos«

M = w / ———-——_, from (14) erie ALAR An oso ecoe nce (20)

=w /————— from, (15)

1 + sin$

d (1 — sin 6)

mn macnn ee eR ee (21)

sin §

do d (i — sin 8)

because S’”’ = S” — d = —— — d = —___—_—_-

sin 6 sin 6

y SU

M =d cot§ cos§_ ,/————— Sram: 420) ated, 4) 3)..2..<cresere weeks elated (22)

\/ 1+ sin§g

This, at last, gives M a value in terms of spore-spacing.

sin § = cosec § — cot§ cos gf sins ov uvita'b An vO px 6 Salenslvink noe eceaena mabealie eee Wee alae n ee anne (23)

1 + sin§

Studies in the basidium 173

This, at last, gives 8 in terms of spore-spacing. The proof is detailed, thus

S” —M d

sin § = from (18), because sing =——

S” sin § ’

S” —M

whence sind =

S” sin 6

S” —-M

Now for = |——— write k

i ore

Then sin 6 = k cosec§ and sing = k cosec @, and from (22)

M d cos2 § 1 sin 6 1

k=1-—=1- ————— x = 1 — cos2§ Ue Jer

bid sin 6 1 + sin$ d V 1+ sin§

Therefore

sin. = cosee 5 ( 1 — cos2 6 /—— )

1 + sing

= cosec § — cot§ cos § /————

1+ sind

These equations are summarised in Appendix 1. The values of the various

factors are given in Appendix 2. With the three data d, o and n = 4, all other

parts of the basidium-unit can be calculated except the lengths of the spores,

sterigmata ‘and basidium.

If the values of the factors, as given in Appendix 2, are plotted against

those of « as graphs (which is the easiest method of obtaining intermediate valucs),

it will be found that they form very gradual curves. This stability of form is the

character of the homobasidium, which is one of the more constant structures in

botany.

The curve sin 8 = f o has its greatest curvature where ¢ = 0.73 = sin 6

The curves sin 8 = f o and sin §: = f o intersect at o = 0.807 when sin

me == )S1n.2.8.=5) 0.72.

When « = (0.5146, there is the special case in which the diameter of the

basidium equals the circumference of the sterigmatic patch circle; that is, w = 7m.

Thus

m=S” sin§ and w= S” +m=S"’ (1 + sin 8)

w 1 + sing

Therefore — = — = J2(1 +o)+1 from (5)

m sin 6

=2414+ /2¢

If this expression is to give the value of =, then

V20 = x —2.414 = 0.7276, and « = 0.5146.

Also, at this value of o, the length of the arc subtended by « equals 1.004d,

which is practically the value of d. In other words, at the commonly observed

value of o, the longitudinal arc of the sterigmatic patch equals the spore-width,

and the diameter of the basidium, if rolled up into a circle, equals the circum-

ference of the sterigmatic patch circle. Here, again, it would seem that the volume

of the hyaline cap of the basidium must be the basic cause.

174 Gardens’ Bulletin, Singapore — XXVI (1972)

Mycena illuminans P. Henn., as a test

This small luminous agaric, not uncommon in Singapore (Corner 1954), has

fairly large, almost globose spores and the sporing basidia are fairly resistant to

immersion in water. I used it, therefore, as a test of the theory. However, the

basidium is large and the gill-tissue opaque, with the result that I could not

obtain the basidium-circle along with the spore-circles; focussing down to the

basidium-outline crushed the spore-circles. Therefore I have used average measures.

The spore is very slightly wider tangentially than radially and I have kept these

measures apart as d tan and d rad. Then, the basidia project to various levels;

they are not clearly dimorphic, but those that project most have slightly wider

spore-spacing. I have given the average for M as observed; insufficiency of basidia

may account for the discrepancy with the calculated value of M.

The measures were taken from camera lucida drawings, averaged, and then

used to construct an ideal basidium for the species at a magnification of 10,000.

Angles on such a figure could be measured with sufficient accuracy. The figure

so closely resembles that in Figure 3 that I limit myself to the comparison between

the calculated values and those observed, whether in the original drawings or in

the geometric reconstruction. These are given in Tables 2 and 3.

I do not consider any of the slight discrepancies in Table 3 to conflict with

the general theory. I would emphasize how the variability of the basidium-unit,

constructed from interweaving hyphae, is combined in the hymenium into an

average regime. Thus, some basidia were so compressed as to be oblong ellipses

in end-view, with diamond-shaped sterigmatic ‘square’, yet the average of the

two diagonals of this diamond was that of the square. Spores vary in size and

spacing, yet the averages obtain.

The Boletus-spore

The spores of most boleti are elongate with the smooth and slightly thickened

wall brown, ochraceous or pink. The adaxial face bulges slightly, but the abaxial

is remarkably straight. Other species, placed in Strobilomyces, Boletellus, Heimiella

and Porphyrellus, have a verrucose, echinulate, reticulate or striate ornamentation

which is also coloured. These spores have recently been studied in great detail

by Perreau-Bertrand (1967), who distinguishes five layers in the wall of the orna-

mented spore. For my present purpose I simplify this complexity into the two

layers of customary application, namely the outer colourless and apparently

structureless exospore (which is the ectospore and perispore of Perreau-Bertrand)

and the coloured endospore with its ornamentation, which I regard as an extension

of the endospore into the exospore; the ornaments make the exospore of Perreau-

Bertrand. The point is that, as Perreau-Bertrand discovered, these spores have a

hyaline outer layer, 0.5—1.5u thick, within which the coloured layer forms as the

endospore. Often this outer layer becomes mucilaginous and is scarcely noticeable

in aqueous mounts. The question arises whether the coloured and apparently

external wall of the smooth boletoid spore represents the outer hyaline or the

inner coloured layer of the wall of the ornamented spore. Perreau-Bertrand found

that some smooth boletoid spores had a hyaline mucilage-sheath to suggest that

the coloured wall was endospore. I confirm this observation from many Malayan

Studies in the basidium 175

species and add positive proof that the coloured layer is endospore (see the

following observation iii), Evidently the smooth boletoid spore is an endospore:

that it is peculiar is shown by the following points.

(i). For most boletoid spores d = 0.36 w to 0.58 w, instead of the usual 0.6 w

to 0.65 w. The spore, therefore, is compressed.

(ii). In the case of the ornamented spores of Strobilomyces and other genera,

d = 0.59 w to 0.71 w but, if the inner spore is taken, then d = 0.48 w to 0.60 w

as with the smooth boletoid spore. This means that there are two spacing factors,

o’ for the outer spore of diameter d’ and o for the inner spore of diameter d.

However, some species of Boletellus have spores that are merely finely striate and

such spores are equivalent only to the endospore of Boletellus; and this genus,

incidentally, becomes practically inseparable from Boletus.

(iii). In some pink-spored species, such as B. albo-ater Schw. and B. nanus

Mass., the spore grows to a size that exceeds the final spore by c. lp in length

and breadth. This larger spore is hyaline and, very soon, there forms within it a

coloured, thickening layer which becomes the outside of the mature spore, while

the outer hyaline layer turns mucilaginous and loses its firm limiting membrane,

In these cases the larger transient spore has d = 0.46 w to 0.60 w and the mature

spore has d = 0.37 w to 0.50 w. This proves that the endospore makes the

coloured wall of the boletoid spore and, as corroboration, some of these spores,

are extremely finely striate and could be referred to Boletellus, except for the

spore-colour. I think that this phenomenon of spore-shrinking is widespread among

smooth-spored boleti and that the limits of Boletellus, as of most segregates from

Boletus, are unsatisfactory.

(iv). The ornamented spores of Strobilomyces and Porphyrellus sect. Graciles

have a small adaxial patch without ornamentation; it is absent from the spores of

Boletellus and Heimiella. The adaxial patch forms where the spore-wall contacts

Or approximates with S”’ and, therefore, in these two genera exospore and endo-

spore must coincide more or less in this area. Radial spacing must be the same

for both exospore and endospore, in which respect these spores are less specialised

than those of other boleti.

(v). The elongate spores are strictly parallel with the long axis of the

basidium and, as already noted, the abaxial side is straight. This is evidence of

the compression of the spore-circles which is to be expected from the tubular

hymenium.

In order to discover what may be happening in the development of the

Boletus-spore, I studied in detail several species with a diversity of spores. Table 4

summarises the results. Figures 5-11 show the simplified geometry of the basidia.

The reconstruction of such figures I find to be the only satisfactory way to appre-

ciate the intricacy of the basidium-unit. The species which I studied have the

following particulars.

Strobilomyces velutipes Cke. et Mass., with purple brown, verrucose, ellipsoid

to subglobose spore; adaxial patch present.

Porphyrellus sect. Graciles, an undescribed species with imperfect observations;

spores purple brown, verrucose but with smooth apex and base, and with a narrow

adaxial patch.

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Studies in the basidium

178 Gardens’ Bulletin, Singapore — XXVI (1972)

Boletus emodensis. B. and B. longicollis. Ces. with olive brown, striate-costate,

ellipsoid spores as in Boletellus.

B. retisporus Pat. et Baker with olive brown, reticulate and ellipsoid spore

(Heimiella).

B. albo-ater Schw., B. nanus Mass. and an undescribed species (Boletus 18b

of my notes), with pink, smooth boletoid spores but with evanescent exospore and

very faintly striate endospore.

B. aureomycelinus Pat. et Baker, B. phaeocephalus Pat. et Baker, B. ravenelii

B. et C. and B. umbilicatus Mass. with olive brown smooth ellipsoid to boletoid

spores. [It is to be noted that B. phaeocephalus is the species which I called

B. funerarius Mass. in my earlier paper (Corner 1947); there is a mixture of

type-painting and type-specimen for B. funerarius, the former being B. phaeoce-

phalus}.

Phylloporus cingulatus Corner with smooth, brown ellipsoid spores.

The first point to note from Table 4 is that, except for Strobilomyces, the

observed values of w exceed those calculated from the spore-circles. The circles

are smaller than would be expected. The sterigmata have been approximated by

an adaxial shift, upwards along the basidium-axis, and therefore the spore-circles

have been narrowed and the spores compressed. The compression is most marked

in B. emodensis (Figure 11) and B. phaeocephalus (Figure 5) in which S$” lies

within w.

The point is illustrated by the position of the sterigmatic patch. In all species,

if a sterigmatic patch circle is drawn on S”’ with radius S”’ w, it greatly exceeds

the arc of the sterigma on the basidium-apex. I have shown two sterigmatic patch

circles in the figures, m for o’ spacing and (m) for o spacing. The (m) circle

is the determinant that fits the sterigmatic arc, while the m circle is the unstable,

excessive factor. However, the centres of m and (m) vary from S”’ to M itself.

Thus B. emodensis (Figure 11) shows the greatest compression with the centre of

(m) on S”’ and radius S$” M.

The second point is that in the species with a distinct, if evanescent, exospore

the value of d (inner spore) gives a better approximation to that of M than does d

(outer spore). Reference to the figures or to the table in Appendix 2 will show

that the values of M and w to be expected from d are excessive; this means that o’

spacing is too large and gives place to o spacing. For some physical reason the

exospore is more or less unstable and the spore, constructed on o spacing, is an

endospore. It is not clear, however, that there is any geometrical reason for the

development of ornamentation; presumably it is a physical reaction of the mem-

brane to the instability of o’. Comparison with other species in the groups with

ornamented spores may reveal some common peculiarity.

In all the species with smooth spores the plane of S$” approaches that of w

in evidence of the compression of the spore-circles. However, as noted, B. emodensis

with striate-costate spore agrees in this respect with the apparently smooth-spored

B. phaeocephalu; and, hence, this relation cannot be connected with ornamentation.

Yet B. phaeocephalus is one of the species in which the endospore can be seen

to be exceedingly finely striate.

Studies in the basidium 179

TABLE 4

Spore-measurement in Boletus

Observed values Theoretical values*

( US ) ( p)

d o Ca Bs

S. velutipes 8.00 16.72 | 8.74 | 11.8 | 14.0 | 10.88 | 13.20 | for d’

6.80 16.60 | 9.90 11.76 | 13.95 | ford

ratio to d’ 1 0.50 | 2.09 | £1.09.) £48 | £1.75 | 136 | 1.65

ratio to d & * 1 0.76 LAS | 1.75. |. 206

(number) t .. | (120) | (104) | (26) | (26)} (61) | (1)

B. emodensis i. wel eee 13.91 | 6.63 | 9.54 | 14.46 | 10.3 | 10.1 | ford’

6.08 13.91 | 7.83 9.38 | 11.26 | ford

ratio to d’ ah nob | 0.38 Gob + 1.31 | 199.| 1.42 | 149

ratio to d z. ee 2 0.65 £23.% 3.51 62.96

(number) xe .. | (152) | 102) | G4 | B4 | 44 | (79)

B. longicollis i. .- eetO.33 17.44 | 7FAtAt.Sa7| 14.5 | 8:97 | 11.31 | ford’

8.70 16.77 | 8.07 9.95 | 12.26 | tord

ratio to d’ Pp. i 0.20 0:69 | 1.12 | 1.40 | 0.87 | 1.10

ratio to d a so 0.36 0.93 }. £33 | 1:67

(number) i ~~ | eee) | 2 (24) | (24) | (24) | (100)

B. retisporus 9.18 17.61 | 8.42 | 10.58 | 15.51 | 10.39 | 12.81 | for d’

7.58 17.61 | 10:02 11.99 | 14.33 | ford

ratio to d’ s pe 1 0.36 Oy | tes |, Feo | | 1.40

ratio to d A. “ 1 0.65 1320) DAO OTS

(number) : .» | (164) | 152) :| (8) | G8) | GD | G4

Boletus 18b 5225 11.00 | 5.75 | 9.06 | 10.00 | 6.98 | 8.47 | for d’

3.90 U2. |. tere 8.28 | 9.62 | ford

ratio to d’ 4: a 1 0.48 TAO) P73) RS R33) 1.61

ratio to d ba 2% 1 1.01 15835) 72.32-)) 2.56

(number) fi .. | 364) ? (86) | (86) | (40) | (86)

B. albo-ater 4.77 10.91 | 6.14 | 8.58 | 10.34 | 7.34 | 8.82 | for d’

3.97 10.91 | 6.94 8.11 | 9.47 | ford

ratio to d’ _ I 1 0.618 £29 BO) i BAT 404-54) 1.85

ratio to d a eS te 0.945 1.77 | 2.16 | 2.60

(number) > fer (raar | (lazy) (62)... (62) 7" GE | G1)

B. nanus 5.38 10.96 | 5.58 | 8.35 | 10.98 | 6.81 | 8.32 | for d’

4.03 10.96 | 6.93 8.10 | 9.48 | ford

ratio to d’ 1 0.441 1.04 | 1.55 | 2.04] 1.26 | 1.54

ratio to d ne ne ae 0.924 72 | BOT 142.73

(number) on .. | (250) | (180) | (46) | (46) | (42) | (92)

B. phaeocephalus .. pout Mes 11.65 | 7.10 | 8.68 | 12.20 | 8.37 | 9.87

ratio tod “4 ay 1 0.810 Poo | 1.98 | 20 | 184) 217

(number) + .. | (102) | (160) | (S50) | (50) | (82) | (66)

B. umbilicatus et , at) ad 13.06 | 7.69 | 8.86 | 12.33 | 9.12 | 10.85

ratio to d F out ay 0.720 1.43, | 1,65. |..2:30:1.. 1,70.) 2.02

(number) +e wo |? (oo) |. GO) (9) (9) (6) | (38)

180 Gardens’ Bulletin, Singapore — X XVI (1972)

Table 4 — continued.

Spore-measurement in Boletus.

Observed values Theoretical values*

fx! ei? (py)

d o Sie M w Ww

B. ravenelli - to Ae Gaze 10.22 | 6.01 | 6.81 | 9.96 8.49

ratio to d df OPK, & 0.720 1.43 | 3.62 | . 2.37 2.02

(number) + .. | (132) | (92) | (40) | (40) | (22) | (69)

B. aureomycelinus all ae 10.89 | 6.00 | 7.90 | 10.47 8.69

ratio to d cf. ae all 0.575 1.24 | 1.62 | 2.14 1.78

(number) se .. | (126) | (56) | (28) | (28) | (36) | (64)

P. cingulatus he sw 1 Sz 11.98 | 6.80 | 8.96 | 11.63 9.95

ratio to d a AS ae 0.615 L3i:t i e8 Zoe 1.91

(number) - .. | (188) | (80) | (40) | (40) | (7)! (73)

Ss’ AN <

* calculated from M=S’” — and w=

Sed iS?

d’ outer spore, d inner spore.

(number) refers to the number of observations.

Various errors in observation and construction can be imagined to explain

the differences in positioning of the sterigmatic patch circles, but without under-

standing of the physical forces involved I do not think that they help the solution.

I mention one possibility because it is a step in the adaxial displacement of the

sterigmata. In Figure 5 I have drawn w’ w’ as the false base of the basidium-apex;

it is the base that makes the basidium-apex hemispheric instead of ellipsoid and,

as will be seen, it has the effect of shifting the sterigmata upwards and adaxially

to compress the spore-circles. The sterigmatic patch, nevertheless, seems to fit

not the arc of / HA’ w’ but the upper half of this arc. It appears that the compres-

sion of the cone-effect of the basidium-unit in the concave hymenium acts on the

sterigmatic patches before emergence of the sterigmata. This is a part of the more

general problem of the hymenium-level where some force causes basidia to stop

elongating and to mature; thus cystidia are basidia that have exceeded this level,

often before it has become established in the developing hymenium, and remain

sterile. The hymenium-level is another apparently electrostatic effect of the fruit-

body.

B. phaeocephalus (Figure 5). At first sight this appears to be a normal recon-

struction but the S” circle lies within the w circle, a spacing circle on S”’ with

radius S”’ w greatly exceeds the sterigmatic patch, and the observed spacing,

whereas a spacing circle on M with radius Mw almost fits the tangent to the

spore; and the sterigmatic arc is much shorter than Hw. I detected no exospore

but comparison with Figure 6 shows that the spore of B. phaeocephalus is an

endospore developed from an adaxially displaced sterigmatic patch. This construc-

tion probably represents the normal state for the smooth boletoid spore.

18]

Boletus phaeocephalus, with w’w’ as the false base of the basidium-apex; x 5000.

Studies in the basidium

Figure 5.

182 Gardens’ Bulletin, Singapore — X XVI (1972)

B. albo-ater (Figure 6). This reconstruction shows the excessive o’ spacing

which leads to the disappearance of the exospore and its substitution by the

endospore with (m) spacing. The centre of the (m) circle is not on S’”” but midway

between S’”’ and M (on the inner limit of the endospore). Construction of « “ on

the false base gives the sterigmatic arc with the radius of the half angle contacting

the M ordinate.

B. nanus (Figure 7). In this reconstruction I have drawn a normal spore on

the right with its evanescent exospore and durable endospore. The centre of the

(m) circle almost coincides with M; that of the m circle is slightly abaxial from S$”’,

but for both the radius is now to S” which coincides with w; a spacing circle

with centre on S”’ and radius S”’ w is clearly of no consequence. On the left I

have drawn an imaginary ellipsoid spore (with endospore and _ exospore)

and an imaginary globose spore; the volume of the latter (6.8 x 7.ly) and

that of the ellipsoid endospore (8.0 x 4.6u) are equal to the volume of the normal

endospore. They are positioned according to the spacing factor o’ for the normal

exospore and its theoretical value of w (as from the table in Appendix 2). These

spores are eccentric because S” is 14.47» for o’ and 12.59 for o. The recon-

struction shows how a globose spore of normal spacing relations with the basidium,

such as might have occurred in the ancestor of Boletus, may have been transformed

into the smooth boietoid spore by compression of the spore-circles commensurate

wiih the concave hymenium.

B. retisporus (Figure 8). In this reconstruction (S”’), for the inner spore,

almost coincides with M; they are 0.3 apart. It appears that sterigmatic spacing

is more compressed than spore-spacing; thus M is displaced adaxially towards S”’

and the wider gap which results between exospore and endospore may allow the

development of the reticulate ornamentation all over the spore without a smooth

adaxial patch. In this case the (m) circle contacts the w’ ordinate of the false base.

It is likely that B. longicollis agrees with B. retisporus though the measures of S’”

and M (in Table 4) do not show it. The basidia of B. longicollis collapse very

easily in aqueous mounts and my measures of the spore-circles are probably too

small. (B. longicollis has also been described as B. altissimus Mass. and B. singa-

porensis Pat. et Baker).

Strobilomyces velutipes (Figure 9). The reconstruction is complicated by the

necessity to distinguish the spore-circles S$’, S” and S”’ for the exospore from (S’),

(S”) and (S”’) for the endospore. Two reconstructions are shown. That on the

right fits the smooth boletoid spore and that on the left is the true spore. The

spore on the right has exospore and endospore concentric, but this would allow

ornamentation to be developed all over the spore, as with B. retisporus. The normal

spore has a smooth adaxial patch, where the membrane of the exospore can be

seen to touch the endospore, and the warts are graded in height from the shortest

on the adaxial face to the tallest on the abaxial. The endospore has, therefore,

to be constructed eccentrically to the exospore. The m circle for the normal spore

gives too large a spacing but the (m) circle with centre on S”’ and radius S”’ w

gives the correct spacing for the endospore. This arrangement means compression

of S’ to (S’) without compression of M, and S”’ remains unaffected. The spore

is produced on a wide basidium with much less compression than with the smooth

boletoid spore and is, accordingly, broadly ellipsoid and even subglobose. The

whole basidium-unit is the least modified and, presumably, the most primitive

among boleti.

183

mS" MwS (S)S'

Figure 6. Boletus albo-ater, with exospore and endospore; x 5000.

Studies in the basidium

Gardens’ Bulletin, Singapore — X XVI (1972)

184

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—

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he normal endospore but altere

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185

Studies in the basidium

(5) S'

mim SSM w S"

Figure 8. Boletus (Heimiella) retisporus; x 4500.

Gardens’ Bulletin, Singapore -— X XVI (1972)

186

-----------------

ee ee ee core

Figure 9. Strobilomyces velutipes with a normal spore on the left and an abnormal spore,

constructed as a smooth spore, on the right; x 4800.

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™

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aS)

Gardens’ Bulletin, Singapore — X XVI (1972)

188

Boletus emodensis; x 5000.

Figure 11.

Studies in the basidium 189

Having no detailed analysis of the curvature of the basidium-apex in this

species, the reconstruction has been made on a hemispheric apex. It is probably

a reconstruction on the false base with the effect that the radius which bisects «

meets the sterigmatic arc where intersected by the M ordinate.

Porphyrellus sect. Graciles (Figure 10). Having no fresh material to study

in this interesting case allied with Strobilomyces, I used alcohol-formalin material

of an undescribed Malayan species. This did not give reliable measures for the

spore-circles. I reconstructed the basidium-unit from measures of d and w and

the ratio w = 1.655d for d’ tan, which gives o’ = 0.5. The measures were as

follows: —

exospore 17.36 X 8.41,» (radial), X 9.16 (tangential);

endospore 17.36 X 7.26, with truncate conical warts 0.7— 1.24 high (mean

0.954) on the sides and abaxial face, 0.1-0.3 on the adaxial side; (average of 30

spores);

basidium 32 X 15.16u, with the sterigmatic circle 11.84 wide (average of 50

basidia and 40 sterigmatic circles).

The reconstruction on a hemispheric apex for the basidium is similar to that

of Strobilomyces but the spore is compressed and elongate through the adaxial

shift of M towards S’”. The point is shown by the value of S’ which is 27» in the

reconstruction though, theoretically, it should be 28.6u. The adaxial shift of M is

the principal change leading to the smooth boletoid spore. Thus, this section of

Porphyrellus appears to be half-way between the state of the basidium-unit in

Strobilomyces and that in Boletus with smooth spore.

B. emodensis (Figure 11). The high compression of these spores is shown by

the upward and adaxial displacement of the sterigmata on the wide basidia.

Though I have constructed the basidium-apex as a hemisphere, the false base

would need to be lifted even higher, as shown by the dotted line, to fit the

sterigmatic patches; the dotted line gives w= 10.lu, which is the usual width

of basidia in smooth spored boleti. Then, as further evidence of compression in

this figure, the real value of w slightly exceeds that of S”; the m circle has centre

M and radius Mw; the (m) circle has centre S”’ and radius S’” M.

B. longicollis belongs in this same alliance of Boletellus with striate-costate

spores. So, apparently, does B. obscurecoccineus Hoehn. with faintly striate spores

equivalent to the endospore of B. emodensis. It would seem that in this alliance

the series can be traced from the broadly ellipsoid spore to the smooth boletoid.

Unfortunately other species in Malaya have n = 2 or 3, and comparable results

could not be obtained.

Phylloporus

This genus is close to Boletus in its wide sense and differs in the lamellate

hymenium. Among fifteen Malayan species for which I have accurate measures

(Corner 1970), thirteen have d = 0.42w to 0.56w, but in P. hborneensis d = 0.58w

and in P. coccineus d = 0.71w. Evidently there is the tendency to the compression

of the spore-circles in this genus without the development of the fully tubular

hymenium. I worked out the details for P. cingulatus (Table 4); they showed the

boletoid features of close agreement between the values of S” and w and of the

location of the M ordinate through the m circle on the false base.

Gardens’ Bulletin, Singapore — X XVI (1972)

190

— ee Se Se Oe Ee = Te

Figure 12. Marasmius nigripes; x 5000-

Studies in the basidium 191

If, as I consider, the poroid Boletus has been derived from a lamellate

agaric of Paxillus-Phylloporus affinity, then it seems that the characteristic of the

Boletus-spore was developed in this ancestor, but both Paxillus and Phylloporus

have smooth spores. This conflicts with the idea, put forward in previous pages,

that the primitive Boletus had an ornamented endospore. Therefore the monotypic

South American Phylloboletellus Singer stands out because it has the phylloporoid

hymenium and the striate-costate spore of Boletellus (Singer 1964). According to

Singer’s measurements, in Phylloboletellus chloephorus d = 0.78w and this implies

both o’ and o spacing. Certainly it suggests a primitive boletoid fungus, though

apparently without adaxial patch. This is supplied by Gloeocantharellus with

ornamented spores and apparently no exospore. For three of its five species

d= 0.59 to 0.65w (Corner 1969). There appears to be a single spacing factor

with external ornamentation, as in Russula and without any of the elaboration

of the boletoid spore.

Marasmius nigripes Schw.

The remarkable spores of this fungus illustrate the use of geometrical analysis

(Figure 12). In adaxial and abaxial view the spores are cruciform; in side-view

they are trigonous and in end-view crescentic. They develop from the spore-

rudiment into a shortly clavate form (6 X 3), typical of many species of

Marasmius; this form is shown in dotted outline in the figure. Then shoulders

develop to extend laterally and abaxially while the apex lengthens into the third

arm and the spore-body swells to full size. There are no radial arms against the

boundaries of S’ and S”’, Figure 12 is based on averages from ten basidia in

side-view and ten in end-view; the basidium apex is drawn as semicircular.

On this reconstruction I then drew (i) the radial tangents to the spore-body

and spore-arm; (ii) the sterigmatic patch circle (centre S”’, radius S’” w); (iii) a

spore-body circle b inscribed on S” between S’”” and S’, and found to my surprise

that it fitted the tangent to the spore-arm; (iv) a spore-arm circle v (centre S”,

radius S” to the apex of the arm), and found to my surprise that it contacted

adjacent spore-bodies and almost reached the long axis of the basidium; (v) an

outer spore-circle a (centre A, circumscribing the spore-arms); (vi) an outermost

circle u circumscribing the v circles, as the outermost limit of the basidium-unit.

This figure leads to the following conclusions: —

(i) the m circle may be the prime spacing circle for the spores; that it does

not contact the tangent in the drawing may be due to the difficulty in locating

the exact tangent in this region where the curvature of the spore is changing.

(ii) the m circle gives © = 0.42 or w = 1.5ld, which corresponds with

w = 1.52d for average measures of spores (9-12 x 8-llu) and basidia (25-32 xX

8-10,).

(iii) the 5 circle, contacting the spore-arm tangent, gives « = 0.44 or w

== 1.55d.

(iv) the actual spore-body (with radius measured along the dotted line joining

adjacent spore-centres in Figure 12) gives c = 0.61 or w = 1.84d.

(v) the spore starts with theoretical spacing of o for the mature basidium;

it then reduces this to o’, grows radially to the allowance of oc’, but this extension

occurs only about the mid-length of the spore; the spore-body is itself compressed

tangentially to a value between o’ and co.

192 Gardens’ Bulletin, Singapore — X XVI (1972)

(vi) in final form, as shown by the v circles, the spore comes to fulfil by

some physical means the whole spore-sector.

The compression of the spore-body, mentioned in (v), may bear on the nature

of the long, narrowly clavate spores of many species of Marasmius, as another

instance of spore-compression, similar to that of Boletus. I have seen a fine mucilage

sheath round many Marasniius spores. On germination of the spore of M. nigripes,

it is only the spore-body that swells.

References

Buller, A.H.R. (1922) Researches on fungi. Vol. Il. Longmans, Green and Co.;

London.

Buller, A.H.R. (1924) Researches on fungi. Vol. II.

Buller, A.H.R. (1931) Researches on fungi. Vol. IV.

Corner, E.J.H. (1936) Hygrophorus with dimorphous spores. Trans. Brit. mycol.

Soc. 20, 157-184.

Corner, E.J.H. (1947) Variation in the size and shape of spores, basidia and

cystidia in Basidiomycetes, New. Phytol. 46, 195-228.

Corner, E.J.H. (1948) Studies in the basidium I. The ampoule effect. New Phytol.

47, 22-51.

Corner, E.J.H. (1950) Monograph of Clavaria and allied genera. Ann. Bot. Mem. 1.

Corner, E.J.H. (1954) Further descriptions of luminous agarics. Trans. Brit. mycol.

Soc. 37, 256-271.

Corner, E.J.H. (1964) The life of plants. Weidenfeld and Nicolson; London.

Corner, E.J.H. (1966) A monograph of cantharelloid fungi. Ann. Bot. Mem. 2.

Corner, E.J.H. (1968) A monograph of Thelephora (Basidiomycetes). Nova

Hedwigia Beih. 27, 1-110.

Corner, E.J.H. (1969) Gloeocantharellus. Nova Hedwigia 18, 793-805.

Corner, E.J.H. (1970) Phylloporus and Paxillus in Malaya and Borneo. Nova

Hedwigia (in press).

Corner, E.J.H. (1972) Boletus in Malaysia. Government Printer. Singapore.

Malencon, G. (1958) Le dévelopment des spores chez les Phylactéries. Bull. Soc.

mycol, Fr. 74, 423-435.

Perreau-Bertrand, J (1964) Recherches sur la différentiation et la structure de la

paroi sporale chez les Homobasidiomycétes a spores ornées, Annls Sci. nat.

Bot. ser. 12, 8, 639-745.

Singer, R. (1964) Phylloboletellus. Nova Hedwigia 7, 129.

Studies in the basidium 193

Appendix 1

Summary of the relations between the spores and the basidium (n = 4) d = spore-width;

m = diameter of the sterigmatic patch circle; w = basidium-width; M = diameter of the

sterigmatic circle; S’, S”, S”’” = diameters of the outer, middle and inner spore-circles

respectively.

eo = the sterigmatic patch angle; § = half the spore-subtending angle; @ = the spore angle;

@ =the spore-spacing factor.

Text reference

a 0 Sr a a a ae (1)

fain’ d

S*=d/2(1 Se ame aN Ra ss fw badip dnd w ves db uvnsde sus auc huetbeacs¥amder GnSevkde (2), (6)

sin §

d (1+sin 6)

f= 5" 4+d=d f2ai +o)t+d= ae rise =a tiv + GOSEO) = 8 SCORE hoc 5 ccc escsiewcne (7)

sin

d(1 — sin 6)

8” =S" —~d=d /2(1+ 0) —d = ————-= d (cosec § — 1) er eee Ce. Sore eee (8)

sin §

Ss +e 2) and J —S”= 2d

a Bed Lhd SV Ae

—=1+sin§ = 1 — sing a eae —- =w

Ss” Fe So

wcos «

M= = =e | Se a ART eee anrrren. Aa a remnren (14)

2 1 + oor 7

M = dcotscosd eS acs sea Mt AOR aBES cn alth veecapacertncadons ee. ie (22)

1 + sin §

Me OF ot (oes § 07) Sf J 21 SY OOTY oooh eens seceshsnagah en denecens (9)

A i cosec 6

=—— -l= aE en Ane 5 eee BOR ii hioe ek Aton sinc dratindawahedaths cxwsisesa (10)

df 2 2.

1 + cos«

I RE IE) AEE RE RO. ARE. A ORE See eee, PE (15)

) Fae oe — cos « ;

ee eI ON Gal Savin pais as eied bah vi ekabeavadehccvcsedet ninteVeeseadicceungescoemerencscses (11)

Nie he

ee i eae TT ee ee, eee ee eee (12) (13)

d 1 S’—M 1 — cose

SAT cecceeccc ese ec ete netec ene ea ene nateneneneeeens (17)

5S” (2( +0) S” sn@ 1+cos«

i, ag 1a m 1 — sin 5 A fa

a ia cdl sts stdensstcwsssstescnrascacececcanecsteccsseccesescaceante (15)

, tm. J2+3n8 5

sin @ = se = cosec § — cot §cos § hcp ood cE ek feuedi onpine (18), (23)

d S” sin § 1 + sin§

5S’ —M

Gardens’ Bulletin, Singapore — X XVI (1972)

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GARDENS’ BULLETI

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. Vol. XXVI, Part I 15th September, 1973

CONTENTS

; PAGE

4 Dctrmons Ee: EEeuCRCY and Habitat of Tree Species in the Rain Forest

a Be of Ulu Kelantan - - - . - 195-210

_ Horrrum R. E.: A New Bamboo from Mt. Kinabalu - - - - 211

Marie A. Martin: Notes on the Vegetation of _the Cardamon Mountains,

a _ Cambodia - : : - - 5 - 213-222

_ Hsvan Ken & Heastert E. AS The x ylopia malayana fruit: significance of its

dehiscence - 223-225

¢ _Sitior0 Kurata: Nepenthes from Borneo, Singapore and Sumatra - . 227-232

a HsuAN KENG: Annotated list of seed plants of Singapore (I) - - - 233-237

* ‘Draxsriet De? Korthalsia hispida Becc. in Malaya - - = - 239-244

ay R so A. N. and Lee War CuIN: Pollen Vogue and Gomalnation in Some

ee Orchid Hybrids - - - - : 245-257

“Nayar M. P.: A Review of the Genus Creaghiella Stapf. (Melastomataceae) - 259-261

) CK H TIONG-KHENG: New Plant Disease Records for Sarawak for 1970 and 1971 263-268

KOCHUMMEN K, M. & Wuitmore T. C.: Notes on the Systematy of Malayan

oP . be Phanerogams XVIII-XXxII - - 269-287

ANNE : JoHNSON: A survey of the occurence of epiterranean soil algae in Singa-

f E: pore Island - _- - - - - - - . 289-302

LiL J.: Pollination in Dioecious Figs: Pollination of Ficus fitulosa by

i i \y Ceratosolon: hewitti - - - - - - - : 303-311

To be purchased at the Botanic Gardens, Singapore

Price: S $18

Published by Authority

é ‘ : : Printed by the Singapore National Printers (Pte) Ltd

Sr 1973

ae een

Frequency and Habitat of Tree Species

in the Rain Forest of Ulu Kelantan

T. C. WHITMORE

Forest Research Institute, Kepong*

Abstract

In 1967 a survey was made as part of the Forest Resource Reconnaissance Survey of

Ulu Kelantan district. 26, 628 trees were enumerated > 4ft girth on sample strips 1 chain

wide, in primary rain forest mostly below 2800ft elevation. For this analysis the survey

area of 1672 acres has been divided into 4 classes on geology and altitude, comprising

sedimentary and granitic rocks and below or above 750ft. The occurrence of different tree

species, genera and families in these 4 classes is listed in a series of Tables. No such

extensive analysis has previously been made. There are many groups which are markedly

more common in one or other of the 4 habitat classes and several groups have a distri-

bution in Ulu Kelantan which counters subjective preconceptions. It is suggested that at

any altitude species’ distribution is likely to be more closely correlated with relief and

soil than with geology.

INTRODUCTION

Our knowledge of the habitat preferences of Rain forest trees in southeast

Asia is very largely based on the subjective impressions of forest botanists not

backed up by measurements of tree frequency in particular habitats or precise

records of the nature of the habitats themselves.

An enumeration of all trees over and including 4 ft girth on a small sample

of the Lowland Rain forest of Ulu Kelantan district, northeast Malaya has been

the stimulus to evaluate the relative frequency of the various tree families, genera

and species occuring there, and to analyse their distribution between the two

major kinds of rock in the district, and at low or higher elevations. This is of

course, background information of fundamental importance when attempting

to summarise our knowledge of particular species for such publications as the

new Tree Flora of Malaya (Malayan Forest Records 26, Whitmore 1972, 1973 a).

Species with horticultural and some with known pharmacological potential

enumerated in this survey have already been discussed in Whitmore (1971).

It is well known in Malaya, as a matter of observation, that different species

have different altitude preferences, and there is a growing awareness that the

rock type on which a forest occurs, to some extent at least, appears to determine

its species composition, in a manner tentatively analysed below where we attempt

to investigate a little further these suspected discriminants to species distribution.

THE FOREST SURVEY

The forest survey was conducted between June and November 1967 by

Forest Resource Reconnaissance Survey teams under the able direction of Mr.

N. Cerra, working from Kepong as an American Peace Corps volunteer, as part

of an operation which has now covered the lowland forest of the whole country.

* Present address: Mead Hall, Saffron Walden. Essex CB10 IUX, England.

195

196 Gardens’ Bulletin — XXVI (1973)

The forest was divided into a series of ‘phototypes’ from the study of aerial

photographs. Teams then went into Ulu Kelantan and systematically sampled

these types throughout the district, at a low intensity. The sample lines were 1 chain

wide and up to several miles long, but divided into 10 chain lengths, so individual

plots of 10 sq. ch. (1 acre) are the basic unit. The lines were cut across the ‘grain’

of the country so as to sample ridge tops, hillsides and valleys; altitude was

recorded; the precise position of each line was recorded on a set of 1 inch maps.

Access was mainly by river, and from any one camp a series of lines was

surveyed; the inset of Fig 1 shows the location of each cluster of survey lines.

The total sample area was 1,672 acres; 26,628 trees were enumerated and named

according to the preferred vernacular list in Kochummen & Wyatt-Smith (1964).

The upper limit of most lines was below 2,800 ft, a few went up as high as 3,800 ft.

Thus the whole survey is restricted to the Lowland Tropical Rain forest formation

(Symington 1943, Burnham & Whitmore 1969).

Because this was a reconnaissance survey no attempt was made to take a

rigorously, objectively selected sample. It was adequate to cover different photo-

types throughout the whole district. It follows that it is unwise to multiply up

from the results of this sample to find tree numbers over the whole of Ulu

Kelantan. The results are, however, likely to be indicative for this part of Malaya.

No pretence is made that the present analysis indicates frequency and habitat

everywhere in Malaya; indeed, as pointed out below, there are, in Ulu Kelantan,

several marked departures from what are suspected to be the preferences of species

and groups elsewhere in Malaya.

The forest data were divided into stands for each of which a table was

prepared stating, inter alia, the number of stems of each vernacular-named group

occurring and the total basal area of all the trees enumerated. The stand tables

have been the basis for the present analysis. Each stand is either one whole

survey line or part of one. Each one contains homogeneous forest of one of the

forest types recognised by the Forest Resource Reconnaissance Survey.

For the present analysis the stands were divided into two groups, those

below 750 ft elevation and those above, to separate lowland dipterocarp forest

from hill and upper dipterocarp forest in the sense of Symington (1943).*

Regrettably, nearly all the stands transgress the 750 ft contour, and have had to

be included on the side where they mostly lie. This can only blur the altitudinal

preferences of species and groups and weaken the clarity of the analyses presented

below.

ULu KELANTAN

Ulu Kelantan district was in 1967 mostly still under primary rain forest.

The major areas of settled agriculture are shown in the inset to Fig 1. Logging

had then not long begun, though by early 1972 much of the district had been

let out on logging concessions. The total area of this administrative district is

4,817 sq. mi. and about 80 per cent was then still forested. Within the forested

part of Ulu Kelantan there is little flat land except in the Lebir/Aring basin in

the east; mostly the rivers run through narrow, steep-sided valleys. The main

mountains lie to the west and east (Fig 1) and this is also, roughly, the distribu-

tion of granite, of Triassic age, which forms the Main and east coast ranges

* Symington drew the boundary at 1,000 ft. I have divided at 750 ft on the advice of

Mr. P. F. Burgess who considers the latter a more realistic elevation especially, as here, in

hilly country.

Tree Species in the Rain Forest of Ulu Kelantan 197

5000 ft

1200 ft

600 ft

Fig. 1. Ulu Kelantan District, location and relief. Taman Negara also shown, S. and E. of

Ulu Kelantan and partly over-lapping. Inset, location of sample sites, rock types (granite

dark, sedimentary rocks pale) and approximate main area of cultivation (hatched). Geology

from Alexander 1963. Cultivation limits from Wyatt-Smith (1964).

198 Gardens’ Bulletin — XXVI (1973)

of the Peninsula respectively. The centre of the district is mainly of Permian and

Triassic sedimentary rocks, which are described further below. The geology is not

known in detail, but the granite/sedimentary rock boundary is fairly accurately

known; only eight of the 222 survey stands lay so close to the boundary that

their geology is doubtful. The stands were classified as lying on either granite

or sedimentary rock and the analysis of species frequency was made for four

groups, low granite (below 750 ft), high granite (over 750 ft), low sedimentary

and high sedimentary; in these four classes 113.8, 448.8, 716.4, 393.0 acres had

been sampled respectively; it can be seen that low granite was distinctly under-

sampled.

SPECIES AND FAMILY FREQUENCY

So few have been the extensive, quantitative analyses of composition and

habitat in Malayan Lowland Rain forest that, when quoting the relative frequency

of species, one is thrown back on observations scattered through the pioneer work

of Foxworthy (1927).

Table 1 shows the 21 species in order of decreasing frequency which have

10 or more stems per 100 acres in this survey. The first column gives the average

density, the next four columns give, for the ten commonest species, the density

in the four habitats. A major weakness in this table is that many species were

not individually recognised in the forest. Dipterocarpaceae dominates the table,

partly because it is a very common family but partly also because every species

has a vernacular name. Burseraceae and Sapotaceae are both very common in

Malayan Lowland Rain forest but their species were not distinguished from each

other. Nevertheless, the top third of the table is probably roughly correct, only

lower down would one expect other species, especially of Burseraceae and

Sapotaceae. Note that the order of the first six species does not vary between the

habitats, though the order does vary lower down. The important exception to this

generalisation is Shorea curtisii* (seraya) which is almost confined to hill ridges

where it sometimes forms almost pure stands; this species is so common in hill

forest that it still easily comes top of the list, despite its low frequency in lowland

forest. The next two species (S. parvifolia, S. leprosula) are both light red merantis

which are fast growers and light demanders and come up gregariously in clearings.

They are amongst the commonest trees everywhere in Malaya.

In Table 2 the 42 families encountered in the survey are arranged in order of

decreasing frequency. Again, the first column gives the average density (in stems

per hundred acres rounded off) and the next four columns relate to the four

separate habitats.

One sees at once why these forests are known as dipterocarp rain forests, the

family Dipterocarpaceae is four times as abundant, amongst the trees > 4 ft girth,

than Leguminosae, which comes second, and this is true of all four habitats. As

in the species table the order of frequency of the families differs between habitats

low down the table, though the first four families are in the same order in every

one. Of the two biggest tree families in Malaya Euphorbiaceae comes surprisingly

high in the list compared to other surveys which have been made (Whitmore,

1973 b); Rubiaceae, which is actually slightly larger than Euphorbiaceae in

Malaya in number of species, comes much lower, this is because very few of them

reach 4 ft girth. Leguminosae always come high in a survey of this nature, but

* Species names throughout follows the new Tree Flora of Malaya (Whitmore 1972

and 1973a) or, for families not yet revised, Kochummen & Wyatt-Smith (1964).

Tree Species in the Rain Forest of Ulu Kelantan 199

Table 1. The commonest of the species identified in the forest, those with on average more

than ten stems per hundred acres

granite sedimentary

ee” Now" high” low high

1. Shorea curtisii(Dipterocarp.) .. xr 125 21 126 20 345

2. Shorea parviflora (Dipterocarp.) . . m 108 91 93 127 95

3. Shorea leprosula (Dipterocarp.) . . a} 80 a7 88 i 82

4. Koompassia malaccensis (Legum.) he 51 36 39 66 45

5. Shorea dvatis (Dipterocarp.) * e 40 26 33 51 34

6. Dipterocarpus cornutus (Dipterocarp.) 1 35 40 13 56 24

7. Elateriospermum tapos (Euphorbi.) =e 33 16 45 20 49

8. Dryobalanops oblongifolia (Dipterocarp.) .-. 31 17 3 66 6

(Shorea multiflora (Dipterocarp.) Ae 23 16 11 30 25

a ie palembanica (Legum.) oN ye 23 18 26 23 22

11. Shorea platyclados (Dipterocarp.) A 21 — — — —

( Koompassia excelsa (Legum.) .. yg 18 —_ _ — _-

, | Cynometra malaccensis (Legum.) a 18 — — — —

14. Dipterocarpus crinitus (Dipterocarp.) $i i5 — — = —

15. Shorea bracteolata (Dipterocarp.) o 14 — — — —

16. Shorea guiso (Dipterocarp.) at iu 13 — — — —

17. Parashorea lucida (Dipterocarp.) cs 12 — — — —-

18. Pometia pinnata (Sapind.) . on 11 — -— “= os

Mesua ferrea (Guttif.) ter ~¢: 10 “— — — —

19.< Endospermum malaccense (Euphorbi.) ne 10 — = — —

| Ochanostachys amentacea (Olac.) dé 10 a a a5 =

Family name abbreviated in brackets; stems per hundred acres rounded off to nearest whole

number.

Myrtaceae and Lauraceae are usually lower and Burseraceae higher. The complete

absence of Flacourtiaceae, with several big trees with well known vernacular

names, is curious. Alangiaceae, Proteaceae, Symplocaceae and ‘Theaceae

would also all have been expected in Ulu Kelantan. It is a well known weakness

of Forest Dept. survey teams in Malaya to assign an unknown tree to either

medang or kelat, and it is likely that trees of these missing families, and one must

assume Flacourtiaceae too, have been mistakenly included in Lauraceae or

Myrtaceae and thus contribute to the unexpectedly high figures for these two

tamilies.

200 Gardens’ Bulletin — X XVI (1973)

Basal Area and Tree Numbers per Acre

Sedimentary rock has a higher basal area of trees > 4 ft girth per acre than

granite, due to the much lower basal area per acre on granite over 750 ft than

elsewhere (Table 3). In Ulu Kelantan (as shown below, Table 9) the important

hill forest timber species Shorea curtisii is relatively rare on granite; doubtless

this contributes to the low basal area.

It can be seen from Table 3 that stems per acre are more numerous above

750 ft than below. These figures reflect a bias in the sample. In the hills big trees

tend to be commonest on ridges and sparse on hill sides and in valleys and it can

clearly be seen on Table 4 that survey lines above 750 ft have tended to follow

ridges rather than to cut across the lie of the country, which becomes increasingly

difficult with increasing elevation. The slightly larger number of stems per acre on

low sedimentary rocks than on low granite, and other slight differences in Table

3 may not be statistically significant. An analysis of variance would be required

to demonstrate that they are not due merely to chance variation in the data, and

this has not been attempted because of the difficulties involved in extracting the

figures from the survey summaries.

HABITAT PREFERENCES OF SPECIES AND GROUPS

Limitations to the Analysis

The main analysis with which we are here concerned is that which attempts

to elucidate how far individual species are confined to particular habitats, in this

case defined on altitude and underlying rock.

Two main weaknesses of the analysis have already been pointed out; firstly,

that the forest data have been arranged in stands many of which cut across the

boundary at 750 ft between lowland and hill forest; secondly, that the forest staff

do not recognise many individual species outside Dipterocarpaceae but do

recognise genera (e.g. petai: Parkia; sepetir: Sindora, etc.) generic groups (e.g.

mempening: Lithocarpus, Quercus) or whole families (e.g. nyatoh: Sapotaceae).

The result of this second weakness is, as of the first, an obfuscation of revealed

habitat preferences. Individual species (or indeed groups within them) must be

expected to show the greatest specificity for habitat, and to lump species together

can only lead to blurring.

A third and serious source of weakness is that the rocks have only been

divided into the two great categories granite and sedimentary. There are consider-

able variations in the granites of Malaya, often over very short distances, in

texture and composition. No attempt has ever been made to map these. The

sedimentary rocks are even more varied and include, in Ulu Kelantan, the pre-

Carboniferous Teku schists in the north, which are the only large area of schist

in the country, plus large areas of predominantly argillaceous Permian sediments

and of predominantly arenaceous Triassic sediments. The various sedimentary

rocks are usually interbedded and have in places been metamorphosed, especially

along the boundaries with the granite. In places they are variously calcareous. No

record was made of the rock type of the survey lines, indeed it is unlikely to

have been homogeneous along their length, and the published geological records

are not sufficiently detailed for high precision. It is for these reasons that the

analysis is restricted to granite versus sedimentary rocks, as the boundary

between these two major types is believed to be well and accurately known.

Tree Species in the Rain Forest of Ulu Kelantan

Table 2. Frequency of Families

Leguminosae

Myrtaceae

Lauraceae

Anacardiaceae

Burseraceae

Euphorbiaceae

Olacaceae

Fagaceae

Sterculiaceae

Guttiferae

Moraceae

Sapotaceae

Sapindaceae

Se A Se a ee

Lee ce I ie eee cee ee

i en

Annonaceae

ys Dilleniaceae

" UMyristicaceae

18. Polygalaceae

19. Tiliaceae

20. Apocynaceae

21. Lecythidaceae

22. Bombacaceae

23. Thymelaeaceae

24. Rutaceae

25. Celastraceae

26. Combretaceae

27. Ebenaceae

28. Rhizophoraceae

29. Hypericaceae

30. Meliaceae

31. Loganiaceae

32. Rubiaceae

33. Simaroubaceae

34. Juglandaceae

35. Malvaceae

36. Ulmaceae

37. Melastomataceae

38. Oxalidaceae

39. Coniferae

40. Magnoliaceae

Theaceae

sat { Linaceae

Dipterocarpaceae . .

average

634

161

122

37.8

wid

122

8.0

1.0

5.0

4.1

3.1

3.0

2.8

2.0

1.9

1.7

1.6

1.5

1.3

0.6

0.5

0.2

0.9

.04

.02

.0006

201

granite sedimentary

low high low high

439 602 587 815

139 153 182 140

73 122 124 i33

57 94 4 68

25 69 64 109

48 90 67 66

50 67 39 71

47 35 46 36

40 60 30 31

41 69 37 37

21 33 38 47

34 43 29 30

27 40 33 22

24 24 a | 20

16 29 14 21

11 14 iF 8.9

23 16 14 9

‘| 8 16 12

13 8 12 10

6.1 9.8 Ts 7.9

11 6.3 8.8 5.9

12 i 53 3.1

1.8 6.0 6.2 2.8

3.3 1.8 6.8 is

0.9 1.3 eres 4.6

0.9 3 4.2 1.8

1.8 2.9 2.0 4.3

1.8 1.3 2.5 2.3

— 20 1.9 13

3.6 2.9 1.6 =

1.8 0.2 1.8 0.8

1.8 2.6 0.6 1.8

—- 11 1.4 1.8

-— a2 -— —

4.9 0.5 0.1 -—

— 0.5 0.1 —

—- 0.4 0.1 —

0.9 0.2 1.3 0.8

= pe _ 1.8

_- —- 3.9 —

= -—— 0.1 —

—— — 0.1 —

In Tables 2 and 5-14 the figures given are for stems per 100 acres.

202 -Gardens’ Bulletin — XXVI (1973)

Table 3. Basal area and tree numbers per acre

granite sedimentary

low high low high

Basal area (sq ft) i <2 A Fe iP 47.9 52.9 56.7

average each rock .. af ae - 49.5 54.3

overall average 2M bd B - Se 52.7

Stems per acre. . a se v So bee 17.3 14.9 17.3

average each rock .. at 4 ie 14.6 15.8

overall average 7 bu ae bi 15.9

Table 4. Representation (as percentage of total acreage) of different topographic sites

granite sedimentary

ao ST SO ge LS Fe —

Both rocks

we, | 0- 500- 1,500- 2,500- 0- 500- 1,500- 2,500-

Coral’ “aititude ihye Meah son 1300. 7.500 4.00) MO” sah Sar

i. they &%. nie. & 8 eet | Wee ee gine J

67.\.. hillside fcc. top4PZedeoBOee. Te avi TneomtpZQa 1ftisn ollidestiiateeeeee

22) nadkdgecepyidel8ad wideaigine ask. 23 24. te Fe ee

area sampled

(acres)... dx. -—i.. 41.8-u2518+ 91Sruns48dcanc a@ienaeeda ie 0

based on an unpublished analysis by P.F. Burgess of 1087.1 out of the total 1,672 acres.

A fourth potential source of weakness is that valleys, hillsides and ridge

crests might have been sampled in different proportions in the four classes, and if

species have topographic preferences a bias would emerge in the results. Examin-

ation of an unpublished analysis by Mr P. F. Burgess of 1087.1 acres (out of the

total 1.672 acres), see Table 4, shows that, fortunately, the proportion of the

topographic classes does not vary largely either with rock type or altitude for

the two altitude groups 0-750 ft and over 750 ft considered here. But, because

there are differences of the order of 10 percent, comments on the present analysis

are restricted to major differences in species’ density. It has been noted already

that the area of low granite sampled was disproportionately low, and this is borne

in mind in the comments which follow.

All these limitations must be borne in mind in the analysis which follows,

which is presented as a series of tables showing stems per 100 acres. Individual

species are shown as far as they were recognised (the Dipterocarpaceae dominat-

ing the lists), then genera (either straight from the survey data or by adding

together species’ frequencies), then families. It is interesting to see how far genera

and families do show habitat preferences, and whether these support the pre-

conceptions forest botanists have about such things, though in general these big

groups cannot be expected to show much uniformity in habitat.

Tree Species in the Rain Forest of Ulu Kelantan 203

It must also be remembered that only trees 5 4 ft. girth were sampled.

The effect of this may be illustrated by considerating two genera of the Sapindaceae.

Xerospermum (gigi buntal) is actually far commoner in the forest than Nephelium

lappaceum (rambutan hutan), but quite the reverse is shown in the result of this

survey, because its trees rarely attain 4 ft girth and so only a few exceptionally

large individuals have been counted, whereas rambutan hutan grows to a larger

tree and so a larger fraction of the total population of the species has been

included.

In general little is known about species or group preferences for sedimentary

or igneous rocks in Malaya as this is the first analysis made of a large body of

data. Comments on the tables are therefore restricted to particular points which

confirm or conflict with our existing subjective impressions.

In fact, rock type itself probably exerts a less direct controlling influence on the

distribution and frequency of plant species than the nature of the relief and

of the soil (which is partly dependent on relief) which themselves are not directly

related to rock type as expressed in this analysis.

In the hot, wet climate of Malaya all soils are very strongly leached and

consequently poor in plant nutrients; physical soil properties are probably usually

more important to plant life than chemical ones, notably the depth to which the

rock has weathered and the particle sizes formed by weathering. Granites tend

to weather very deeply and to form well-structured soils, such as are seen along

road cuttings in the Main Range, though quartz-rich areas may yield very

shallow and coarse soils. Sedimentary rocks tend to weather less deeply, especially

where metamorphosed; shale-derived soils are much more finely textured than

sandstone-derived soils. Further, the relief of granite country tends to be less

rugged than of sedimentary where precipitately sided knife-edge crests often

replace broad, rounded ridges. The task of collecting detailed data on topography

and soils, such as would be needed for a fully analysis, would be formidable

in rain forest country, and has never been attempted over more than a few acres.

On small areas the problem of correlating plant distribution with habitat is con-

founded, in these excessively species-rich forests, by the probability that there is

not enough space in a small area of any particular habitat for all the species to

grow which are ecologically suited.

The Analysis

The first important conclusion results from a glance at Tables 5-12. It can

be seen at once that nearly all species, except some of the rarer ones, occur in

all four habitats, but that the frequency of occurrence varies in nearly all cases.

This shows how misleading qualitative impressions of species’ distribution can be,

and how important it is that ecological analyses of the rain forest are not based

solely on presence or absence.

Groups showing a marked preference for granite.

(i) Below 750 ft, Table 5:

Hibiscus floccosus (kangsar) and Pterygota horsfieldii (kasah) are valley species

not known previously to prefer granite.

The genus Hopea is usually considered commonest at high elevations; the

preference here for lowland forest is unexpected.

204 Gardens’ Bulletin — X XVI (1973)

Table 5. Groups showing a marked preference for granite and below 750 ft elevation

granite sedimentary

@) low high low high

a) species

Dipterocarp. Shorea assamica .. uh JO9 if 0.2 3.3 =

S. kunstleri i] 0; , fiom 8.7 re» =

Euphorbi. Endospermum malaccense Pe im eee 12 8.7 7.6

Maly. Hibiscus floccosus . . af s: 4.9 0.5 0.1 --

Mor. Artocarpus rigidus. . fd og 43 8.0 7.1 5.6

A. scortechinii (elasticus) dag ue 7.9 5.6 2.4 1.5

Sterculi. Pterygota horsfieldii fi. s 5.3 2.9 0.4 0.3

(b) genera

Dipterocarp. Hopea . . it a - 1G 3A 27 22 18

Legum. Parkia . . au aft pea sblg 11 7.3 6.1

Myristicaceae B, 4 a iO 223 16.1 14.4 9.4

All the species of the forest survey are included in the tables 5-12 except for a few obviously

mistaken identifications.

(ii) Above 750 ft, Table 6:

Shorea platyclados (meranti bukit) is the highest growing Malayan Shorea

and a very important timber tree of upper dipterocarp forest from 2,500—4,000 it;

it is locally gregarious; its almost complete absence from sedimentary rocks in

Ulu Kelantan is certainly not true of Malaya generally. Inspection of Table 4

shows that none of the subsample Burgess has analysed is on sedimentary rocks

over 2,500 ft, and the result for meranti bukit is therefore almost certainly an

artefact due to sampling bias.

Table 6. Groups showing a marked preference for granite and above 750 ft elevation

granite sedimentary

low high low high

(a) species

Dipterocarp. Shorea platyclados a3 . eh 73 0.4 4.3

Rubi. Anthocephalus chinensis v% ae 1.8 pay 0.3 1.8

(b) genera

Anacardi. Campnosperma .. me . 2.6 6.9 2.0 1.3

Jugland. Engelhardtia te! * 5 err ee But — —

Sterculi. Heritiera : Mey i 25 11 10

Annonaceae B “ 7 i (GS 29 14 21

Anthocephalus chinensis (cadamba) (kelempayan) is a species of valleys

and alluvial soils in Borneo; we have noticed it common on hill slopes, e.g. at

Genting Sempah, and this preference for hill forest is borne out by this survey.

Engelhardtia (dungun paya) is well known as a genus of hill and mountain

forest in Malaya.

Tree Species in the Rain Forest of Ulu Kelantan 205

(iii) Just granite, Table 7:

None of the preferences revealed here was previously suspected.

Table 7. Groups showing a marked preference just for granite

granite sedimentary

(a low high low high

a) species

Bombac. Bombax valetonii . a rae Pe 6.9 0.4 0.8

Dipterocarp. Dipterocarpus appendiculatus . . be 4.4 0.2 — —

Sterculi. Pterocymbium javanicum TF 8.8 7.3 2.0 1.0

(6) genera — nil

Bombacaceae ra ist a a 12 12 5.3 a

Fagaceae . . is ve his ne 40 60 30 31

Meliaceae ret sik ee fe 3.6 2.9 1.6 —

Sterculiaceae _ pe < es 41 69 37 37

Groups showing a marked preference for sedimentary rocks.

(i) Below 750 ft, Table 8:

Dryobalanops oblongifolia (keledan) is a riverine species. The other Dryobala-

nops in Malaya D. aromatica (kapur) is known strongly to prefer sedimentary

rocks; both are found mainly east of the Main Range.

Mesua ferrea (penaga) is a species of poor soils and dry sites, this is consonant

with its marked preference for sedimentary rocks.

Table 8. Groups showing a marked preference for sedimentaries and below 750 ft elevation

granite sedimentary

low high low high

(a) species

Dipterocarp. Dryobalanops oblongifolia. . M 17 Chas 66 5.6

Shorea bracteolata . . rs a 19 2.9 21 13

S. multiflora - sre 2 16 11 30 25

S. parvifolia m. he be 91 93 127 95

S. singkawang he om me — — 3.9 0.8

Guttif. Mesua ferrea me MS “ 4.4 3.3 14 12

Magnoli. Aromadendron elegans ‘ia a — — 3.9 -—

(6) genera

Combret. Terminalia be. f. si = — 3.9 _—

Dipterocarp. Dipterocarpus oe ane he 68 48 94 69

(ii) Above 750 ft, Table 9:

Shorea curtisii (seraya) is the most important timber tree of hill dipterocarp

forest; its distribution seems to be correlated with sites liable to periodic drought,

which tend to be mainly shallow, sedimentary-derived soils and hill ridges, as is

amply confirmed by the present survey (P. F. Burgess, unpublished observations).

Swintonia, Vatica and Calophyllum are well known to be commoner in the

hills. Their preference for sedimentary rocks was not known, though all are known

as genera of ‘poor’ soils or sites.

206 Gardens’ Bulletin— X XVI (1973)

The preference of the genus Shorea and of the family Dipterocarpaceae for

this habitat completely obscures the markedly differing preferences of the many

individual species enumerated and reflects the abundance of S. curtisii.

Table 9. Groups showing a marked preference for sedimentaries and above 750 ft elevation

granite sedimentary

@ low high low high

a) species

Dipterocarp. Shorea curtisii se! + 33 21 126 20 345

S. guiso 2% ” fy 7.9 11 11 16

(5) genera

Anacardi. Rengas* IP se y 11 21 22 35

Swintonia y be Me. 9.7 27 28 63

Dipterocarp. Shorea - 8 = ; pee 468 372 653

Vatica .. oh aS ie 1.8 6.0 7.7 14

Guttif. Calophyllum a ie, Td 17 29 23 34

{c) families

Anacardiaceae val ™ - 53 25 69 64 109

Dipterocarpaceae af: re me <5 Oe 602 587 815

Coniferae ne ae ne se i fee — — 1.8

* Gluta, Melanochyla, Melannorhoea, Semecarpus

(iii) Just sedimentary rocks, Table 10:

Koompassia malaccensis (kempas) is one of the commonest trees everywhere

in Malaya. Its preference for sedimentary rocks was not suspected and may not be

typical of the whole country.

Myrtaceae is a family which, throughout Malesia, becomes prominent in

‘poor’ sites, its preference in Ulu Kelantan for hills and for sedimentaries is not

unexpected.

Table 10. Groups showing a marked preference just for sedimentaries

granite sedimentary

low high low high

{a) species

Dipterocarp. Dipterocarpus crinitus ra . 4.4 5.8 20 21

Shorea dasyphylla . . aso an 1.8 1.6 2.1 2.3

S. faguetiana ; aks as 0.9 0.2 1.5 poe

S. macroptera a , + 0.9 Yi 3.3

S. ovalis ce ae le 26 33 51 34

S. ovata wd fs sa Meee — 14 1.8

S. sumatrana , a a — 0.8 0.3

Legum. Koompassia malaccensis - ne 36 39 66 45

Mor. Antiaris toxicaria m a — 0.1 0.8

Artocarpus integer . . ” << — 0.4 0.3

Olac. Ctenolophon parvifolius oe bas 1.8 2.0 5.7 4.3

(b) genera

Celastr. Lophopetalum 331 £ 0.9 1.3 ee 4.6

Legum. Koompassia + aye =e 54 56 84 63

Polygal. Xanthophyllum ot ™ 7.0 7.6 16 12

Rhizophor. Pellacalyx 1.8 0.9 2S 2.0

Myrtaceae Lo is ae if 73 122 124 133

Tree Species in the Rain Forest of Ulu Kelantan 207

Groups showing a marked preference for forest below 750 ft, Table 11:

Dipterocarpus oblongifolius (keruing neram) is a riverbank species.

Pometia pinnata var pinnata (kasai) is, in Malaya, a valley species.

Barringtonia (putat), as a genus, is commonest in valleys and along rivers.

Euodia (pepauh) is nowhere common in Malaya. It has been considered a

genus preferring the hills, an impression contradicted by this survey.

Pentace (melunak) likewise has been considered mainly a hill genus.

Table 11. Groups showing a marked preference for elevation below 750 ft irrespective of rock

granite sedimentary

low high low high

(a) species

Dipterocarp. Dipterocaropus oblongifolius . . se 09 — 0.4 -—

Dipt. cornutus + kunstleri .. gn 48 eA 60 29

Shorea balanocarpoides wy ‘es 7.9 if 5.2 3.3

Olac. Ochanostachys amentacea bes a 20 4.4 15 5.6

Sapind. Pometia pinnata v. pinnata... fo 18 9.1 i 9.7

(6) genera

Lecythid. —_ Barringtonia ¥ ae me 11 6.3 $s 5.9

Logani. Fagraea af Ai es 1.8 0.2 1.8 0.8

Oxalid. Sarcotheca <i ts afi 0.9 0.2 1.3 0.8

Rut. Euodia.. a a. re ait 1.8 6.8 i wes

Tili. Pentace AY a; * 4 13 7.6 12 9.8

Olacaceae - a &. At 46 35 46 36

Groups showing a marked preference for forest above 750 ft, Table 12:

Elateriospermum tapos, (perah), Cratoxylum formosum (derum) and Tristania

(pelawan) are well known to prefer hills.

Lauraceae characterise the Lower Montane or oak-laurel forest of Malaya

so their increase with elevation is also as expected.

Table 12. Groups showing a marked preference for elevations above 750 ft irrespective of rock

granite sedimentary

low high low high

(a) species

Dipterocarp. Dipterocarpus grandiflorus .. am 7 11 6.8 13

D. sublamellatus + verrucosus oo 0.7 0.1 4.3

Parashorea lucida .- . ny o— 13 4.6 7.6

S. leprosula i a vr af 88 77 82

Euphorbi. _Elateriospermum tapos ae Bs 16 45 20 49

Hyperic. Cratoxylum formosum 7" a 2 0.1 1.5

Mor. Artocarpus lanceifolius i a 12 25 15 iy

Rhizophor. Carallia brachiata . . sis ) — 04 — 0.3

Simaroub. Irvingia malayana . . ais ._ —_ 1 1.4 1.8

(5) genera

Dipterocarp. Anisoptera as ee ~ 21 37 20 48

Eben. Diospyros y a0 ¢. 1.8 2.9 2.0 4.3

Myrt. Tristania a“ 3 oo 1.1 0.8 1.8

Sterculi. Scaphium i - 7 13 30 20 24

Lauraceae a - fs a 57 94 72 68

208 Gardens’ Bulletin — XXVI (1973)

Ubiquitous Groups, Table 13:

This class includes several very common and easily recognised big tree

species (Shorea pauciflora, nemesu; Dyera costulata, jelutong: Cynometra

malaccensis, kekatong; Intsia palembanica, merbau; Koompasia excelsa, tualang;

Scorodocarpus_ borneensis, kulim) which no doubt have helped to create the

impression, sometimes held, that Malayan timber trees can grow almost anywhere

in the mesic Rain forest climax forests, though this impression is also undoubtedly

strengthened by the fact, revealed by this survey and commented on above, that

habitat affects frequency but seidom prevents occurrence.

No significance is attached to the long list of ubiquitous genera and families

for reasons already discussed.

Table 13. Ubiquitous groups

granite sedimentary

low high low high

a) species

Apocyn. Dyera costulata .. ae Es 6.1 8.7 6.7 6.9

Dipterocarp. Dipterocarpus baudii oi se a5 7.8 6.0 2.0

Shorea pauciflora - . e a 19 10 11 23

Euphorbi. _Baccaurea griffithii a - 3.5 2.0 2.4 2.3

Sapium baccatum .. , an 4.4 2.0 Zz 5.9

Legum. Cynometra malaccensis =e on 18 19 18 18

Intsia palembanica “a age 18 26 23 22

Koompassia excelsa ox :a 18 17 18 18

Olac. Scorodocarpus borneensis ~ ce 22 25 19 21

Strombosia javanica ae 34 2.6 1.3 3.2 2.5

S. rotundifolia - li Le — aud 3.2 2.5

Sapind. Nephelium lappaceum 4 ts 2.6 7.6 5.9 5.1

Pometia pinnata v. alnifolia_ . . aaa 2.6 6.0 9.1 5.6

{b) genera

Bombac. Durio .. - " - 4.4 4.5 4.0 2.0

Dilleni. Dillenia ay 7 =: 11 14 17 8.9

Fag. Castanopsis - a ee 14 9.6 11 12

Legum. Dialium he = 3 22 26 26 23

Sindora wm ae - 7.0 9.8 7.1 5.9

Sterculi. Pterospermum wa a % 1.8 se | 33 1.3

Apocynaceae . -: -. 6.1 9.8 7.3 7.9

Burseraceae ae 7 . os 43 90 67 66

Euphorbiaceae aie . - < 50 67 39 71

Leguminosae a os . #% 139 153 182 140

Guttiferae oy a “s st 21 33 38 47

Moraceae i aS if ai 34 43 29 30

Rhizophoraceae Jt es a o 1.8 1.3 2.5 2.3

Rubiaceae .* tye .. .- 1.8 2.6 0.6 1.8

Sapindaceae 4 - a = 24 24 27 20

Thymelaeaceae ys mM ott ii 1.8 6.0 6.2 2.8

Tree Species in the Rain Forest of Ulu Kelantan 209

Table 14. Groups which tend to avoid granite below 750 ft

granite sedimentary

low high low high

Legum. Pithecellobium splendens i} -f 0.9 4.5 3.8 2.0

Anacardi. Mangifera ae oe ne 1.8 7.8 z21 6.1

Pentaspadon iy . We 4 1.8 6.0 4.9 3.1

Burser. Santiria se § oe S 1] 29 pa 25

Hyperic. Cratoxylum Me ry oe == 2a 1.9 1.5

Mor. Ficus .. Aa Se tA: — psa | Zh 3.6

Myrt. Eugenia re “g eR 73 P| 123 131

Thymelaea. Aquilaria is 7 ee 0.9 4.7 uD 2.0

Table 14 shows groups which avoid low granite but are otherwise ubiquitous.

The area of this habitat sampled is much lower than of the other three habitats,

and therein lies the simplest explanation of this category.

ACKNOWLEDGEMENTS

I am grateful to Enche Salleh b. Mohd. Nor for allowing me to have a set

of the stand tables prepared from Mr. Cerra’s forest survey; to Mr. P.F. Burgess

for encouraging me to undertake the analysis, for commenting on a draft and for

allowing me to make use of his unpublished analysis of part of the same data;

also for considerable assistance with the computation to Enche2 Loh Hoy Shing,

T. Suppiah, P. Selveraj, Chan Yee Chong and Zainuddin b. Sohadi.

REFERENCES

Alexander, J.B. (1963) Geological Map of Malaya, 6th edition (Diamond Jubilee).

Burnham, C.P. & Whitmore, T.C. (1969) The altitudinal sequence of forests and

soils on granite near Kuala Lumpur. Malaya. Nat J. 22, 99-118.

Foxworthy, F.W. (1927) Commercial Timber Trees of the Malay Peninsula. Malay.

For: Rec. 3.

Kochummen, K.M. & Wyatt-Smith, J. (1964) Pocket Check List of Timber Trees,

Malay. For. Rec. 17.

Symington, C.F. (1943) Forester’s Manual of Dipterocarps. Malay. For Rec. 16.

Whitmore, T.C. (1971) Fruit trees and some trees of pharmacological potential ‘n

the rain forest of Ulu Kelantan. Malay. Nat. J. 24, 222-4.

Whitmore, T.C. (1972) Ed. Tree Flora of Malaya. Malay. For Rec. 26, Vol. 1.

Whitmore, T.C. (1973 a) Ed. Tree Flora of Malaya Malay. For. Rec. 26, Vol. 2.

Whitmore, T.C. (1973 b) Euphorbiaceae, in Whitmore, T.C. ed. Tree Flora of

Malaya. Malay. For. Rec. 26, Vol. 2.

Wyatt-Smith, J. (1964) A preliminary vegetation map of Malaya with descriptions

of the vegetation types. J. Trop. Geogr. 18, 200-13.

210

Gardens’ Bulletin — XXVI (1973)

APPENDIX

Species and groups with fewer than one stem per 100 acres.

No significance is attached to the different frequency of these between the

four habitats.

Anacardiaceae

Bouea

Annonaceae

Xylopia ferruginea

Cananga odorata

Apocynaceae

Alstonia

Bombacaceae

Coelostegia

Neesia

Coniferae

Agathis dammara

Podocarpus imbricatus

Dipterocarpaceae

Dipterocarpus apterus

D. chartaceus

D. gracilis

Shorea atrinervosa

. glauca

. laevis

. Maxwel'iana

. scrobiculata

S. multiflora

S. resina-nigra

ANN MN

Euphorbiaceae

Macaranga gigantea

Mallotus leucodermis /muticus

Guttiferae

Garcinia

Hypericaceae

Cratoxylum arborescens

Leguminosae

Adenanthera

Ormosia

Linanceae

Ixonanthes icosandra

Melastomataceae

Memecylon

Meliaceae

Amoora

Lansium domesticum

Sandoricum koetjape

Moraceae

Artocarpus kemando

Streblus elongatus

Rubiaceae

Adina

Randia scortechinii

Sapindaceae

Xerospermum

Theaceae

Gordonia concentricicatrix

Thymelaeaceae

Gonystylus

Ulmaceae

Gironniera

A New Bamboo from Mt. Kinabalu

R. E. HOLTTUM

Royal Botanic Gardens, Kew, U.K.

In 1956 I described five species of a new genus Racemobambos, slender bam-

boos from mountain forest, three of them from Mt. Kinabalu (Gard. Bull. Sing. 15:

267-273). When visiting that mountain in November 1972 I found a fourth species

in flower, on the ridge below Kambarangoh, a route followed by many persons who

have climbed the mountain during the past forty years. I did not find young culms

from which culm-sheaths could be described, and hope someone else will be able

to remedy this deficiency. The strongly tessellate lower surface of the leaves (at

least when dry) and their quite large size, are distinctive, and should be sufficient

to identify a plant which is not flowering. A comparison of living plants with

R. hirsuta is desirable.

Racemobambos tessellata Holttum, sp. nov.

Vaginae basium culmorum non visae. Culmus florifer c. 8 mm diametro,

glaber; rami ad quodque nodum c. 8, inaequales, usque 80 cm longi, longiores

ramosi; ramuli ultimi basi foliiferi apicem versus floriferi; folia usque 16 cm longa,

1.9 cm lata, subtus valde tessellata, basi auriculis parvis setas tenues flexuosas usque

10 mm longas ferentibus praedita, in apicem setiformi 5 mm longum terminata;

ramuli floriferi usque 8 cm longi, leviter hirsuti basin versus tantum; spiculae usque

3.5 cm longae; glumae vacuae 2, inferior 6-7 mm longa, 5-venosa, superior 8—9

mm longa, omnino glabrae; rachillae internodia 5 mm longa, apicem versus

adpresse sericea; lemmata 9-10 mm longa. marginibus omino minute ciliata, cetera

glabra, apice non setiformia; paleae 9-10 mm longae, carinis ciliatis; lodiculae c.

3 mm longae, ciliatae; antherae 5 mm longae.

TYPE: Mt Kinabalu, Sabah, below Kambarangoh, in ridge forest at 6500 ft,

Holttum 46, Nov. 1972. (SING; dupl. K.).

In its foliage this species is near R. hirsuta, which is only known from the

original Clemens collection on Penibukan ridge; but R. hirsuta is densely appressed-

hairy throughout the branches of the inflorescence and on the dorsal surface of

lemmas, and its leaves are not tessellate on the lower surface. The leaves of R. tessel-

lata have very distinct cross-veins connecting the smaller longitudinal veins with

which they make a tessellate pattern. R. tessellata has a'so a much more amply

branched inflorescence than R. hirsuta and its lemmas lack the short setiform tip

found in R. hirsuta.

211

210

Species asd cual IN r 3 4

sienificunce 16 atac hed 4p its di em tréc sey

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Notes on the Vegetation of the Cardamom Mountains, Cambodia

Marie A. MARTIN

Centre national de la Recherche Scientifique de Paris

Summary

The following pages present some floristic and phytogeographic observations made in the

course of ethnobotanical work. They must be considered merely as preliminary notes to a

future more detailed study of a massif of which only the periphery has been previously visited

by naturalists. It is therefore felt that, notwithstanding their incompleteness, these notes will

represent a useful contribution to our botanical knowledge of the western Cambodian moun-

tains, as little in the literature, and none of it in English, presently exists.

Introduction

In 1965-1966, thanks to a grant from the National Institute for Scientific

Research in Paris, I made my first ethnobotanical expedition to Cambodia. The

investigations were then principally carried out in the area around Leach in

Pursat province, and concentrated on the plants used by the lowland Cambodian

peasants. I therefore had occasion, through the study of their uses, to become

acquainted with tropical plants, and a flora, at first totally unknown to me,

and particularly with the species of the Dry Dipterocarp forests, the monsoon

forests rich in Leguminosae and Lythraceae, the swamp forests, and also those

of certain secondary types.* Some forays into the Cardamoms at this time

convinced me of the intrinsic interest, as much to anthropology as to botany,

that this, the most extensive montane area in Cambodia, presented, Further,

during my second mission which began in February, 1969, I decided on this

region, as little known to botanists as to ethnologists, as my study area. The

forests to the south of the range alone have been recently studied (1), apart

from some notes that are available on the vegetation of the Tamyong river basin

in Pursat province (2). The following pages can hardly claim to contain an

exhaustive study of this flora; they are merely observations that have been

made within the framework of an ethnobotanical investigation: on-the-spot

records that have for the most part been made along the lanes and tracks figured

in map 2. It need hardly be added that the lists of species are very incomplete, but

as far as possible the dominant or characteristic species have been noted; some.

even of these, are yet to be identified.

The herbarium specimens collected are retained in the Natural History

Museum in Paris. Those already identified are incorporated in the herbarium of

the Phanerogam Laboratory; the remainder, which will be examined on my

return to France, can meanwhile also be consulted in this laboratory, It is for

this reason that I cite collection numbers for all unidentified species.

*T also made very limited number of collecting trips in lowland mixed rain forest.

1. P. Dy Phon, 1969. La végétation du sud-ouest du Cambodge. Thesis, Toulouse

University, 261 pp. (unpublished). (This has subsequently been published in Ann. Fac. Sc.

Phnom Penh 1970, no. 3, pp. 1-135 & 1971 no. 4, pp. 1-77 — ed.)

2. M. Schmid, 1969. Considérations sur la végétation du centre-ouest Cambodgien. Ann.

Fac. Sc. Phnom Penh, 2: 81 - 100.

213

214 Gardens’ Bulletin— XXVI (1973)

The field-work was carried out without any outside scientific assistance

except for some botanical excursions made on the southern flanks of the massif

with Dr. P. S. Ashton. I am most grateful to him for this opportunity to exchange

our botanical experience, and for his encouragement; I am also indebted to

him for the English translation of this manuscript.

The Vegetation Types

No extensive rainfall figures are available for the Cardamom massif itself,

merely limited observations from the Thmar Beng region near the centre and

some valuable summaries from Khemarak Phouminville on the coast. The annual

rainfall for the former (extrapolated from incomplete data) is approximately

4 m., and 5 m. for the latter; neither do detailed soil studies exist. Further,

for non-scientific reasons it has not been possible to explore the montane vegetation

above 1000 m. altitude.

The vegetation is here delineated according to the physiography; the main

types to be considered are:

Forests of humid valley bottoms, notably riverain forests,

Forests of the slopes.

Forests of ridges.

Some notes are also provided for:

Moist submontane grasslands.

Monsoon forests (foréts claires) and savanna forests (foréts séches).

1. Riverain forests

These form a narrow fringe along the margins of the rivers. Where the

valley is narrow and enclosed, the species of the hillslopes descend to the waterside;

these will be described later. When the valley is broad, the forest which borders

the river contains several characteristic species, one of which, Altingia (siamensis)

is nearly always present. The structure is that of a true closed evergreen forest,

with emergent species such as:

Altingia (siamensis)

Anisoptera sp. or Hopea odorata Roxb.

Tristania (burmanica)

Hopea is strictly riparian and disappears when one penetrates into the forest.

The following species are prominent in the middle and understorey:

Syzygium zeylanicum (L.) DC

Pandanus (tectorius)

Fagraea racemosa Jack ex Wall.

Ficus sp.

Cinnamomum sp.

Xanthophyllum glaucum Wall. ex Hassk.

With the exception of Hopea and X. glaucum the other species cited here

are nearly always present and form the characteristic floristic component.

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Vegetation of the Cardamom Mountains 217

2. Evergreen forests of ridges and plateaux

(a) Forest with Fagaceae, Lauraceae, Myrtaceae, Meliaceae. This occurs also on

some slopes free of rocks. It is possible to recognize several strata:

An upper and a lower canopy stratum; these when taken together are dense.

Trees of the former are often variable both in height and diameter, yet

form a more or less complete cover except in certain types where the

understorey is very dense. The lower canopy stratum only sometimes forms

a complete cover, Taken together, the espacement between individual trees

is typically 3-5 m. The boles are cylindrical and straight, without prominent

buttresses though often somewhat flared at the base; in some areas between

a quarter and a third of the lower canopy trees are leaning. The crowns of

the upper canopy stratum are generally light, with diffuse foliage; those of

the lower tend to be lanceolate, and the stratum taken as a whole is more

dense. For the most part the first branches appear in the upper two-thirds

to three-quarters of total height of the trees; they generally arise from the

same point at the bole apex, with the exception of certain monopodial

species in the lower canopy, notably Garcinia vilersiana Pierre and Sageraea

elliptica (A. DC.) Hook. f.

The understorey stratum includes many Rubiaceae and spiny palmlets as

well as tree ferns. Along the roadsides a Melastomataceous shrub Medinilla

coerulescens Guill. is very abundant.

The herb and shrub stratum consists essentially of zingibers, some ferns, and

a semi-woody gregarious Strobilanthes which can reach 24 m. and which

often by itself consistitutes the shrub layer. In open glades a Cyperaceae

(1537) is usually noted.

Epiphytes are abundant both in the upper and lower canopy strata: orchids

of many kinds, the smallest growing in among the moss; many ferns, in

particular Asplenium, Platycerium and Drynaria; mosses and hepatics are

conspicuously abundant.

Herbaceous and semi-woody climbers are frequent, twisted round the tree

trunks; large woody climbers, reaching into the upper-most branches, are

less abundant.

Appearance of the understorey: When the forest is completely closed, as

is generally the case, the understorey though lush is easy to pass through

in spite of the presence of spiny palms. When light penetrates the canopy to

any extent, the more the understorey (both woody and herbaceous) is

illuminated the more it burgeons; the plants become entangled and progress

becomes exceedingly difficult.

This evergreen forest with Lauraceae, Fagaceae and Myrtaceae is the most

widespread in the massif, It is rich in species, of which the most frequent

are:

MAIN CANOPY:

Cinnamomum aff. javanicum Dysoxylum procerum Hiern.

Cinnamomum sp. (1443) Schima crenata Korth.

Syzygium sp. (1553)

Syzygium aff. cochinchinensis

Parki

Quercus chrysocalyx Hick. & A. Camus arkia streptocarpa Hance

Castanopsis pierrei Hance Baccaurea oxycarpa Gagnep.

Dysoxylum cauliflorum Hiern. Aquilaria krassna Pierre ex Lecomte

Dysoxylum aff. hoaensis Garcinia hanburyi Hook. f.

Podocarpus imbricatus Bl.

218

UNDERSTOREY:

Areca triandra Roxb. or Pinanga

duperreana Pierre ex Becc. (The two

species never occur together.)

Daemonorops aft. pierreanus

Cyathea podophylla (Hook.) Copel.

HERB AND SHRUB STRATUM:

Amomum kravanh Pierre ex Gagnep.

(the Cambodian Cardamom plant,

confined to rich soils).

Amomum spp. (Among which is No.

1507)

Alpinia spp. (Among which is No.

1709)

CLIMBERS AND EPIPHYTES:

Albizzia sp.

Apocynaceae (1528)

Dischidia aff. imbricata (Bl.) Steud.

Dischidia nummularia R. Br.

Certain Annonaceae

Cassytha filiformis L.

Araceae (Pothos spp. ?)

MOSSES AND HEPATICS*:

Acroporium sp.

Bazzania sp.

Chaetomitrium sp.

Cheilolejeunea ceylanica (St.) Schust.

& Kahr.

Cheilolejeunea verdonii (Hoff.) Schust.

& Kahr.

Cololejeunea sp.

Dicranoloma braunii (C.M.) Par.

Drepanolejeunea thwaitesiana

Ectropothecium buitenzorgii (Bel.)

Jaeg.

Gardens’ Bulletin — XXVI (1973)

Cyathea latebrosa (Wall.) Copel.

Some Rubiaceae

Medinilla coerulescens Guill.

Some spiny Smilax spp.

Strobilanthes sp.

Many vascular cryptogams, especially

Selaginella siamensis Hiern. and

Pteridium esculentum (Forst.) Coo-

kayne

Gleichenia aft. norristi

Asplenium nidus L.

Platycerium coronarium Desv.

Drynaria spp.

Many orchids, notably Eria,

Bulbophyllum, Dendrobium and

Appendicula spp.

Fissidens aft. nobilis Griff.

Homaliodendron crassinervium Ther. °

Leptolejeunea subacuta St.

Leucobryum bowringii Mitt.

Leucoloma molle (C.M.) Par.

Plagiochila sp.

Radula acuminata St.

Raphidolejeunea subacuta St.

Trachypus bicolor Reinw. & Hornsch.

Trichocolea pluma Mont.

Thuidium meyenianum (Hamp.) Jaeg.

* Identified by Mr. Pierre Tixier.

Vegetation of the Cardamom Mountains 219

Besides these species there are others which can become locally abundant within

this forest type:

TREES:

A very large Apocynaceae, reaching (?) Aglaia sp.

50 m. tall (1533) Maba sp.

Anisoptera sp. (?) Lauraceae: No. 1548

Dipterocarpus turbinatus Gaertn. f. Leguminosae or Sapindaceae (1534)

Dipterocarpus costatus Gaertn. f. Irvingia malayana Oliv. ex Benn,

Vatica odorata (Griff.) Sym. Castanopsis cambodiana A. Cheval.

Ficus sp. (1562) Lithocarpus cerifera Hick. & A. Camus

Heritiera javanica (Bl.) Kost. Calophyllum sp. and other Guttiferae

Sapium baccatum Roxb. Melia azedarach L.

Adenanthera pavonina L. Baccaurea sapida Muell.-Arg.

SHRUBS AND HERBS:

Sageraea elliptica (A.DC.)} Hook. f. An arborescent Pandanus

Evodia triphylla DC.

Certain species reach 1000 m. altitude (and perhaps higher), such as many

Myrtaceae, some Lauraceae and Guttiferae, Schima crenata, Podocarpus

imbricatus and Sapium baccatum.

On the other hand there are others confined to middle altitudes, including:

All Meliaceae Ficus sp.

Parkia streptocarpa Apocynaceae No. 1533

Dipterocarpus costatus

Most palms (excepting Pinanga duperreana) do not exceed 600-800 m.

By contrast: Jrvingia malayana, Heritiera javanica, Quercus chrysocalyx,

Baccaurea oxycarpa, Melia azedarach disappear near 400-600 m.

Yet others only appear at certain restricted altitudes: the genus Cyathea,

at c. 500 m., Pinanga duperreana, at c. 600 m., Fagaceae, No. 1549, at

c. 800 m.

On red soil derived from basalt, Meliaceae, the larger Lauraceae notably

Cinnamomum afi. javanicum, Tetrameles nudiflora, Sapium baccatum and

Syzygium sp. (1553) are particularly well represented.

(b) Forest with Hopea pierrei Hance, Quercus chrysocalyx Hick. et A. Camus,

Anthocephalus sp. (1739).

One finds this type in the western part of the mountain range, One dominant

species, Hopea pierrei, exceptionally successful, prevails at all levels, as

saplings, poles and mature trees. It grows also to the north, around the

plateau of Veal Veng in the same forest type. The height of its crown

hardly exceeds 20 m. In both areas the soil is a red lateritic clay. Two

species of medium height (c. 30 m.) are associated with Hopea pierrei:

Anthocephalus sp. (1739) and Quercus chrysocalyx. A Tristania (1736) which

is sometimes associated seems to prefer in particular the moister habitats.

Finally, another Fagaceae appears in this type, which it has not so far

been possible to collect.

Apart from these distinctions, one meets in this forest type nearly all the

trees of evergreen forest type (a), and the understorey is equally similar.

220 Gardens’ Bulletin — XXVI (1973)

Locally, several dipterocarps appear: besides Hopea pierrei, Anisoptera sp.

and Dipterocarpus costatus can occur. In the forests to the north, Vatica

odorata is abundant adjacent to Hopea pierrei and Anthocephalus sp.

In rocky places these species are mixed with a bamboo (Bambusa sp.,

1710).

3. Forests on slopes, with bamboo

it can happen that, on the slopes of certain phnoms*, the vegetation may not

be different from that which covers the summits; this occurs particularly where

the soil remains of more or less uniform depth. Generally, however, the slopes

present the appearance of a mosaic of sandstone blocks, the prevailing substrate

of the Cardamom mountains, and of more or less humiferous clay screes — resulting

from the decomposition of certain types of sandstone.

On occasion walls of exposed rock strata are exposed, as if the water, in

seeping down the slopes had exposed a buried relief; on others they occur in

irregular screes. Sometimes also, landslips have carried the sandstone down from

the hilltops towards the lower slopes, where it accumulates; the persisting scars

on the slopes, and the blocks that mark their course, remain as testimony. Be that

as it may, whether the rock rests in situ or has fallen from the ridges, there is

hardly any room for the establishment of vegetation.

Here bamboos (Bambusa sp., 1710, 1696) are dominant. Someiimes there

are few trees, or even none. On the ground an Alpinia and a small species vf

fern can grow among the bamboos, but are local. Above 500 m. one meets some

cyatheas here and there. Two large and aggressive invading ferns, Gleichenia aff.

norrisii and Dicranopteris sp. can grow among the bamboos and even completely

replace them. It appears that this comes about when the soil is particularly

impoverished. Certain spiny plants, such as Rubus spp. (especially bordering

tracks) and certain climbers can render such thicket impenetrable.

Between these communities with bamboos and the majestic evergreen forests

occurring on ridges and here and there on slopes, there can occur a transitional

type where some species of the evergreen forests mix with the bamboos. Quercus

chrysocalyx is the most frequent but all the other species can occur from time

to time; their density is generally low and their height hardly exceeds 20 m.

The spiny palms, and in particular the rattans, are numerous.

These bamboo forests climb to 1000 m., possibly higher seeing that we know

so little about the vegetation that grows above that altitude.

4. The vegetation of the wet grasslands

The tracks that cross the range from time to time open out into plateaux

covered with herbaceous vegetation. Trees are very scattered or for much of the

time absent. On the ground grasses and Drosera (indica) occur in abundance;

an unidentified herb (1664) is generally associated with them. Locally, Leptocarpus

disjunctus Mast. is dominant, at which time it is associated with Centrolepis

cambodiana. Further plants encountered here sometimes are a Euphorbiaceae

(1769) and another small undetermined herb (1764).

This vegetation has been observed to occur over sandy soil and on surfaces

where the iron pan, close to the surface, outcrops here and there.

Where swampy terrain occurs it is hemmed by giant grasses such as Saccharum

arundinaceum Retz., by Phragmites sp. or by Cyperaceae.

*Phnom: a mountain, hill or mound.

Vegetation of the Cardamom Mountains 221

5. Dry deciduous forest, Erythrophloeum cambodianum Gagnep., Irvingia

malayana, Lagerstroemia sp.

On the slopes of certain phnoms to the north of the range grows a semi-

deciduous forest, characteristised by the abundance of leguminous trees,

TREES: the following are dominant:

Erythrophloeum cambodianum Gagnep.

Irvingia malayana

Lagerstroemia sp.

Bombax ceiba L.

(?) Nephelium sp.

and are accompanied by:

Dalbergia nigrescens Kurz Ficus sp.

Xylia kerrii Craib & Hutch. Aquilaria krassna Pierre ex Lecomte

Sindora cochinchinensis Baill. Garcinia ferrea Pierre

Pterocarpus sp. Terminalia nigrovenulosa Pierre

Near the summits Tetrameles nudiflora R.Br. is noticed.

SHRUBS:

Cratoxylon sp.

Memecylon sp.

Some Rubiaceae

A few climbers and spiny plants

The ground is covered by numerous ferns and zingibers, including Amomum

kravanh Pierre ex Gagnep.

Many of the species quoted grow in dry habitats and one comes across them

often in the deciduous formations of the plains of Tonlé Sap and Mekong. In

the mountains they are to be found up to 500 m. altitude on rocky outcrops with

shallow soils.

6. The savanna forests

(a) The pine forests:

Pinus merkusii Jungh. et de Vriese. These are as a rule submontane, but

they do exist also in the lowlands. On the Cardamoms themselves they

follow certain cuestas and plateaux.

In the regions visited Tristania burmanica is the best represented woody

species though Dipterocarpus obtusifolius Teysm. and Melastoma sp. also

occur; /mperata cylindrica P. Beauv. and Selaginella sp. dominate the field

layer. In all, the number of species appears to be low.

Gardens’ Bulletin — XXVI (1973)

(b) The savanna forests with dipterocarps:

Unique to South-East Asia, these, frequently called Dry Dipterocarp forests,

are widespread in Cambodia; they occur to several hundred metres altitude

and are generally found on sandy soils. The best represented species in

them are:

Shorea obtusa Wall. ex BI., which can form almost pure stands.

Dipterocarpus tuberculatus Roxb.

Dipterocarpus obtusifolius Teysm.

Pentacme siamensis Kurz

Dipterocarpus intricatus Dyer, which seems to indicate somewhat better

soil conditions.

The Dipterocarpaceae can alone constitute the single tree stratum, or equally

can be accompanied by:

Careya sphaerica Roxb.

Aporosa sphaerosperma Gagnep.

Terminalia alata Heyne ex Roth

Mitragyna brunonis (Wall. ex G.Don)

Craib

Diospyros erhetioides Wall.

Melanorrhoea laccifera Pierre

Sindora cochinchinensis Baill.

Xylia kerrii Craib & Hutch

Parinari anamensis Hance

Buchanania reticulata Hance

Dillenia ovata Wall. ex Hook. f. &

Thoms.

Dillenia pentagyna Roxb.

Zizyphus cenoplia Mill.

Randia tomentosa Bl.

Feroniella lucida (Teijsm, & Binn. ex.

Scheff.) Swingle

A bush with distinct red bark (1349)

In the understorey Corypha lecomtei Becc. is particularly abundant; it

occurs also in analogous vegetation types to the north of Kompong Thom

away on the other side of the Great Lake. The most abundant and

ubiquitous herb is Arundinaria falcata Nees, though it can sometimes be

replaced by Imperata cylindrica P. Beauv., notably after persistent burning.

It should be recalled that the savanna forests, including the pine forests, are

for the most part burned annually to facilitate the hunting of game which

frequent them. Some foresters (3) believe that these types are mostly

man-induced and that the climatic climax-type is an unfired savanna forest

witli dipterocarps; such a savanna forest, which still occurs here and there

on the plains, includes many Leguminosae:

Xylia kerrii Craib & Hutch,

Sitndora cochinchinensis Baill.

Peltophorum aff. pterocarpa

Afzelia xylocarpa Craib

as well as:

Lagerstroemia spp.

Combretum quadrangulare Kurz

Terminalia sp.

Terminalia nigrovenulosa Pierre

Dalbergia nigrescens Kurz

Dialium cochinchinensis Pierre

Dalbergia aff. cochinchinensis

Dalbergia aft. lanceolaria

Bombax ceiba L.

Shorea talura Roxb.

The shrub (1349) already mentioned.

Sometimes also, Dipterocarpus intricatus Dyer occurs.

A bamboo (1696), encountered also in evergreen forest, grows at times in the

understorey.

P. Maurand, 1968. Politique forestitre 4 envisager au Vietnam, dans l’aprés-guerre..

(3)

Bull. Soc. Et. Indochin., Saigon, 43, 4: 267-309.

The Xylopia malayana fruit: significance of its dehiscence

HSUAN KeEnG* & E. A. HEASLETT**

Xylopia malayana Hook. f. & Thoms. of the family Annonaceae, is a small

tree, 10-20 m tall, with smooth, pale greyish bark, distributed in Malaysia, Singa-

pore, and Sumatra (Sinclair 1955). In Singapore it can be found scattered in

lowland forest around MacRitchie Reservoir and in the Bukit Timah Nature

Reserve.

The mature carpels of this tree are oblong obovoid, 3—3.8 cm long, 1.5—-18 cm

across, slightly ridged on the dorsal side and shortly stipitate at the base. They are

greyish green to yellowish green in colour, sometimes with a pinkish tinge when

fully matured. The surface of the carpels is densely covered with short, velutinous

hairs. The mature carpels, usually 2, 3 to many (12 or so) in a bundle (Figure 1a),

form an aggregate fruit, or botanically a follicetum.

The mode of dehiscence of these mature carpels is interesting. A dehiscent

line at first appears at the median of the dorsal suture, and from there it extends

towards both apical and basal ends (Figure 1b). After the carpel has completely

f) ff

‘y

or,

——

Fig. 1. Xylopia malayana Hook. f. & Thoms. a. An aggregate fruit (consisting of 9 free

carpels); 6. A mature carpel dehiscing along the dorsal suture; c. A dehiscent

carpel with seeds arranged in two rows on both sides of the ventral suture.

* Botany Department, University of Singapore

** Malayan Nature Society, Singapore Branch

224 Gardens’ Bulletin— XXVI (1973)

dehisced along its dorsal suture, it exposes the soft, somewhat juicy, pink to scarlet-

red pulp in which two rows of seeds were embedded, showing 2 to 4 seeds on each

row, alternatively arranged along the ventral suture (Figure 1c). In the freshly

exposed fruit, the seeds are coated with a thin layer of creamy white, waxy sub-

stance. After the seeds are exposed in the air for a day or two, the colour of the

seed coat gradually changes from creamy white to pale chestnut and then to dark

chestnut brown. The fruit wall or pericarp is soft leathery in texture. The two

wing-like valves of the dehiscent carpels gradually curve acutely backwards, often

severely damaging the adjacent tissues near the apex and base of the fruit. There-

fore, in an advanced stage of dehiscence, carpels are often found to be further

split along the ventral suture to a greater or lesser extent.

The trees which we examined in Singapore and in Southern Johore, Malaysia

were fruiting during October and November, 1972. A considerable amount of fruit,

partly or fully dehiscent, was found on the ground under the trees, and some of it

had been damaged by an unknown animal, Since the open carpels possess a faint,

fragrant smell, and display a mass of bright pink to scarlet pulp with several white

to brown seeds, it seems probable that the seeds of Xylopia malayana are dispersed

by birds. (cf. Pijl 1969).

Hutchinson (1959, 1: 134) gives a general description of the fruit of

Annonaceae as follows “‘carpels usually stipitate in fruit, free, rarely united into

1— or many-locular mass, dry or fleshy, rarely dehiscent”. Sinclair (1955, p. 168)

observed that in the Malayan Annonaceae, “the carpels are indehiscent, except in

the case of Anaxagorea, where the carpels are follicles. In some species of Xylopia

and Alphonsea, there is a kind of semi-dehiscence but the carpels do not open by

well-defined sutures’.

Further it is interesting to note as Sinclair pointed out (p. 347) that there is a

strong resemblance between Anaxagorea and Xylopia in their carpels, stamens and

geographical distribution, Both genera, together with several others are classified

under the tribe Xylopieae by Sinclair, but reduced to the Xylopia Group of the

tribe Unoneae by Fries (1959).

It is generally accepted that the follicular fruit probably represents the basic

pattern of fruit among angiosperms (Eames 1961, Takhtajan 1969). Most members

of the modern Magnoliaceae have follicular fruits. The fruit of the ancestral form

of the Annonaceae is very likely of a similar nature. Therefore the presence of

follicular carpels in Anaxagorea (which contains only one or two nearly basally

attached ovules and seeds in each carpel) and Xylopia (which has a special mode

of dehiscent as described above) probably indicate that each of these two genera

retains in part the characteristics of the fruit of the putative ancestral form of the

family.

It should also be pointed out that the main dehiscent line of the carpels in

Xylopia, like that of many members of the Magnoliaceae, is along the dorsal suture.

This is in contrast to the definition of a follicle as given in most glossaries and to

the follicular fruits as found in Anaxagorea and in various other families (e.g.

Illiciaceae, Dilleniaceae, etc.) in which the dehiscent line is invariably along the

ventral suture.

Monographers of the Annonaceae generally agree that the flower structures of

Anaxagorea, Xylopia, and allied genera which constitute the tribe Xylopieae, are

all characterized by the valvate sepals and petals, and are thus considered to be of

fairly advanced nature. The retention of the more primitive characters of the fruit

in these genera may be explained by postulating a discrepancy in evolutionary rate

between the flowers and the fruits.

Plate 1. Xylopia malayana Hook. f. & Thoms.

Above: a fruiting branch with dehiscent carpels and seeds;

Below: an aggregate fruit and a dehiscent carpel.

(scale: 1 cm divisions) Photo by Mr. D. Teow

The Xylopia malayana fruit 225

References

1. Eames, A. J. (1961) Morphology of the Angiosperms. New York.

2. Fries, R. (1959) Annonaceae, in Engler & Prantl, Nat. Pflanzenfam. 2nd ed.

17a If: 1-170.

Hutchinson, J. (1959) The families of flowering plants. 2nd ed. Oxford.

van der Pijl, L. (1969) Principles of dispersal in higher plants, Berlin.

Sinclair, J. (1955) A revision of the Malayan Annonaceae, in Gard. Bull.

Sing. 14 (2): 149-476.

6. Takhtajan, A. (1969) Flowering plants, origin and dispersal (Engl. transl. by

C. Jeffrey). Edinburgh.

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Nepenthes from Borneo, Singapore and Sumatra

Shigeo KURATA*

This paper comprises the result of an expedition to Indonesian Islands in

search of Nepenthes. It records thirteen species and describes two of them as

new taxa. All but one were collected during this research trip made by the author

and G. Mikil (participant from Malaysia) to Borneo, Singapore, Java and Sumatra

in March 1972.

The new taxa are N. campanulata and N. rhombicaulis. The author is grateful

to Drs. C. X. Furtado and K. L. Chang for their kind assistance.

1. Nepenthes alata Blanco, FI. Filip., ed. I, p. 805 (1837).

SUMATRA: West Sumatra, Bukittinggi, en route to Lake Manindjau, 1100 m

alt., Kurata & Mikil 4101, 4102, 4103.

Distribution. Philippines, Malaya, Sumatra and Borneo.

The plants grow on the sand-stone cliff of a river. The two wings on the

pitcher wall are not ciliated at the margins. The leaf is sessile and the stem is

remarkably trigonous with the wider wings. By these characters the Sumatran plants

vary from those of the Philippines.

2. Nepenthes ampullaria Jack, Comp. Bot. Mag., I, p. 271 (1835).

SINGAPORE: MacRitchie Reservoir, 50 m alt., Kurata & Mikil 4030, 4031,

4032. SUMATRA: North Sumatra, near the headwater of Asahan river, 800m.

alt., Kurata & Mikil 4309, 4310, 4311, 4312, 4313.

Distribution. Malaya, Singapore, Borneo, Sumatra and New Guinea.

3. Nepenthes bongso Korth., Verh., Nat. Gesch. Bot. p. 19, t. 14 (1839).

SUMATRA: West Sumatra, Mt. Talang, 1540-1950 m alt., Kurata & Mikil

4245, 4246, 4247, 4248, 4249, 4250, 4251, 4252.

Distribution. Sumatra.

4. Nepenthes campanulata Kurata, sp. nov. (Pls. 1 & 2) — Nepenthes campanu-

lata (nomen nudum) Kurata, The Heredity, Vol. 26, No. 10, p. 44 & 50 (1972).

Planta tenuis, breviter scandens. Caulis 20-35 cm altus, 3-4 mm crassus,

cylindricus, glaber. Folia sessilia, 5-7 cm longa, 1.2—2 cm lata, spathulato-lanceolata,

basi } “+ amplexicaulia, apice rotundata et leviter peltata, coriacea, utrinque glabra;

nervi longitudinales utrinque 2-3, nervi transversales obscure, oblique ascendentes;

cirrhus tenuis, 2.5—4 cm longus, cylindricus, glaber. Ascidia monomorphia, campanu-

lata, 4.5-7 cm longa, parte inferiore cylindrica vel leviter ventricosa, 1—1.5 cm lata,

_*Insectivorous Plants Society of Japan, Department of Biology, Nippon Dental College,

Fujimi, Chiyoda-Ku, Tokyo, Japan.

227

228 Gardens’ Bulletin — X XVI (1973)

parte superiore infundibuliformia, 3.5-5.5 cm lata, luteo-viridia, diffuse purpureo-

maculata, herbacea, glabra, intus parte 4 inferiore glandulosa, costis 2 prominenti-

bus; os circulare, fere horizontale; peristomium fere 0, breviter denticulatum;

operculum eilliptico-oblongum, 1.2-1.5 cm longum, facie inferiore planum; calcar

ca. 1 mm longum, filiforme. /nflorescentia ignota.

Stem slender, shortly climbing, 20-35 cm high, the part with adult leaves

3-4 mm thick, cylindrical, glabrous. Leaves scattered, coriaceous, sessile, spathulate-

lanceolate, 5-7 cm long, 1.2—2 cm broad, the base 2-2 amplexicaul, rounded and

slightly peltate at the apex; longitudinal nerves 2-3 on both sides, pinnate nerves

obliquely ascending; both surfaces glabrous; tendril slender, 2.5-4 cm _ long,

cylindrical, glabrous. Pitchers herbaceous, campanulate, 4.5-7 cm high, 1-1.5 cm

broad at the lower cylindrical or slightly ventricose part, 3.5-5.5 cm broad in the

upper infundibuliform part, with 2 prominent ribs; mouth circular, almost hori-

zontal; peristome almost none, the minute teeth fringing the mouth; inner surface

of the pitcher glandular in the lower 4 part; the pitcher yellow-green with purple-

spots, glabrous; lid elliptic-oblong, 1.2-1.5 cm long, under surface plain; spur

filiform, about 1 mm long. /nfiorescence unknown.

Typus: A. Kostermans 13764 (SING.); foot of Mt. Ilas Bungaan, East

Borneo, 300 m. alt., Sept. 9, 1957.

This new species is closely related to N. inermis Danser of Sumatra by its

slender habit and campanulate pitcher without conspicuous peristome. However

present species differs clearly from the latter by the lamina with peltate portion

near the apex, elliptic-oblong lid of a remarkably small size (its length always

measures less than 4 of the mouth-diameter) and existence of glandless but waxy

zone which occupies greater part of the inner surface of the pitcher. The digestive

glands on the inner surface of the pitcher are all round (not overarched) and

are only distributed in the bottom part with a sparse density of 100-250 glands

per square cm. Also, colour of dried specimen distinctly differs from that of

N. inermis. Danser described the dried specimens of N. inermis as blackish in

all parts (examined by the author), but those of present species are reddish brown

all over. See addendum on p. 232.

Though this specimen collected by A. Kostermans is the only one and has

no inflorescence, these characters mentioned above sufficiently support the author’s

conclusion that this species is a new one. N. campanulata is not a natural hybrid

as there is not any known species having these characters which could claim to

be its parental species.

Distribution of N. campanulata seems limited to a particular area presumably

owing to its habitat which is noted as the sand and limestone walls at an altitude

of 300 meters by the collector, whereas N. inermis was found only on high places

(1800-2590 m. alt.) in the mountains in central Sumatra.

5. Nepenthes carunculata Dans., Bull. Jard. Bot. Buitenz. ser. III, Vol. IX, p. 277

(1928).

SUMATRA: West Sumatra, Mt. Talang, 1900-2000 m alt., Kurata & Mikil

4218, 4219, 4220, 4221, 4222, 4223.

Distribution. Sumatra.

o ee SIE. Ge RATT aC D a IOs IAI IE

Plate 1. Nepenthes campanulata Kurata (x 0.5).

Plate 2. Nepenthes campanulata: a pitcher (x 1.5).

Nepenthes from Borneo, Singapore and Sumatra 229

6. Nepenthes dubia Dans., Bull. Jard. Bot. Buitenz. ser. III, Vol. IX, p. 285 (1928).

SUMATRA: West Sumatra, Mt. Talang, 1500-2000 m alt., Kurata & Mikil

4230, 4231—a, 4231—b, 4233, 4234, 4235, 4236.

Distribution. Sumatra.

7. Nepenthes gracilis Korth., Verh., Nat. Gesch. Bot. p. 22, t. I & IV, ic. 1-38

(1839).

BORNEO: Sabah, Bukit Padang, near Kota Kinabalu, 300 m alt., Kurata &

Mikil 4020, 4021, 4023. SINGAPORE: MacRitchie Reservoir, 50 m alt., Kurata &

Mikil 4050, 4051, 4052.

Distribution. Borneo, Sumatra, Celebes, Malaya and Singapore.

8. Nepenthes mirabilis Druce, Fl. coch., II, p. 606 (1790).

SUMATRA: West Sumatra, Indarung, 10 km east from Padang, a swampy

shrub, 250 m alt., Kurata & Mikil 4256, 4257, 4258, 4259, 4260, 4261, 4262, 4263.

Distribution. Southern China, Indochina, Malaya, Singapore, Java, Sumatra,

Borneo, Philippines, Moluccas, New Guinea and Australia.

This species varies somewhat on each island. In the case of Sumatran plants

the pitcher is very narrow and its peristome has a distinctly flattened surface.

9. Nepenthes pectinata Dans., Bull. Jard. Bot. Buitenz. ser. III, Vol. IX, p. 350

(1928).

SUMATRA: West Sumatra, Mt. Singgaiang, 2400-2600 m alt., Kurata &

Mikil 4202, 4203, 4204, 4205, 4206; Mt. Merapi, 2400 m alt., Kurata & Mikil

4211, 4212, 4213, 4214, 4215, 4216.

Distribution. Sumatra.

10.. Nepenthes rafflesiana Jack, Comp. Bot. Mag., I, p. 270 (1835).

BORNEO: Sabah, Bukit Padang near Kota Kinabalu, 300 m alt., Kurata &

Mikil 4006, 4007, 4008, 4009, 4010, 4011, 4012. SINGAPORE: MacRitchie Reser-

voir, 50 m alt., Kurata & Mikil 4040, 4041.

Distribution. Malaya, Singapore, Borneo and Sumatra.

11. Nepenthes rhombicaulis Kurata, sp. nov. (Fig. 1) — Nepenthes rhombicaulis

(nomen nudum) Kurata, The Heredity, Vol. 26, No. 10, p. 44 (1972).

Planta robusta, alte scandens. Caulis 5-20 m altus, 5-10 mm crassus, quadratus

vel rhomboides, glaber. Folia sessilia, 12-22 cm longa, 3-4 cm lata, lanceolata,

herbacea, basi 4-4 amplexicaulia, utrinque glabra, margine ferrugineo-pubescentia,

supra lucido-viridia, subtus luteo-viridia, nervi longitudinales utrinque 2-3, nervi

transversales obscure, oblique ascendentes; cirrhus tenuis, 10-13 cm longus, cylin-

dricus, glaber, luteo-viridis. Ascidia inferiora et rosularium 6-12 cm longa, 2.5—3.5

cm lata, parte inferiore ventricosa, parte superiore cylindrica, rubro- vel pallido-

viridia, diffuse purpureo-maculata, coriacea, sparse pubescentia, intus parte +

inferiore glandulosa, bialata, alis margine dentatis; os orbiculare obliquum; peris-

tomium 3-5 mm latum, subcylindricum, rubrum, margine exteriore expansum,

undulatum, margine interiore denticulatum; operculum elliptico-oblongum, 1.7—2.5

230 Gardens’ Bulletin — XXVI (1973)

Fig. 1. Nepenthes rhombicaulis Kurata A. upper portion of a climbing stem (x 0.5); B. male

inflorescence (x 0.5); C. female inflorescence (x 0.5); D. rosette leaves with pitchers

(x 0.5); E. male flower (x 5); F. capsules (x 1).

Nepenthes from Borneo, Singapore and Sumatra 231

cm latum, facie inferiore prope apicem appendiculatum; calcar ca. 5 mm longum,

filiforme; ascidia superiora ignota. Inflorescentia 30-40 cm longa, racemosa,

pedunculus 15-20 cm longus, glaber, pedicelli 10-15 mm longi, biflori; sepala 4,

elliptica, 4 mm longa, 3 mm lata, extus minute tomentosa; columna staminae

4 mm longa, antherae 8, uniseriatae; capsula 20-25 mm longa, 5 mm lata, fusi-

formis, brunneo-nitida, leviter puberula, valvae lanceolatae.

Stem climbing up to 20 m high, the part with adult leaves 5-10 mm thick,

quadrate or rhomboid, glabrous; often short shoots and rosettes at the base of

older plants. Leaves of the rosettes and short shoots scattered, the lamina small,

several cm long, lanceolate; Leaves of the climbing stem scattered, herbaceous,

sessile, lanceolate, 12-22 cm long, 3-4 cm broad, the base 4—% amplexicaul; both

surfaces glabrous, the margin ferrugineous pubescent; the upper surface lucid-

green, the under surface yellow-green; longitudinal nerves 2-3 on both sides,

pinnate nerves running obliquely ascending; tendril slender, 10-13 cm long, cylin-

drical, yellow-green, glabrous. Pitchers of the rosettes and the short shoots coria-

ceous, 6-12 cm high, 2.5-3.5 cm broad, ventricose in the lower part, cylindrical

in the upper part, with 2 wings, the margins of the wings toothed; mouth oblique,

circular; peristome subcylindrical, 3-5 mm broad, reddish, outer margin of the

peristome expanded and undulated, interior margin toothed; inner surface of the

pitcher glandular in the lower <_ part; the pitcher red- or pale-green with purple-

spots, sparsely pubescent; lid elliptic-oblong, 1.7—2.5 cm broad, with an appendage

situated nearer to the apex on the under surface; spur filiform, about 5 mm long;

Pitchers of the climbing stem unknown. Inflorescence a raceme, 30-40 cm long,

the peduncle 15-20 cm long, glabrous; pedicels 10-15 mm long, 2-flowered, sepals

4, elliptical, 4 mm long, 3 mm broad, the outer surface minute tomentose; staminal

column 4 mm long, the anthers 8, uniseriate; capsules fusiform, 20-25 mm long,

5 mm broad, bright-brown, slightly pubescent; valve lanceolate.

Typus: §. Kurata 4300 [male] and 430] [female] (Herb. Nippon Dental

College, Tokyo); Mt. Pangulubao near Perapat, North Sumatra, 1700-1900 m alt.,

March 29, 1972.

This species has a peculiar stem with four sharp-angled ridges; two of them

are connected with elongated mid-ribs of two leaves while the remaining two

are connected with leaf margins. As a result of such characters the cross-

section of the stem is either rhomboid or square depending on the level of the

stem sectioned. The pitchers are soft and breakable and are found only on the

rosette leaves and those of the short shoots. The pitchers do not seem to arise from

the leaves of the climbing stem. The rosette leaves and those of the short shoots are

sometimes coloured with purple-green and fringed with distinct reddish-brown hairs

at the margins.

This plant is comparatively common in the subalpine forest of the western

slope of Mt. Pangulubao and its distribution seems widely scattered on the moun-

tains around Lake Toba.

12. Nepenthes singalana Becc., Malesia III, p. 4 & 12, t. IIL (1886).

SUMATRA: West Sumatra, Mt. Singgalang, 2400-2600 m alt., Kurata & Mikil

4207, 4208, 4209, 4210; Mt. Merapi, 2500 m alt., Kurata & Mikil 4211—a, 4211-b.

Distribution. Sumatra.

232 Gardens’ Bulletin — XXVI (1973)

13. Nepenthes tobaica Dans., Bull. Jard. Bot. Buitenz ser. III, Vol. IX, p. 382

(1928).

SUMATRA: North Sumatra, near Perapat, Lake Toba, 1000 m alt., Kurata

& Mikil 4266, 4267, 4268, 4269, 4270, 4271, 4272; Mt. Pangulubao, 1500-1700 m

alt., Kurata & Mikil 4273, 4274, 4275, 4276, 4277, 4278, 4279, 4280; Mt. Sibajak,

1800 m alt., Kurata & Mikil 4286, 4287, 4288, 4289, 4290, 4291, 4292, 4293.

Distribution. Sumatra.

Addendum to NEPENTHES CAMPANULATA

A special expedition was organised by Dr. A. J. G. H. Kostermans in 1957 to

Mount Ilas Bungaan, an area in N. E. Kalimantan. Kostermans was informed by

Dayaks that the rock was named such because it was completely overgrown with

flowers (ilas = rock, bungaan = flowering). It took him 14 days to travel through

unknown country to reach that rock. The steep 300-ft rockface was completely

covered by a yellowish coloured NEPENTHES here described as a new species.

Moreover, when climbing the rockface with cane-ropes, antique rock graves

were found with well preserved wooden carved coffins, still containing skeletons

of which some were headless.

Annotated list of seed plants of Singapore (I)

HSUAN KENG

Department of Botany

University of Singapore

Introduction

The first list of plants of Singapore and adjacent islets, entitled ““The flora of

Singapore”, was compiled by H. N. Ridley in 1900, with supplementary notes

appearing a year later. These were published in the Journal of the Royal Asiatic

Society, Straits Branch, volumes 33 and 34. Subsequently, two separate lists,

entitled “List of Dicotyledons in Syonan” and “List of Monocotyledons in

Syonan”’, were compiled by the Botanic Gardens staff during the Japanese occu-

pation (1942-45), but were never published. Only their typed manuscripts were

preserved in the Botanic Gardens, Singapore.

The present annotated list is essentially based on specimens accumulated in

the herbarium of the Botanic Gardens, on Ridley’s Flora of the Malay Peninsula

(1922-25), on the lists mentioned above and on my own notes. In addition,

numerous monographs, revisions and short articles were consulted. It is my intention

to bring the nomenclature of the plant names up to date as far as possible, and

admittedly, a number of them may have been overlooked. In some cases, I

exercised my own judgement in accepting or rejecting some of the latest nomen-

clatural changes.

For convenience, the arrangement of the families follows that given in my

book “‘Orders and families of Malayan Seed Plants” (University of Malaya Press

in 1969, Kuala Lumpur).

With rapid urbanization and industrialization, a large number of native plants

have already vanished or are on the verge of disappearance. On the other hand,

however, a number of exotic plants were introduced, and some are already well-

established. To differentiate them in the list, native species are printed in bold

face, while introduced ones are in italics. Only native species bear a collection

number of the herbarium specimen (often historical). Where available vernacular

names in Malay/Chinese are supplied. It must be understood that the list of cultiva-

ted plants can not be expected to be complete. For example, among Gymnosperms,

it was recorded in the “Index of Plants” of 1912 by J. W. Anderson, that several

cycadaceous plants such as Bowenia, Ceratozamia, Stangeria, and coniferous plants

such as Cephalotaxus, Sciadopitys, T hujopsis, etc. were growing in Singapore. None

of these have survived, the main reason being that, under local climatic conditions,

many of these introduced plants do not produce cones or flowers, let alone viable

seeds. The perpetuation of such species would then depend on their ability to

propagate by vegetative means or on the recurring introduction of seeds and plants.

The list will be published in instalments, with the intention of eventually

combining them into one volume. I would like to express my appreciation to the

Director and staff of the Botanic Gardens, Singapore, for their willing cooperation

and assistance in the preparation of this list.

233

234 Gardens’ Bulletin — XXVI (1973)

I. Gymnosperms

1. Cycadaceae

Artificial Key to the Genera

A. Leaflets with a midrib

A. Leaflets without a midrib.

B. Leaflets spinescent.

C. Spines minute, to 0.25 cm long

C. Spines conspicuous, to 1 cm long

B. Leaflets entire.

D. Leaflets 15-20 cm long, not jointed

D. Leaflets 8-10 cm long; jointed at base

Cycas revoluta Thunb.

Cycas

Dioon

Encephalartos

Macrozamia

Zamia

A dwarf cycad, with strongly curved or ‘revolute’ leaflets; introduced from S.

China and Japan; often cultivated; sometimes producing cones. 7

Cycas rumphii Miq.

Formerly found wild on sandy shores at Changi (Ridley 4408), Tempinis, and

Tuas, now survived only in cultivation. Dwarf palm-like; stem thick-columnar,

rarely branched. Probably not specifically different from C. circinalis Roxb. of

India and the Pacific Islands. Vern. Pakis laut, Paku raja, § TERRE

Cycas siamensis Miq.

A native of N. Malaya and Thailand; base of stem bulb-shaped: cultivated

occasionally; producing cones. BAKE

Dioon spinulosum Dyer

Native of Mexico; sterile.

Encephalartos villosus Lem.

Native of S. Africa, occasionally producing cones.

Macrozamia denisonii Moore & F. Muell. var. hopei Schust.

Native of E. & N. E. Australia, occasionally producing cones.

Zamia floridana A. DC.

Native of Florida, U.S.A., often producing cones. jk FE

Zamia media Jacq. var. gutierrezii Schust.

Also known as Z. integrifolia Hort.; native of Cuba, sometimes producing

cones.

Zamia media Jacq. var. tenuis Schust.

Also known as Z. tenuis Willd.; native of Bahamas, sometimes producing

cones.

Annotated list of seed plants of Singapore (1) 235

2. Podocarpaceae

Artificial Key to the Genera from Podocarpaceae to Gnetaceae (adapted from

Corner’s ‘Wayside Trees” and modified).

A. Leaves flat, 0.5-10 cm wide, not needle-like.

B. Leaves opposite, usually spreading in one plane; twigs with distinct internodes.

C. Leaves thin, net-veined with a midrib; nodes swollen Gnetum

C. Leaves thick, stiff, faintly parallel-veined; nodes not swollen.

D. Terminal buds blunt; resin copious Agathis

D. Terminal buds pointed; resin scant Podocarpus (in part)

B. Leaves alternately or spirally arranged; internodes always very short.

E. Leave sharply pointed, without a midrib Araucaria (in part)

E. Leaves not prickly, with a distinct midrib Podocarpus (in part)

A. Leaves needle- or scale-like, 2 mm wide or less.

F, Needless over 15 cm long, 2 or 3 in a bundle Pinus

F. Leaves less than 2 cm long, generally much less; not in bundles.

G. Plants with 2 kinds of leaves: some twigs with minute scale-leaves, others with

longer needle-leaves.

H. Leaves in 4 rows (or in two, alternating pairs) Juniperus

H. Leaves spirally or alternately arranged.

I. Needle-leaves flat, alternate, twisting in one plane; scale-leaves with

projecting points Podocarpus (imbricatus)

I. Needle-leaves not flat; scale-leaves not with projecting points.

Dacrydium

G. Aduit plants with only one kind of leaves, either needle- or scale-shaped.

J. Needle-leaves 1-2 cm long, spirally arranged, with projecting points.

K. Branches in whorls Araucaria

K. Branches not in whorls Cry ptomeria

J. Scale-leaves 2 mm long or less.

L. Small branches with their leaves flattened into one plane Thuja

L. Otherwise Cupressus, Calocedrus, Callitris.

Dacrydium elatum Wall.

Native of Malayan mountains and elsewhere; remaining sterile; leaves of two

forms: needle-shaped and scale-like, often on separate branches. jR44

Podocarpus imbricatus B1.

Native cf Malayan mountains and elsewhere; producing male ones only.

JT EE He WE HL

Podocarpus motleyi Dummer

Native of Malayan swamps and elsewhere.

Podo. neriifolius D. Don!

Native of Malayan mountains and elsewhere; producing cones and seed. § H

Podo. polvstachyus R. Br.

A native species, found in Labrador, Kranji (Ridley 13304) and other parts

of the island near the sea and in mangroves; also growing in gardens. Vern.

Sintada.

Podo. wallichianus Presl

Collected at Changi and Jurong (Corner s.n. Apr. 1933); erroneously known

as Podo. blumei Endl.; commonly found in Malayan mountains. 4 #£ 4; #4

1. Several other species of Podocarpus from China, Indonesia and elsewhere are intro-

duced, some even producing viable seeds.

236 Gardens’ Bulletin — X XVI (1973)

3. Araucariaceae

Agathis dammara (Lamb.) L. Rich.

Native of Malayan mountains and elsewhere; cultivated; producing cones,

occasionally viable seeds. A small patch of trees on the back of Raffles Hall,

Univ. of Singapore. Variously called A. loranthifolia Salisb. or A. alba Saff.

in literature. Vern. Damar minyak, ARgh

Araucaria bidwilii Hook.

Introduced from Queensland; Australia. Leaves lanceolate, 2-rowed; known

as Monkey puzzle.

Arau. cunninghamii Sweet

Native of Australia and New Guinea. Needle in dense spirals, sharp; known

as Hoop pine.

Arau. excelsa R. Br.

Native of Norfolk Island, New Zealand. Needles in dense spirals, blunt. Known

as Norfoik I. pine; propagated by cutting; one of the commonest Araucaria in

cultivation. —§9 7##

Arau. kunsteinii K. Soh.

Native of New Guinea, Known as Kiing pine.

4. Pinaceae

Pinus merkusii Jungh. & De Vriese*

Native of Thailand, Cambodia and Elsewhere. Producing cones and occasion-

ally viable seeds. KEK ,

5. Taxodiaceae

Cryptomeria japonica D. Don

Native to S. China and Japan; sterile, rarely producing male cones. ji) #%

6. Cupressaceae

Callitris macleayana Muell.

Possibly together with one or two other species, is cultivated; all from

Australia; sterile. jA}H #4

Calocedrus formosana Florin

Formerly known as Libocedrus formosana Florin; native to Formosa; occa-

sionally producing male cones. }4 fiij

2. Several other species of pines, such as P. insularis Endl. (3-needled, from the Philip-

pines), P. caribaea Morelet (3- or 2-needled, from the W. Indies), P. massoniana Lamb.

(2-needled, from S. China) are sometimes cultivated; often remain sterile, rarely producing

male cones.

Annotated list of seed plants of Singapore (1) 237

Cupressus macrocarpa Hort. ex Gordon

Native to N. America; sterile. K# #47

Juniperus chinensis L.

Native of China; several horticultural forms commonly cultivated; sterile or

producing male cones only; propagated by cutting. [B}}4

Thuja orientalis L.

Native of N. China; commonly cultivated in gardens; often producing male

cones; propagated by cutting. {fl|4

7. Gnetaceae

Gnetum gnemonoides Brongn.

Listed in Ridley’s Flora as G. wrayi Gamble. Recorded from Tuas, Jurong,

Chua Chu Kang (Ridley 6126), etc. Seeds large, 5-7 cm long.

Gnetum gnemon L.

Shrub or small tree (in contrast to other species listed here being woody

climbers) easily mistaken as a dicot. It is either a native or at least was

introduced during the immemorable past. Usually only the female plants are

cultivated near Malay villages (Ridley 6106) because of the edible seeds

(erroneously called as “nuts”). Vern. Meninjau or Belinjau, kif{ii

Gnetum latifolium BI. var. funiculare MGF.

Known as G. kingianum Gamble in Ridley’s Flora. Recorded from Garden

Jungle, Tanglin (Ridley 5688) Kranji. A big climber, the bark used for making

string; leaves large, elliptic. Vern. Akar tutubo, Akar susurus.

Gnet. macrostachyum Hook.

Recorded from Siglap and Changi (Ridley 4822); leaves elliptic; seeds nearly

globose.

Gnet. microcarpum BI.

Formerly known separately as G. campestre Gamb. and G. sylvestris Gamb.

Recorded from Tanglin, Kranji, Garden jungle, Seletar (Ridley 3851), etc. A

large climber; seed golden yellow; leaves narrow; relatively common. Vern.

Akar jullah, (if

Ree

Pi fenrt hl ids « dancnaura (Cand) b4

Native of Vaiaiee cotiae ~F ¢!

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10 litle folevittlis: hie yi

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ns Hoop pide. . hat Gad on

Squaw \adeT Aindit. bohnovest sldrast now D 28 & ——

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Korthalsia hispida Becc. in Malaya

J. DRANSFIELD

Herbarium Bogoriense, Bogor

Indonesia

On 7th January 1973, while in Singapore, I accompanied Dr. E. A. Heaslett

on a one day field trip to Gunung Panti in Johore, Malay Peninsula. Despite bad

weather, we were able to observe about 55 taxa of Palmae in the forest of Panti

Fast. This total indicates the richness of the palm flora: amongst those species

observed was one novelty, a Korthalsia vegetatively distinct from all other Malayan

species of Korthalsia.

Within the genus Korthalsia in Malaya (Furtado 1951) there are two types of

ochrea (extension of the leaf sheath beyond the insertion of the petiole); in one

type such as that found in K. scaphigera Griff., K. echinometra Becc. and K.

scortechinii Becc. (see fig 1 a, b) the ochrea is swollen and hollow and forms a

spiny chamber around the leaf sheath of the leaf above, and is favoured by ants

as a nesting place. The relationship of the ants with the rattan has been described

many times, and it seems likely that particular ants are specific to different Korthal-

sia species. In the other type as found in all other Malayan species of Korthalsia

such as K. flagellaris Miq and K. rigida Bl. (see fig 1c, d), the ochrea is not

swollen and forms a tightly sheathing tube around the leaf sheath of the leaf above:

this type of ochrea may remain entire or becomes split or tattered, and may be

armed or unarmed.

The Korthalsia species new to the Malay Peninsula from Panti East Forest

Reserve, however, has a third type of ochrea, a type found in four species (see

fig 1 e): K. robusta Bl. of Sumatra. K. macrocarpa Becc. of Borneo, K. squarrosa

Becc. of the Philippines and K. hispida Becc. of Sumatra and Borneo. In these

four species, the ochrea is shaped something “like an elongated ass’s ear” (Beccari

1918): the ochrea is not sheathing, being open opposite the petiole, the two edges

inrolling to produce an open-ended spiny tube, pale straw in colour, diverging from

the axis at an angle of about 20 degrees. In K. squarrosa the ochreas are slightly

different in that they do not diverge far from the stem, but the ochreas visible in

Beccari’s plate (Beccari loc. cit.) are just below an inflorescence and such ochreas

are often slightly abnormal. Within this tube-like ochrea in the Sumatran and

Bornean species, ants make their nests. When the rattan is touched, the ants within

rustle in unison in each ochrea by banging their heads against the dry ochrea, each

ochrea slightly out of phase with the next. I have only heard this rustling noise

in K. robusta, K. hispida and K. macrocarpa; I have not heard it in K. scaphigera

or K. echinometra and it is quite likely that the ant species in these last two are

different.

239

240 Gardens’ Bulletin — XXVI (1973)

The Gunung Panti plant is indistinguishable from Korthalsia hispida, a mis-

understood species from West Sumatra, Aceh, and Borneo, first collected by

Beccari (P. S. 673 — Beccari 1884) at Ayer Mancur, West Sumatra, and later

included by Beccari in Korthalsia robusta Bl. non Mart. (Beccari 1918). In July

1972 in Propinsi Jambi, Sumatra, (Dransfield in press) I found specimens of a

Korthalsia, vegetatively fitting the description and plate of K. hispida but with

inflorescences totally different from those of the widespread Sumatran taxon, K.

robusta, the vegetative parts of which were unknown to Beccari. The Jambi

specimens are identical to Meijer 2411, Nunukan, Borneo (Meijer 1956). There are

hence two species in Sumatra, K. robusta and K. hispida, and Beccari appears to

have confused the two in his monograph (1918) — the fertile specimens referred

by Beccari to K. robusta belong to K. robusta; the vegetative specimens referred

to K. robusta belong to another species originally described as K. hispida. In fact

vegetatively K. hispida is more distinct than K. robusta and K. macrocarpa and it

is quite possible that these last two represent one widespread variable species

common to Sumatra and Borneo.

Because of the confusion in this group of Korthalsia it is considered useful to

give a complete description of Korthalsia hispida. Although the relationship of

Korthalsia macrocarpa and Korthalsia robusta is not yet clear, I am presenting

this note in order to draw attention to this easily recognized group of Korthalsia

species.

Korthalsia hispida Becc.

Beccari, in Malesia //, (1884) 71., Beccari, Ann. Roy. Bot. Gard. Calc. XII, 3

(1918) 148. Plate 99. Meijer, W. Penggemar Alam 36 (1957) 60, Fig 3.

Slender, clustering, thicket-forming rattan, branching at ground-level. Stems

to 1.0 cm in diam. without sheaths, with sheaths 1.5—1.8 cm. Internodes 20-25 cm

long. Sheaths bright shiny green when fresh, covered with sparse, scattered, shiny

black spines to 2.0 cm long, easily breaking just above the base and minute c.

0.5 mm long, easily detached spiculae. When young, sheath covered with chocolate

coloured scurfy indumentum. Ochrea 25-30 cm long, like a very narrow elongate

horn, diverging at an angle of c. 20 degrees from the axis, shiny brown within,

straw-coloured without, thorny-spiculate and indumentose as the leaf sheath,

truncate and shallowly bilobed at the apex, papery in texture, open opposite the

petiole, rolled, often filled with vicious biting ants. Leaf to 1.8 m long: cirrus to

90 cm, petiole to 10 cm. Petiole and rachis yellowish-green, semi-circular in cross

section, to 5 mm wide, armed with scattered, reflexed black spines below, above,

spines in groups 3-5 forming claws, separated by c. 3 cm, distance of separation

decreasing above to c. 5 mm at tip of cirrus, Leaflets 7 on each side, alternate

below, subopposite above, separated by 7-10 cm, with ansae 6 mm long, the lower

two leaflets narrowly cuneate-rhomboid, to 3 cm broad, upper leaflets broader,

15-20 cm long by 7-9 cm broad, irregularly plicate, longly acuminate praemorse

at upper margins, bright green above, grey-white indumentose below. Leaves sub-

tending the terminal inflorescence much smaller.

Inflorescence system lax, terminal, single stems dying after flowering. the

upper 3-8 nodes producing partial inflorescences. Partial inflorescences to 30 cm

long, emerging through splits in the upper leaf sheaths; prophylls and other bracts

tubular below, variously split and tattered above, 5-10 cm long, straw-coloured,

papery, densely covered in chocolate-coloured indumentum and small spiculae.

Bracts on axis subtending branches, each terminating in rhachillae, 2-6 in number.

241

Korthalsia hispida in Malaya

10cm

OCHREA TYPES IN KORTHALSIA

Fig. 1

on-

ida,

piny, d) K. sp. (? juvenile of K. rigi

shaped ochrea:

b) K. echinometra. N.B. hole made by ants. N

, Central Sumatra) — ochrea unarmed. Horn-

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fe)

2) [aa]

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sig

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53.

nASHM

242 Gardens’ Bulletin — X XVI (1973)

Rhachillae pendulous, to 15 cm long, 1.2 cm in diam, clothed in a tight spiral of

bracts, adnate laterally, each bract subtending a condensed branching system with

one hermaphrodite flower. Bracts to 1 cm wide and 5 mm high, pale brown with

deeper brown fugaceous scurfy indumentum, irregularly split and lobed at the

apex. Flowers pale brown in colour, exserted between the tight bracts. Sepals 3,

free, imbricate, ovate, shiny, straw-coloured, to 3 mm long. Corolla with a tube

c. 3 mm long and 3 free imbricate rounded triangular lobes above, 4 mm long

and 3.5 mm wide, during growth of the ovary after fertilization, the corolla tube

splitting and being carried out of the bract axils on the tip of the ovary, at the

base of the style. Anthers 6, 2 x 1 mm, borne on a short staminal tube, this shortly

epipetalous. Pollen pale yellow. Ovary at anthesis, to 2 mm high, with style to

4 mm, chestnut brown, scaly. Style grooved longitudinally and tipped with 3

minute approximate stigmatic lobes. Fruit when not damaged crowned with remains

of style, surrounded by corolla and stamens, at least 1.2 cm long, and 1 cm wide.

Mature fruit unknown.

Specimens examined

Type not seen, though Plate thereof in Beccari (1918) examined.

SUMATRA

Aceh: Nainggolan s.n. April 1931, Bukit Plawi (BO).

Jambi: Dransfield 2620. 22.7.72. Kampung Penetai, Kabupaten Kerinci,

streamside, Hill Dipterocarp Forest, alt 200m (BO, L, BH).

BORNEO

East Kalimantan: Meijer 2411. 5.12.53. Nunukan, near S. Binusan.

dry lowland Dipterocarp Forest (BO).

MALAY PENINSULA

Johore: Dransfield 3037. 7.1.73. Gunung Panti East. Lowland,

reshwater swamp forest. (SING, BO, L, BH).

Korthalsia hispida differs from the other species having long horn-like ochreas,

in the hispid appearance of the sheaths, ochreas and inflorescence bracts, produced

by the dense minute spiculae, and in the lax, not approximate, slender infiorescence

branches. K. macrocarpa, K. robusta, and K. squarrosa all have squat inflorescences

made up of robust, fat, approximate rhachillae, unlike any other Korthalsia species

(see Beccari 1918).

The presence of this group of long-ochrea Korthalsia species in Malaya is not

surprising; the absence of collections of it, is, especially as it grows in a very

famous, much-visited botanical locality.

Acknowledgements

I should like to thank Dr. E. A. Heaslett for introducing me to Gunung Panti East,

and Ahmad Shukur and Samsuri, plant collectors, Botanic Gardens, Singapore for

their help in collecting palms on Gunung Panti: thanks are also due to Sdr.

Damhuri, draughtsman, Herbarium Bogoriense, for preparing the figures.

Korthalsia hispida in Malaya

BAKO CARELESS

0) aay) ae ee Ae ~ ad) in »S

Pesan pics eR SN on

vig sae

SOE SAK ALA cy

RANE Deas Wek Ph aw

Ves, “sr¢ A Py > Ae

ella eat

Fig. 2. KORTHALSIA HISPIDA BECC.

243

Illustration of a node and most of one internode and the subtended partial inflore-

scence, taken from near the apex of a flowering stem; in all 3, nodes were producing

partial inflorescences. N.B. the hispidity on all sheaths, ochreas, and bracts, Dransfield

2620, Jambi, Sumatra. Drawing by Damhuri.

244 Gardens’ Bulletin — X XVI (1973)

References

Beccari, O. (1886): Piante Ospitatrici, Malesia 2, 62-78.

Beccari, O. (1918): Asiatic Palms — Lepidocaryceae. Part 3. The Species of the

genera Ceratolobus, Calospatha, Plectocomia, Plectocomiopsis,

Myrialepis, Zalacca, Pigafetta, Korthalsia, Metroxylon, Eugeis-

sona, Ann. R. bot. Gard. Calcutta 12.

Dransfield, J. (in press): Notes on the Palm Flora of central Sumatra. Reinwardtia

& (4).

Furtado, C.X. (1953): Palmae Malesicae 11. Korthalsia. Gdns’ Bull., Singapore 13

300-324.

Meijer, W. (1957): Rotans van Nunukan. Penggemar Alam 36 51-64.

Pollen Viability and Germination in Some Orchid Hybrids

A. N. RAo and LEE War CHIN*

Department of Botany

University of Singapore

Introduction

Singapore has become an important orchid breeding center producing many

well-known hybrids of international acclaim. Such successful results are achieved

by the continued efforts of scientists at Botanic Gardens and local horticulturists

over a period of three decades. In Singapore the first hybrid that appeared was

Vanda Miss Joaquim in 1893 (Holttum, 1953). Nevertheless, our knowledge about

the cytogenetics and pollen physiology of local orchid hybrids is very limited. The

importance of such studies need not be over-emphasized since pollen viability

and germination form the essential requirements to produce good and the required

hybrids. Not every cross pollination will bear fruit and it is the common experience

of every orchid breeder that more than 60 per cent of the pollinations made would

end up with no fruits or fruits without seeds. One of the main reasons for such

failures could be the pollen sterility and some of these problems are under study

(Rao and Goh, 1970).

Very few studies are made on orchid pollen germination. Molisch determined

the optimal sucrose concentration for 6 European species (Withner, 1959). Miwa

(1937) germinated the pollen of certain orchid species and their hybrids to test

the longevity and viability of stored orchid pollen. Curtis and Duncan (1947)

studied 11 species in 7 genera. The present paper outlines the morphology of

pollinia, pollen structure, viability and germination of some well-known local

hydrids.

Materials and Methods

Pollinia of the following hybrids were collected from Mandai Gardens now

known as Singapore Orchids Pte. Ltd.: (i) Arachnis Maggie Oei var. Red Ribbon,

(ii) Arachnis Maggie Oei var. Yellow Ribbon, (iii) Aranda Hilda Galistan, (iv) A.

Lucy Laycock (long spray), (v) A. Lucy Laycock (short spray), (vi) A. Wendy

Scott var. Greenfield, (vii) Aranthera Lilliput, (viii) Vanda Poepoe and (xi) Vanda

Josephine (from the garden at Botany Department, University of Singapore).

Fresh pollinia from open flowers were collected between 2-3 p.m. and used

within 1 hr. after collection, using half a pollinium in each case.

The following culture media were used: (a) distilled water as the control

medium (5) stigmatic extract — the appropriate number of fresh stigmas were

ground in 10 mls. of distilled water and the clear filtrate of the mixture was then

used for germination. The number of stigmas used in such preparation varied from

* Currently at Botanic Gardens, Singapore.

245

Pollen Viability and Germination in some Orchid Hybrids 247

Figures 19-25: Pollen germination in Vanda Josephine. Fig. 19: in dist. water, note

ihe short tubes. Fig. 20: in stigmatic medium at 2 stigmas/10 mi of dist. water; all 4 grains

in the tetrad germinated. Fig. 21: in 4% sucrose a & b different sized pollen tubes with

tapering ends. Fig. 22: in salt-boron medium, a— pollen tubes typical match-stick shaped,

some with vacuoles; b — short tube, with broad ends. Fig. 23: in GA (10-7 mg/L) medium

short pollen tubes with exudates (arrow). Fig. 24: in borax medium, with all the grains in

tetrad germinated. Fig. 25: in kinetin (10-6mg/L) medium with pollen exudate (arrow),

See opposite page

Figures 1-18. Figs. 1-9: Pollinia of different orchid hybrids. Figs. | & 2: Arachnis

Maggie Oei var Red Ribbon and var. Yellow Ribbon respectively. Figs. 3-6: Aranda Hilda

Galistan; Lucy Laycock (long spray); Lucy Laycock (short spray); Wendy Scott var. Greenfield

respectively. Fig. 7: Aranthera Lilliput. Fig. 8: Vanda Poepoe. Fig. 9: Vanda Josephine.

a-column, b-} stigma, c-pollinia. Figs. 10-18: Tetrads showing microspore arrangement and

supernumerary spores. Fig. 10: Arachnis Maggie Oei var Red Ribbon, Fig. 11: A. Maggie

Oei var. Yellow Ribbon. Fig. 12: Aranda Hilda Galistan. Fig. 13: A. Lucy Laycock (long

spray), Fig. 14: A Lucy Laycock (short spray). Fig. 15: A. Wendy Scott var. Greenfield. Fig

16: Aranthera Lilliput, Fig. 17: Wanda Poepoe. Fig. 18: Vanda Josephine.

248 Gardens’ Bulletin— XXVI (1973)

3, 1, 2, 3 stigmas/10 mls. of distilled water. In some cases all the four concentrations

were used, in others only 1 or 2. The total length of the stigmas ranged from 3-8

mm in different hybrids. Their sizes and weights are:

Length of Wt. of

Plants stigma I stigma

(in mm) (in mg)

Arachnis Maggie Oei var. Red Ribbon 8 198

A. Maggie Oei var. Yellow Ribbon 8 180

Aranda Hilda Galistan 5 101

A. Lucy Laycock (long spray) 5 96

A. Lucy Laycock (short spray) 5 109

A. Wendy Scott var. Greenfield 6 87

Aranthera Lilliput 3 16

Vanda Poepoe 8 84

V. Josephine 8 85

(d) Borax medium — Sodium tetraborate was used to prepare the media with

concentration ranging from 10° to 10 mg./L.

(e) Salt boron medium (modified Knop’s medium). Formula: H;BO,; — 100,

Ca(NO;)2.H,O0 — 300, MgSO,. 7H,O — 200, KNO; — 100mg/L.

(f) Different growth substances — well known auxins like: Indole — 3 — Acetic

Acid (IAA), Indole — 2 — Butyric Acid (IBA), Indole — 3 — Propionic Acid

(IPA), o¢ — Naphthalene Acetic Acid (NAA), 2,4 — Dichlorophenoxy acetic

acid (2,4-D), 2,45 — Trichlorophenoxy acetic acid (2,4,5-ITPA), source of

gibberellin Gibberellic Acid K salt (GA), and source of Kinin — Kinetin (K)

were used in the concentrations that ranged from 10° to 10 mg/L. Either all or

only certain concentrations were found to be effective.

Solid watch glasses (4 x 4 x 1.75 cm) were used to germinate pollinia. The

pollinia were cut into small pieces and dispersed in | ml of the medium. A glass

cover was used to prevent evaporation of the medium and the watch glass was

placed in a humid chamber to prevent evaporation and condensation. Diffuse light

of 235 F. C. S. (during day-time) and temperature of 28°C prevailed in the labora-

tory. Duration of experiment was limited to 24 hours since most pollen did not

germinate or their pollen tubes did not grow after this period. Each experiment

was repeated at least twice.

At the end of 24 hours, the percentage germination of pollen tetrads and

pollen cells, pollen tube length and morphology were studied. Lactophenol with

cotton blue was used for staining. At least 225 tetrads were studied for each

treatment to assess the percentage germination and 90 tubes for tube length

determination.

Observations and Results

Pollinia and pollen grains: In orchids the pollen tetrads develop into mature

structures called compound pollen grains and these are united to form pollinia. In

all of them two pollinia are present per flower (Figs. 1-9). The pollinia of different

hybrids were excised for comparative study. In size the Aranthera pollinium was

smallest among the 9 hybrids studied (Fig. 7). Pollinia of Aranda hybrids were

smaller than those of Arachnis or Vanda (Figs. 1-6, 8, 9). The disc was flat in

Arachnis, triangular in Aranda, and of irregular shape in Vanda. Even among the

Pollen Viability and Germination in some Orchid Hybrids 249

Table 1. Morphology of the pollen tetrads, size and arrangement of pollen cells in them.

Morphology of Pollen Tetrads

Nittinos of Orchids . Size, ‘ica he grains in tetrads

isobi- tetra- .

i SLB lateral. | hedeal T-shaped) linear

Arachnis Maggie Oei var. Red Ribbon... | 37 x 23 1.61 92 6 1 1

Arachnis Maggie Oei var. Yellow Ribbon | 40 x 27 1.48 90 8 zZ —

Aranda Hilda Galistan i! - i |e oy 2&5 dee 82 16 — ys

A. Lucy Laycock (long spray) can Sa 2a TAZ 72 Ze 2 4

A. Lucy Laycock (short spray) oe Loe Ok oe ee 88 8 2 2

A. Wendy Scott var. Greenfield vets & 22 eS Most of them are supernumerary

spores. Normal tetrad — _ only

about 1%

Aranthera Lilliput “t ye lA ee 20 Sua 89 9 1 1

Vanda Poepoe .. a ere ~~ 2 b6 68 32 1 1

Vanda Josephine «t Re ee aes ay 72 ad 1 oe

L — Length of pollen tetrad

B — Breadth of pollen tetrad

L/B — Length /breadth ratio.

Aranda hybrids the pollinia of each hybrid were distinctly different from the others

in having disks and stalks of different shape and size (Figs. 3-6). The pollinia

were teased to separate the tetrads. These are diagrammatically represented in

Figs. 10-18. Each figure shows the variation in size, shape, arrangement as well as

the number of cells in each tetrad or the unit (is one which had more than 4 cells).

In addition, the cells of the tetrad or unit also varied in size. Some of them e.g.

Vanda Miss Joaquim, Vanda Poepoe, Aranthera Lilliput had one small and three

big grains or vice-versa (Figs. 14, 16-18, 22, 28, 29). Two big and two small grains

were also common in some (Figs. 13, 22). Supernumerary spores were also common

in many (Figs. 12-14, 16, 18) and the individual unit had 5, 6 or 8 cells. In

such units, which had supernumerary spores, at least two of them were big and

of comparable size with the spores of the majority of the tetrads and the remaining

were generally small (Figs. 12, 13, 18). Pollen tetrads show isobilateral, tetrahedral,

linear and ‘T’ shaped arrangement. The percentage occurrence of each type is

recorded in Table 1. Comparatively, the two latter conditions were rare and the

isobilateral type was most common. The average size of pollen tetrad ranged

from 28 x 28 » to 42 x 27 uw. The linear tetrads were much smaller in size when

compared with the others (Figs. 12, 14, 16).

Pollen germination: Positive responses resulted in the pollen germination of

Vanda hybrids and these details will be considered first, followed by the results

of other hybrids.

(a) Distilled water (control medium). In V. Poepoe and V. Miss Joaquim 27 and

30% of the tetrads germinated with an average tube length of 15 and 19 yp

respectively (Tables 2, 3; Figs. 19, 26, 30, 31).

250

Table 2.

Gardens’ Bulletin — XXVI (1973)

different media after 24 hrs. at room temperature.

No. of grains germinated per

tetrad, in %

Vanda Poepoe. Percentage germination and average pollen tube length in

Media Concentrations %G pL

1 2 3. jattat

Distilled water 18 5 2 2 27 15

(control)

| 1 stigma /10 mis. 20 21 14 10 65 57

distilled H2O

Stigmatic 2 stigmas /10 mls.| 27 30 13 13 83 68

extract distilled H2O

ee “We | =s

3 stigmas /10 mls.| 21 20 8 4 53 44

distilled H20

pe 28 20 4 12 64 56

at 23 25 24 13 85 75 :

Sucrose 6% 8 16 24 41 89 83

8% 15 11 12 | 12 50 | 48

Le) eee seh? oe ia at | tthe

10% 11 9 | 11 8 49 40

1x 10—5mg./L| 26 6 2 BO paige bh ag

atl iw 320 fi Hiawft 29 pial salle aoe

1x 10—4mg./L 20 12 2 2 | ey pees

'Yivo Soe ee} SAAS Gee oh ee | /

1x 10—3 mg./L 27 10 0 2 39 | 36

Borax 1x 10—2mg./L 20 14 4 2 40 36

1x 10—1mg./L 2a 12 8 2 47 38

1x 10—° mg./L 24 16 8 13 50 55

1x 10—! mg. /L 16 14 8 z | 40 37

Salt-boron medium ZZ 32 10 8 he 72

% G — Perc. germination

L — Avg. pollen tube length in y,

Pollen Viability and Germination in some Orchid Hybrids 251

Table 3. Vanda Poepoe. Percentage germination and average pollen tube length in optimal

concentrations of different media, after 24 hrs. at room temperature.

No. of cade: 3 per

E etrad, in |

Media PY le a : 70G pl

| 1 2 3 4

Distilled water | 18 5 2 2 ZY 15

(control) 2

TAA 11x 10—6 mg. IL| My We td | taal nk 71 71

IBA \1x 10—5mg./L| 21 25 «| 45 es es hls

IPA 1x 10—6mg./L| 24 17 iaab 3 ss | al

NAA 1x 10—4mg. 1 10-4 mg. JL 27 26 20 5 78 82

2,4-D fins SED i 10—6mg./L| 31 | 21 15 ala 60

2,4,5-TPA lay 1x t0—Sime/L |. 27 | 25 6 7 65 oll 40

GA ee TX 0? mall 3g 23 5 7 ia 75

Kinetin fix tae n | 24 15 9 | 7 | 55 * 34 G,

% G — Perc. germination

p. L — Avg. pollen tube length in be

(b) Stigmatic extract medium: For both the hybrids the optimal concentration of

stigmatic extract was 2 stigmas/10 ml. of distilled water. The percentage germination

for V. Poepoe and V. Miss Joaquim was 83% and 79% with an average tube

length of 68 and 87 » respectively and these results were better than in the control

medium (Tables 2, 3; Figs. 20, 27, 33). The response at other concentrations was

variable in terms of percentage germination.

and 89% of the tetrads germinated with an average tube length of 83 » (Fig. 28).

In V. Miss Joaquim 92% of the tetrads germinated in 4% sucrose medium

(optimal concentration) with an average tube length of 74 u (Fig. 21a, b). Other

variations observed in media with higher or lower concentrations are given in

Tables 2, 3 and in all the concentrations used the response was better than in

the control medium.

(zd) Borax medium: In media with different concentrations of borax better results

than control were obtained in V. Poepoe. The optimal concentration for the two

hybrids were 1 and 0.1 mg/L respectively (Figs. 24, 34) and the response of

V. Miss Joaquim was much better than V. Poepoe (Tables 2, 3).

(e) Salt-boron medium: When compared with control or borax medium the values

obtained in this medium were better in the case of V. Poepoe (Table 2). In case

of V. Miss Joaquim the percentage germination was almost like that of control but

tube length improved. These results were poor when compared with those obtained

in optimal concentration of borax medium (Table 3, Fig. 22a, b).

252

Gardens’ Bulletin — XXVI (1973)

Table 4. Vanda Josephine. Percentage germination and pollen tube length in different

media after 24 hrs. at room temperature.

| |

Media Concentration

Distilled H2O

(control)

stigma /10mls.

distilled H2O

wI-

—_

stigma /10mls.

distilled H2O

Stigmatic

extract

2 stigmas /10mls.

distilled H2O

stigmas /10mls.

distilled H2O

Sucrose 6%

ft 10—3 mg. /L

1x 10—2mg./L

Borax 1 x (10—1 mg. /L |

1x 100 aa 21

[Ax 10 mg. /L

No. of grains germinated per

tetrad, in %

Gf el

2 3 4

5 on

| |

5 2 0 36 | 19

11 2-01 x14 | 38 55

9 6 21 | «48 77

| | |

6 20 | | 7

jp YH

| |

in | 24 2 39 | 47

| ea.) ee |

15 | 31 21 75 | 62

8 22 | 7-4-2 | 74

21 14 36 80 | 37

MESSE, A at a8! ae A

1 ‘2 1 | 20 | 21

Je AE oS Eee

0 0 0 11 14

17 | 0 | 1 46 | 27

———__,—___|—____—

8 8 | 0 60 | 41

19 24 | 18 | 85 | 84

| |

13 goin of pu bay | 63

20 OG DONO SNOW Bi? eee

2 1 1 13 | 19

6 5 | 1 38 Se

Salt-boron medium |

% G — Perc. germination

b& L — Avg. pollen tube length in »

Pollen Viability and Germination in some Orchid Hybrids 253

(f) Media with different growth substances: Cultures in NAA (104 mg/L) showed

the highest percentage germination than in other auxin media, attaining 74% and

the longest tube length of 82 ». Effect of IBA was similar of that of IAA giving

better results than 2,4-D and 2,4,5-TPA media (Table 4). In case of V. Poepoe

for IAA, IPA, 2,4—-D and 2,4,5—-TPA the optimal concentration was 10° mg/L. Of

these four auxin media the highest percentage (71%) germination and the longest

tubes (71 ») were formed in IAA medium and the lowest percentage germination

was obtained in IPA medium (55%); the shortest tubes were produced in 2,4,5-TPA

medium (40 yp).

Table 5. Vanda Josephine. Percentage germination and pollen tube length in optimal

concentration of different media after 24 hrs. at room temperature.

No. of grains germinated per

Media Pe iane a ‘va all %oG | pL

1 2 3 4

Distilled water 29 5 2 0 36 19

(control)

IAA ee ie ee eee 12 12 7 64. | 43

IBA spe tie 10—5mg./L| 32 sal ab? ae ae 52 | 35

IPA 1x 10—5mg./L| 20 15 i 3 49 59

NAA Piet OO ae ie ie eh AL ee ee ae ae 47

eS. iV 10—Saie (Lee. | a8 it fue ae pea 51

2,4,5-TPA Pe HO— Smet | 3 hl. bk,.6 3 60 45

GA lt 10—Sme. fl |) 29 16 | 5 1 a Sa 39

Kinetin ie 10—6mg./L| 14 15 | 18 | 17 64 64

% G — Perc. germination

p. L — Avg. pollen tube length in mM

The optimal concentration for GA and K was 10° mg/L (GA), resulting in

61% germination and 75 » tube length and 55% germination and 34 yu tube

length respectively (Table 4). The different auxin, gibberellin and kinetin media,

at various optimal concentration, gave better results than the control medium. When

compared with borax medium, poorer results were obtained in the two auxin

media, i.e. IPA or 2,4,5-TPA. In all the others, including GA medium, better results

were recorded. Kinetin medium was less favourable than the borax medium and

similarly, gibberellin and kinetin media were not so favourable when compared

with stigmatic extract or sucrose media (Tables 2, 3).

In case of V. Miss Joaquim germination and elongation of pollen tubes in-

creased appreciably in auxins, GA and K media, when compared with the con-

trol or salt-boron medium. The optimal concentration was between 10° and

10*mg/L. Higher concentrations were inhibitory for germination and tube growth.

This was true in all the media except in case of IAA where even at 0.1 mg/L

the percentage germination and tube length were very similar to that of the

control (Table 5). In IAA and kinetin media better germination resulted when

compared with the other auxins or GA media. The media with auxins, GA or K

(at optimal concentrations) were not so favourable as stigmatic extract, sucrose

or borax media (Tables 4, 5).

254 Gardens’ Bulletin — XXVI (1973)

Hybrids other than Vanda

The pollen germination studies in case of: (a) Arachnis Maggie Oei var. Red

Ribbon, (b) Arachnis Maggie Oei var. Yellow Ribbon, (c) Aranda Hilda Gallistan,

(d) Aranda Lucy Laycock (long spray), (e) Aranda Lucy Laycock (short spray), (f)

Aranda Wendy Scott var. Greenfield and (g) Aranthera Lilliput, gave very poor

results in the different media used. In most of them no germination was seen with

the exception of Arachnis Maggie Oei var. Yellow Ribbon the pollen of which

showed small protrusions in the control medium. In borax medium there was

good germination and pollen tube growth at two lower concentrations i.e. 10

(25% G, 11 p) and 10+ (61% G, 32 ») mg/L. In salt boron medium 52%

germination and 29 » tube length resulted. In other media (stigmatic extract with

1, 2 or 3 stigma/10 ml of H, O; sucrose 2-10%) there was no germination. Like-

wise, the different hybrids did not germinate in the auxin, GA or K media at

various concentrations mentioned previously.

Morphology of pollen tubes:

Certain variations were noticed. In V. Peopoe a single case of branching

was observed in NAA medium (10+ mg/L) and slight swellings of pollen tubes

were common in NAA and 2, 4-D media (10° mg/L). Tips of pollen tubes were

pointed in sucrose medium. In V. Miss Joaquim oil globules were present in the

grains and pollen tubes. Tips or middle portion of pollen tubes were swollen in

salt-boron and IAA media (Fig. 22a, 32) and forking of the tube was seen in

kinetin medium (10° mg/L). Exudates were common in GA and K media

(Figs. 23, 25).

Discussion

Structural variation of pollinia in different orchids has been described and

the present study reveals the fact that hybrids retain the same genetic characters

of parents (Adams, 1959). Further study of other hybrids may substantiate this

fact since very few hybrids are critically examined from this point of view.

Occurrence of pollen tetrads is common in Orchidaceae (Withner, 1959). The

presence of supernumerary spores, frequency of their occurrence, and an analysis

of their structure has been studied in the diploid and tetradploids of Vanda hybrids

(Kamemoto, 1956). In five of the nine hybrids studied presently the supernumerary

spores were present and it is interesting to record that only few of them

germinated when compared with the regular tetrads.

Of the nine hybrids investigated germination was seen only in Vanda Poepoe,

V. Miss Joaquim and to a certain extent in Arachnis Maggie Oei var. Yellow

Ribbon. Others did not germinate in the different media used. This loss of

germinability may result from immaturity of pollen, long preservation, or it can

be an inherent property resulting from complicated and forced hybridisation

(Miwa, 1937). It is possible that the reason mentioned last is applicable here

since fresh pollinia were used in all the experiments. Except Vanda, most of the

other hybrids studied presently do not set fruits either by self or cross pollination.

In control medium the percentage germination was low and tube lengths

shorter in both the hybrids of Vanda, In those cases where the pollen grains

germinate with short tubes, it is said that the grains contain a certain amount of

nutrients that promote initial growth but this is insufficient for the growth of

longer tubes (Brink, 1924a). All the growth adjuncts used presently, both organic

Figures 26-35: V. Poepoe. Fig. 26: Germination in dist. water (control). Fig. 27: in

stigmatic extract (2 stigmas/10 ml of dist. water), Fig. 28: in 6% sucrose medium, note beak

like tubes. Figs. 29-35: V. Miss Joaquin. Fig. 29: Tetrads of different sizes. Figs.

30 & 31: Germination in dist. water (control) and one of the tetrads enlarged to show short

tubes in Fig. 31. Fig. 32: Germination in IAA medium (10-5mg/L), showing pollen tubes with

bulbous tips. Fig. 33: in stigmatic extract (2 stigmas/10 ml dist. water). Fig. 34: in borax

medium (10-1! mg/L). Fig. 35: Aborted grains of different sizes.

Pollen Viability and Germination in some Orchid Hybrids 255

and inorganic, were found to be stimulatory when compared with the control

medium. Their effects on percentage germination and tube growth were of different

magnitude. Better results were obtained for Vanda hybrids in stigmatic extract

medium which indicated that the necessary stimulii required for germination and

tube growth were present. The stimulatory effect of stigmatic extract on pollen

germination was observed in other orchids studied earlier (Curtis and Duncan,

1947; Rao and Lee, 1972). Details pertaining to other angiosperms are also

reviewed (Johri and Vasil, 1961).

Sucrose undoubtedly had a stimulatory effect on germination and tube growth

of Vanda hybrids. The optimal concentration varied from 4-6% and best results

were obtained in sucrose media. This is in conformity with earlier findings of

other workers (Miwa, 1937; Curtis and Duncan, 1947). Externally supplied sugars

do serve as a source of nutrition for pollen germination in most of the species

studied so far and the important role of sugars is vividly discussed (Johri and

Vasil, 1961; O’Kelly, 1955, 1957; Visser, 1955).

In case of V. Miss Joaquim, very good germination and tube growth resulted

in boron medium. Vanda Poepoe showed an improvement both in germination

and tube growth but the effect was not as good as stigmatic extract or sucrose

media. Arachnis Maggie Oei var. (Yellow) Ribbon also showed good germination

in the presence of low concentration of borax but the tube growth was poor.

Salt-boron medium was more effective than borax medium in V. Poepoe and

it was contrary in case of V. Miss Joaquim. In case of the latter the presence of

salts reduced both the percentage germination and tube length, though boron was

present as part of the medium. In case of V. Poepoe borax alone was not stimulatory

but its synergistic action with the salts was obvious. Certain salts, especially

calcium (ions), have a stimulatory effect on pollen germination in majority of

angiosperms so far investigated (Brewbaker and Kwack, 1964; Mascarenhas and

Machlis, 1964; Vasil, 1960, 1964). Negative chemotropic response to calcium ions

has also been observed in Lilium, Zea and others (Rosen, 1968, Cook and Walden,

1965). The results obtained in Vanda Miss Joaquim support the above observations.

The fact that pollen grains are naturally endowed with auxins is wellknown

and in certain cases like Antirrhinum, Bryophyllum the auxin concentration is so

high that it is inhibitory to germination (Curtis and Duncan, 1947; Smith, 1942;

Vasil, 1960). In the two Vanda hybrids studied, all the auxins (concentration 10-’

to 10+ mg/L) had a stimulatory effect on germination and tube growth. The

results obtained presently are similar to the observations made in other angiosperms

like Cucumis melo, Eriobotrya japonica, Triticum and others (Loo and Hwang,

1944; Dikshit, 1956; Vasil, 1960). Curtis and Duncan (1947) found that NAA did

not increase pollen germination in the three orchids namely, Cyrtopodium punctatum,

Cattleya guatmalense, and Phalaenopsis hybrid. However, this could be attributed

to the comparatively high concentrations used (0.1 — 10 mg/L) when compared

with present studies.

Though the effect of gibberellic acid and other gibberellins on germination

and elongation of pollen tubes has been extensively studied in other angiosperms,

they have not been used before for orchid pollen (Chandler, 1957; Kato, 1955).

In both the hybrids of Vanda the percentage germination and tube growth improved

in low concentration of GA media. Johri and Vasil (1961) postulated that

incorporation of kinetin in nutrient medium together with auxins may hasten the

division of generative cell. Kinetin improved the percentage germination as well as

tube growth of Vanda Miss Joaquim and the results were much better than in

control, salt-boron, or certain auxin media and similar results were recorded in

256 Gardens’ Bulletin— XXVI (1973)

V. Poepoe. Thus the present studies would indicate that in the Vanda hybrids the

organic nutrients would improve the pollen germination and tube growth to a

greater extent than the inorganic or the other growth substances. As for the

pollen of other hybrids they are viable as indicated by selective staining. Different

physical conditions or growth adjuncts than the ones used presently may be

necessary to induce their germination and these are being investigated.

Summary

A study of pollen morphology, structure and germination is made in the

well-known orchid hybrids. They are Arachnis Maggie Oei var. Red Ribbon,

Arachnis Maggie Oei var. Yellow Ribbon, Aranda Hilda Galistan, A. Lucy Lay-

cock (long spray), A. Luck Laycock (short spray). A. Wendy Scott var. Greenfield,

Aranthera Lilliput, Vanda Poepoe and Vanda Josephine. Experiments were con-

ducted to germinate them in the laboratory conditions. Of these, only Vanda

Poepoe, Vanda Miss Joaquim gave positive results and very limited results were

obtained in case of Arachnis Maggie Oei var. Yellow Ribbion. Pollen of other

hybrids did not germinate. Different substances that are recognised to promote

pollen germination were used at different concentrations and distilled water formed

the control medium. Percentage germination, tube length, and morphological

variations of such structures are determined and described. In these hyrbids,

for pollen germination the organic sources (sucrose, stigmatic extract) appear to be

most effective than the inorganic ones (Borax, salt-boron combination, auxins,

GA or K). Auxins improved the germination and of all the auxina used IAA

was most effective. Similarly GA and K improved the germination in both the

hybrids of Vanda.

Grateful acknowledgment is made to Mr. John Ede who permitted us to

collect the orchids from Mandai garden and Mr. A. G. Alphonso, Acting Director,

Singapore Botanic Gardens for the correct identification of the hybrids and some

technical information. We are thankful to Mr. R. Tay, Mr. D. Teow and Mrs. M.

Goh for their technical assistance.

Literature Cited

Adams, H. H., 1959: Aspects of variation in the Orchidaceae 73-154 in The

orchids — A Scientific survey. (ed.) C. L. Withner. The Ronald

Press Co. New York.

Brink, R. A., 1924: The Physiology of Pollen I. The requirements for growth.

Amer. Jour. Bot., /7, 210-228.

, 1924a: The Physiology of Pollen III. Growth in vitro and in vivo.

Ibid., 11, 351-364.

Brewbaker, J. L. & B. H. Kwack, 1964: The calcium ion and substances influencing

pollen growth. Jn Physiology and Fertilisation (ed.) H. F. Linskens,

North Holland Co. Amsterdam, 143-151.

Chandler, C., 1957: The effect of gibberellic acid on germination and pollen tube

growth. Contrib. Boyce Thomp. Inst., 19, 215-223.

Cook, E. S. & Walden, D. B., 1965: The male gametophyte of Zea mays L. II.

In vitro germination. Can. Jour. Bot., 43, 779-787.

Pollen Viability and Germination in some Orchid Hybrids Zar

Curtis, J. T. & R. E. Duncan, 1947: Studies in the germination of orchid pollen.

Bull. Amer. Orchid Soc., 16, 595-597.

Dikshit, N. N., 1956: Effects of hormone on germination in loquat pollen. Curr.

Sci., 25, 27-28.

Holttum, R. E., 1953, Flora of Malaya — Orchids of Malaya. Govt. Printing Office,

Singapore.

Johri, B. M. & I. K. Vasil, 1961: Physiology of pollen. Bot. Rev. 27, 325-381.

Kamemoto, H., 1956; Cytology of Vanda Nellie Morley and Vanda Emma Van

Dercuter. Amer. Orchid Soc. Bull. 25: 234-243.

Kato, F., 1955: Responses of plant cells to gibberellin. Bot. Gaz. 117: 16-24.

Loo, T. and T. Hwang, 1944: Growth stimulation by manganese sulfate, IAA and

colchicine in pollen germination. Amer. Jour. Bot., 3/: 356-367.

Masceranhas, J. P. & Machlis, L., 1964: Chemotropic response of pollen of

Antirhinum majus. L. to calcium. Plant Physiol. 39: 70-77.

Miwa, A., 1937: Test of germinating power of orchid pollen. Orchid Rev. 45:

345-349,

O’Kelly, J. C., 1955: External Carbohydrates in growth and respiration of pollen

tubes ‘in vitro’. Amer. Jour. Bot. 42: 322-326.

, 1957: Boron effects on growth, oxygen uptake and sugar absorption

by germinating pollen. Amer. Jour. Bot. 44: 239-244.

Roa, A. N. & C. J. Goh, 1970: Progress of orchid research in Singapore, Proc.

Seminar on Res. programmes in Singapore. Nanyang Univ. P. 111-113.

, & Lee Wai Chin, 1972: Experimental studies in the germination

of orchid pollen. La Cellule. 57: 292-310.

Rosen, W. G., 1964: Chemotropism and fine structure of pollen tubes. 159-166.

in pollen physiology and fertilisation (ed.) H. F. Linskens. North

Holland Co. Amsterdam.

Smith, P. F., 1942: Studies on the growth of pollen with respect to temperature,

auxins, colchicin and vitamin B., Amer. Jour. Bot. 44: 637-650.

Vasil, I. K., 1960: Studies on pollen germination of certain Cucurbitaceae. Amer.

Jour. Bot., 47: 239-247.

, 1964: Effect of Boron on pollen germination and pollen tube

growth. Linskens, H. F. (ed.). Pollen physiology and fertilisation.

North Holland, Amsterdam. 107-119.

Visser, T., 1955: Germination and storage of pollen. Mededeel. Landbouwhoo-

geschool (Wageningen), 55: 1-68.

Withner, C. L., 1959: The orchids. A scientific survey. Ronald Press Co., New

York.

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A Review of the Genus Creaghiella Stapf.

(MELASTOMATACEAE)

M. P. NAYAR

Central National Herbarium, Botanic Garden,

Howrah

Abstract

A new species of Creaghiella is described from Borneo. Anerincleistus philip-

pinensis Merr. is transferred to the present genus as Creaghiella philippinensis

(Merr.) Nayar. A key to the species so far known is also presented.

Introduction

Stapf Gin Hook. f. Ic. Pl. 25: t. 2455, 1896) established the genus Creaghiella

on the basis of the specimen Creagh s.n. from Maruda Bay, Sandakan in honour

of Creagh, Governor of Sabah. The genus Creaghiella is allied to Anerincleistus

but differs from it in having subequal or unequal stamens with dorsally gibbose

and ventrally 2-lobed connective, whereas in Anerincleistus the connective is

inappendiculate ventrally and minutely spurred or tuberculate.

The three species C. purpurea Stapf, C. setosa Nayar and C. philippinensis

(Merr.) Nayar form a homogeneous group, having similar facies with densely setose

or tomentose, usually purplish branches, 4-merous flowers, subequal or unequal

stamens with dorsally gibbose and ventrally 2-lobed connective.

The purplish branches, leaves and inflorescence give colour to the different

species of this genus and this can be introduced as a garden plant. This is an

endemic genus with two species (C. purpurea Stapf. and C. setosa Nayar) in Sabah

region of Borneo and a single species C. philippinensis (Merr.) Nayar in Palawan

Island of Philippines. Palawan Island represents the nearest land-bridge from

Borneo to other islands of Philippines. Creaghiella is a genus of densely hirsute

shrubs growing in creeks and sand dunes near sea-shore.

Key to the species of Creaghiella

I. Inflorescence terminal, umbellate.

iI. Calyx tube shaggy tomentose, calyx lobes long; inflorescence axis short,

stout; flowers and capsules larger C. purpurea

II. Calyx tube shortly setose, calyx lobes short; inflorescence axis long,

slender; flowers and capsules smaller C. setosa

I. Inflorescence axillary, fascicled C. philippinensis

259

260 Gardens’ Bulletin — X XVI (1973)

Enumeration of species

1. Creaghiella purpurea Stapf in Hook. f. Ic. Pl. 25: t. 2455, 1896. (Creagh s.n.

Holotype K.)

Distribution: Endemic to Borneo. Borneo: Sabah, Maruda Bay, Tanjong Batu,

Sandakan, 5 May 1895. Creagh s.n. (K) Kg. Malanta, alt. 6 m., 30 May 1933 Olik

NBFD 3327 (K, L.); Elopura, Sandakan F.D., alt. 20 m., 18 Dec. 1947, Kadir

NBFD A.672 (K); Ibid., alt. 50 m., 20 Jun. 1949, Castro NBFD A478 (K); Ibid.,

Kadir NBFD A.944 (K); Elopura, Sandakan, Kabili F.R., Austin Cuadra NBFD

A.515 (K); Ibid., Keith NBFD 4529 (K, L.); Kabili-seilok F.R., alt. 2 m., 12 Jul.

1937, Enggoh NBFD 7298 (K, L.) Batu Sapi road, Sandakan, 19 Aug. 1960, W.

Meijer SAN 21542 (K.).

This is a beautiful shrub of 2-4 m. tall with red flowers and pinkish branches

and inflorescence.

2. Creaghiella setosa Nayar, sp. nov. Affinis C. purpureae Stapf. sed floribus

capsulisque minoribus, lobis calycinis minoribus, calycis tubo minute setoso,

pedunculis longioribus tenuibus differt.

Frutex Rami teretes, dense hirsuto-tomentosi. Folia ovata, 10-20 cm X 6-13

cm, basi rotunda vel subcordata, apice acuminata, margine denticulata, supra

velutino-hispida, subtus hispida, 7-nervia, venulis transversis supra et subtus

distinctis, membranacea; petiolus 2-6 cm longus, dense hirsuto-tomentosus. Jnflo-

rescentia terminalis, 7-15 cm longa, pseudoumbellata, dense hirsuta. Calycis tubus

campanulatus, 5—5.5 mm longus, dense setosus, setis 0.5 mm longis limbus 4 lobatus,

lobis triangularibus, 1-2 mm longis. Petala oblonga, 2.5-3 mm X 1-1.5 mm, apice

ciliata. Stamina 8, subaequalia, filamentis 6-7 mm longis, antheris lanceolatis 6-8

mm longis, apice rostratis, connectivo non producto, antice 2 lobato, postice

gibboso. Ovarium calycis tubo septis 8 adnatum, coronatum. Capsula 4 X 3.5 mm,

dense setosa. Semina minuta, 0.3-0.4 mm longa, numerosa.

TYPUS: G. Mikil SAN 31422 (Holotype L.)

Shrub. Branches terete, densely hirsute tomentose. Leaves ovate, 10-20 cm

x 6-16 cm, base rotund or subcordate, apex acuminate, margin denticulate,

upper surface velvety hispid, under surface hispid, 7-nerved, transverse venules

on the upper and lower surface distinct, membranaceous; petiole 2-6 cm long,

densely hirsute tomentose. Inflorescence terminal, 7-15 cm long, pseudoumbellate,

densely hirsute. Calyx tube campanulate, 5—-5.5 mm long, densely setose, bristles

0.5 mm long, 4-lobed, lobe triangular, 1-2 mm long. Petals oblong, 2.5-3 mm X

1-1.5 mm, apex ciliate. Stamens 8, subequal, filament 6-7 mm long, anther

lanceolate 6-8 mm long, apex rostrate, connective not produced, connective ven-

trally 2-lobed and dorsally gibbose. Ovary adnate to the calyx tube by 8 septa,

disc present. Capsule 4 X 3.5 mm, densely setose. Seeds minute, 0.3—-0.4 mm long,

numerous.

Distribution: Borneo: Sabah. Klias, Beaufort Dist., Sept. 1962, G. Mikil SAN

31422 (L); Beaufort, alt. 66 m., 28 Sept. 1932, Bakar NBFD 3399 (K, L.); Padas

gorge, south side, Tenom Dist., Sept. 1959, W. Meijer SAN 19827 (K, L.); Pangi,

Tenom Dist., alt. 500 m., 5 Sept. 1954, G. H. S. Wood & Wyatt Smith SAN

A.4311 (L.).

Review of Creaghiella (Melastomataceae) 261

C. setosa is closely allied to C. purpurea, but it is immediately recognisable by

its smaller flowers, smaller capsule and short setose hairs in the calyx tube and long

slender inflorescence axis and smaller calyx lobes; whereas in C. purpurea the

flowers and capsule are larger, calyx tube is shaggy tomentose and inflorescence

axis is short and robust.

3. Creaghiella philippinensis (Merr.) Nayar comb. nov.

Anerincleistus philippinensis Merr. in Philipp. Journ. Sc. Bot. 12: 337, 1917.

Distributon: Philippines: Palawan, Malampuya, Sept. 1910, Merrill 7241 (Syntypes

K, BM.); Ibid., May 1913, Merrill 9412 (Lectotype K); Mt. Capoas, Apr. 1913,

Merrill 9552 (Syntype K.).

In the original description it is mentioned that the stamens are dorsally tuber-

culate and ventrally inappendiculate. On careful scrutiny it is seen that the anthers

are dorsally gibbose and ventrally 2-lobate; whereas in the genus Anerinceistus

the subulate anthers end dorsally in a tubercle and ventrally they are inappendi-

culate. Besides the nature of floral parts, the nature of its leaves and its habit

indicate that this taxon is allied to Creaghiella purpurea and C. setosa of Sabah,

Borneo.

In C. purpurea and C. setosa the inflorescence is terminal, whereas in this

taxon the inflorescence is axillary. But in all the three species the flower buds in

juvenile stage are subsessile and the pedicels prominently elongate during anthesis.

C. philippinensis is allied to C. purpurea, but is easily distinguished by its axillary

inflorescence, larger petals (1.5 cm X 1.1 cm) and short ventral appendages;

whereas in C. purpurea the inflorescence is terminal, petals are much smaller

(3.5-4 mm X 2 mm) and the ventral appendages are prominent.

Acknowledgements

I wish to express my gratitude to Sir George Taylor, Director, Royal Botanic

Gardens, Kew, for all facilities during my stay at Kew from 1961-67. I am indebted

to the authorities of the following Herbaria for their hospitality during my visits

and for the loan of herbarium specimens: Rijksherbarium, Leiden and The British

Museum (Nat. Hist.) London. My thanks are also due to Dr. K. Subramanyam,

Director, Botanical Survey of India, for his encouragement.

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New Plant Disease Records for Sarawak

for 1970 and 1971

KUEH TIONG-KHENG

Department of Agriculture, Sarawak, E. Malaysia

Lists of plant disease records for Sarawak have been given by Johnston (1960)

and Turner (1963, 1964, 1966, 1967, 1969 and 1971). The list below consists of

previously unrecorded fungal diseases together with diseases caused by algae and

plant parasitic nematodes, observed or collected by the writer from the time of his

taking over the office of Plant Pathology Section of Semongok Agricultural Research

Centre in mid August, 1970 until the end of 1971. Thirteen of these records appear

in the Annual Report of the Research Branch, Department of Agriculture, Sarawak,

for the year 1970.

The causal organisms are listed under their respective host plants which are

arranged in alphabetical order of their botanical names. The frequency of occur-

rence is given together with the Commonwealth Mycological Institute Herbarium

serial number, where identification has been performed by the Institute. Two

species of fungi and one nematode species were identified at the Royal Botanic

Gardens, Kew, and the Commonwealth Institute of Helminthology, Herts, respec-

tively.

In the list that follows, One, Occ., Comm. stand for One record, Occasionally

and Common respectively.

Alternanthera bettzickiana var. spathulata

Wilt Corticium rolfsii Curzi One —

Ananas comosus Merr, (Pineapple, Nanas)

Leaf rot Curvularia eragrostidis (P. Henn.)

J. A. Meyer One 142889

Annona squamosa L. (Sweet Sop)

Red rust Cephaleuros virescens Kunze One —

Arachis hypogaea L. (Groundnut, Kachang Tanah)

Wilt Fusarium solani (Mart.) Sacc. One 159548

Begonia glabra Ruiz. et. Pav.

Wilt Corticium rolfsii Curzi One —

Bixa orellana L. (Annatto)

Leaf blight Pesialotiopsis theae (Saw.) Steyaert One 163728a

Leaf blight Colletotrichum capsici (Syd.) Butler &

Bisby One 163728b

Brassica juncea Coss. (Kai Choy)

Leaf rot Choanephora cucurbitarum

(Berk. & Rav.) Thaxt. Occ. —

263

264 Gardens’ Bulletin — X XVI (1973)

Canangium odoratum Baill. (Kenanga)

Velvet blight Septobasidium sp. One —

Capsicum annuum L. (Chilli, Chabai)

Leaf spot Leptosphaerulina trifolii (Rostrup) Petr. One 153073

Capsicum frutescens L. var. baccatum (Ornamental pepper)

Leaf spot Cercospora capsici Heald & Wolf Occ. 163286

Carica papaya L. (Papaya)

Leaf spot Sphaerulina sp. One 159602b

White root disease Fomes lignosus (Klotzsch) Bres. One —

Chrysanthemum sp.

Flower blight Cladosporium oxysporium Berk. & Curt. One 160403a

Flower blight Fusarium equiseti (Corda) Sacc. One 160403b

Leaf spot Cercospora chrysanthemicola Yen One 163291a

Leaf spot Septoria chrysanthemella Sacc. One 163291b

Citrus microcarpa Bunge (Musk-lime, Limau kesturi)

Pink disease Corticium salmonicolor Berk. & Br. One —

Cocos nucifera L. (Coconut, Kelapa)

From roots of wilting Cunninghamella echinulata (Thaxt.)

palm Thaxt, ex Blakeslee One 159555

From roots of wilting Ceratocystis paradoxa (Dade) Moreau One 159557

palm

From roots of wilting Fusarium monileforme Sheld. One 159562

palm

From fabric bark of Fusarium solani (Mart.) Sacc. One 159556 & 159559

wilting palm

Codiaeum variegatum Blume (Garden Croton)

Velvet blight Septobasidium sp. One 163287

Coffea robusta Linden (Robusta coffee, Kopi)

Leaf blight Glomerella cingulata

(Stonem.) Spauld. & Schrenk One 163289

Elaeis guineensis Jacq. (Oil palm)

Stem rot ? Thielaviopsis sp. One 148041

Dry basal rot Ceratocystis paradoxa (Dade) Mareau Occ. 159563

Ficus glossularioides Burm. f.

Leaf spot Glomerella cingulata

(Stonem.) Spauld. & Schrenk One 163726

Ficus uncinata Becc.

Leaf spot Cercospora sp. One 163727

Fimbristylis globulosa Kunth (Globular Fimbristylis, Rumput sandang)

Inflorescence smut Cintractia exicola (Berk.) Cornu Occ. 163730a & 163729

Inflorescence stalk Stauronema sp. One 163731

blight

Inflorescence smut Fusarium heterosporum Nees ex Fr. One 163730b

Inflorescence smut Fusarium oxysporum Schlecht. One 163730c

New Plant Disease Records, Sarawak, 1970 and 1971

Fimbristylis miliacea Vahl (Lesser Fimbristylis, Rumput tahi kerbau)

Wilt Corticium rolfsii Curzi

Flemingia congesta Roxb. (Bush cover crop)

Thread blight Marasmiellus scandens

(Mass.) Dennis & Reid

Glycine max (L.) Merr. (Soybean)

Leaf spot Phyllosticta glycinea Tehon & Daniels

Leaf spot Ascochyta sp.

Leaf blight Choanephora circinans

(Nag. & Kawak.) Hess. & Benj.

Leaf blight Curvularia eragrostidis

(P. Henn.) J. A. Meyer

Associated with leaf Fusarium equiseti (Corda) Sacc.

blight

Guizotia abyssinica (L.f.) Cass. (Niger seed)

Collar rot Corticium rolfsii Curzi

Impatiens balsamina L. (Balsam)

Leaf spot Cercospora fukushiana (Mats.) Yam.

Wilt Corticium rolfsii Curzi

Imperata cylindrica Beauv. (Lalang)

Leaf spot Colletotrichum graminicola

(Ces.) Wilson

False smut Cerebella andropogonis Ces.

Inflorescence mould Curvularia pallescens Boedijn

One

Occ.

One

Occ.

One

Occ.

One

Leersia hexandra Swartz. (Tiger’s tongue grass, Rumput lidah rimau)

Leaf spot Pyricularia oryzae Cav.

Mangifera indica L. (Mango)

Leaf anthracnose Glomerella cingulata

(Stonem.) Spauld. & Schrenk

Manihot utilissima Pohl. (Cassava, Tapioca, ubi Kayu)

Leaf blight Pestalotiopsis sp.

Leaf spot Phyllosticta manihot Speg.

Tuber rot Sphaerostilbe repens Berk. & Br.

Wilt Corticium rolfsii Curzi

Michelia alba DC. (White chempaka, Chempaka puteh)

Thread blight Marasmiellus scandens

(Mass.) Dennis & Reid.

Musa sapientum L. (Banana, Pisang)

Leaf speckle Periconiella muse M. B. Ellis

Leaf spot Pestalotia leprogena Speg.

Heart leaf disease Drechslera mus@-sapientum

(Hansf.) B. Ellis

Myristica fragrans Houtt. (Nutmeg, Buah pala)

From roots Gliocladium sp.

265

160402e

160402f

160402a

160402d

160402c

163734

163282

154092a

154092b

Comm. —

One

163733

Comm. 163292a

Occ.

One

163292b

153071

153070

163281

159551

266

Gardens’ Bulletin— X XVI (1973)

Nephelium lappaceum L. (Rambutan)

Fruit blight

Fusarium sp. One 163279

Oryza sativa L. (Rice, Padi)

Brown grain

On grain

Wilt

Leaf spot

Brown grain

Pithomyces sacchari (Speg.) M. B. Ellis One 159593b

Pestalotiopsis sp. One 159593c

Corticium rolfsii Curzi Occ. —

Curvularia eragrostidis

(P. Henn.) J. A. Meyer One 142890

Pestalotiopsis ? disseminata

(Thum.) Steyaert One 142891

Pestalotiopsis palmarum

(Cooke) Steyaert One 159549

Pachyrrhizus erosus Urban. (Yam bean)

Wilt

Colletotrichum capsici

(Syd.) Butler & Bisby One 163280

Panicum repens L. (Creeping panic grass, Rumput halia)

Wilt

Blast

Corticium rolfsii Curzi Occ. —

Pyricularia oryzae Cav. Comm. —

Paspalum scrobiculatum L. (Carpet grass)

Seed blight

Fusarium heterosporum Nees ex Fr. Comm. 154093b

Pennisetum purpureum Schum. & Thonn. (Napier grass)

Leaf spot

On dead stems &

leaves

Curvularia eragrostidis

(P. Henn.) J. A. Meyer Occ. 163288a

Cochliobolus lunatus Nelson & Haasis One 163288d

Pithomyces chartarum

(Berk. & Curt.) M. B. Ellis One 163288c

Septoria sp. One 163288b

Volvariella esculenta (Massee) Sing. One —

Peperomia pellucida H.B. & K.

Wilt

Corticium rolfsii-Curzi One —

Piper betle L. (Betel, Sireh)

Pink disease

Corticium salmonicolor Berk. & Br. One —

Piper nigrum L. (Black pepper, Lada Hitam)

On dead branches

On dead spikes, leaves

& branches

Black spike

Leaf spot

Pleurotus (Hohenbuehelia) testudo Berk. One Kew

Marasmius campanella Holterm. . | One Kew

Cochliobolus geniculatus Nelson One 159552

Glomerella cingulata

(Stonem.) Spauld. & Schrenk One 159553

Pogostemon cablin Benth, (Patchouli)

White root

Root knot

Wilt

Fomes lignosus (Klotzsch) Bres. One —

Meloidogyne javanica (Treub., 1885)

Chitwood, 1949 Occ. CIH

Corticium rolfsii Curzi One —

New Plant Disease Records, Sarawak, 1970 and 1971

Ricinus communis L. (Castor oil plant)

Leaf spot

Fruit blight

Leaf spot

Fruit blight .

Fruit blight

Alternaria ricini (Yoshii) Hansford

Colletotrichum capsici

(Syd.) Butler & Bisby

Ascochyta sp.

Fusarium oxysporum Schlecht.

Botryodiplodia theobromae Pat.

Glomerella cingulata

(Stonem.) Spauld. & Schrenk

Occ. 159600b

One 154094a

One 159600a

One 154094b

One 154094c

One 154094d

On Clypeolella ricini Cicinobella sp.

Rosa sp. (Rose)

Leaf spot Mycosphaerella sp.

Sanchezia nobilis Hook. f. (Kadok gajah)

Leaf rot Corticium solani (Prill. & Delacr.)

Bourd, & Galz.

Sorghum sudanense Stapf (Sudan grass)

Leaf spot Fusarium sp.

Sorghum vulgare Pers. (Sorghum)

Leaf spot

Panicle disease

Cercospora sorghi Ell. & Ev.

Fusarium semitectum Berk. & Rav.

Alternaria Iongissima Deighton &

One

159601b

159597b

163278b

153068

153069b

Collar rot

MacGarvie

Corticium rolfsii Curzi

Uncaria gambir Roxb. (Gambir)

Leaf rot

Vanda vars. (Orchid)

Leaf spot

Corticium solani (Prill. & Delacr.)

Bourd. & Galz.

Cochliobolus geniculatus Nelson

Vanilla planifolia Andr. (Vanilla)

Rust

Uredo scabies Cke.

Zea mays L. (Maize, Jagong)

Leaf spot

Curvularia senegalensis (Speg.) Subram.

Diplodia zeicola Saccas

Zingiber officinale Rosc. (Ginger, Haliya)

Leaf scorch Hendersonia zingiberi Sawada

Leaf spot Coniothyrium fuckelii Sacc.

Leaf spot Ophiobolus sp.

Leaf scorch Tetraploa aristata Berk. & Br.

Leaf scorch Leptosphaeria sp.

Leaf blight Pyricularia zingiberi Nishikado

Red rust Cephaleuros virescens Kunze

Leaf blight Leptosphaeria typhae Karst.

Leaf spot Phyllosticta zingiberi Ramakrishnan

Leaf blight

Cochliobolus geniculatus Nelson

One 153069c

One —

One 153074

One 159596

One 163284a

One 163284b

One 163285a

One 163285b

One 163285c

One 163283b

One 163283c

One 159594b

One 159594c

One 159594a

Comm. 153072a

163283d

Comm. 153072b

163283a

267

268 Gardens’ Bulletin — X XVI (1973)

Acknowledgements

I am grateful to the Director of Agriculture, Sarawak, for granting permission

to publish this list and the Directors and staff of the Commonwealth Mycological

Institute, the Royal Botanic Gardens, Kew, and the Commonwealth Institute of

Helminthology, Herts, without whose assistance in identifying many of the species,

the list could not have been published.

References

Annual Report (1970). Research Branch, Department of Agriculture, Sarawak.

Johnston, A. (1960). A preliminary plant disease survey in Sarawax. F.A.O. Rome

(mimeographed).

Turner, G. J. (1963). New records of plant diseases in Sarawak for the years 1960

& 1961. Gdns’. Bull. Singapore, 20 285-288.

Turner, G. J. (1964). New records of plant diseases in Sarawak for the year 1962.

Gdns’. Bull. Singapore, 20 369-376.

Turner, G. J. (1966). New records of plant diseases in Sarawak for the years 1963

& 1964. Gdns’. Bull. Singapore, 21 393-402.

Turner, G. J. (1967). New records of plant diseases in Sarawak for the year 1965.

Gdns’. Bull. Singapore, 22 123-128.

Turner, G. J. (1969). New records of plant diseases in Sarawak for the years 1966

& 1967. Gdns’. Bull. Singapore, 24 181-184.

Turner, G. J. (1971). Fungi and plant diseases in Sarawak. Phytopath. Papers No.

13, C’wealth Mycol. Inst., Kew, Surrey, England. Jan. 1971.

Notes on the Systematy of Malayan Phanerogams

XVIII — XXII *

K. M. KOCHUMMEN & T. C. WHITMORE

Forest Research Institute, Kepong

Abstract

Calophyllum austrocoriaceum and C. rupicolum var. elatum are new taxa. C. intramar-

ginale is reduced to C. inophylloide var. singapurense.

A division of Malayan Garcinia into groups is described. G. burkillii, G. cantleyana and

its var. grandifolia, G. cataractalis, G. hendersoniana and G. murtonii are new species. G.

dumosa (and its Malayan allies), G. nigrolineata, G. penangiana, G. bancana, G. opaca and

G. pyrifera are annotated with several new synonyms proposed; G. opace var. dumosa is

proposed as a new variety.

In Mesua notes are provided on the alliances of M. assamica, M. elegans, M. kunstleri

and M. lepidota; and M, kochummeniana, M. nivenii, M. nuda and M. purseglovei are new

species.

Barringtonia payensiana is a new species.

Crudia viridiflora is a new species, with simple leaves.

In Olacaceae, Strombosia maingayi is shown to be the legitimate name for the tree

long known as S. rotundifolia; the differences from S. multiflora are discussed.

In Proteaceae, Heliciopsis cockburnii, H. montana and H. whitmorei are described as

new species.

This fourth instalment of our series of notes completes the taxonomic ground-

work for volume 2 of the Tree Flora of Malaya, which takes the project over the

half way mark. The longest note is on the Guttiferae, mainly Garcinia, the second

biggest family (after Euphorbiaceae) treated in volume 2. Yet another new legu-

minous species has been discovered, this time a simple-leaved Crudia. Once again

we express our thanks to Dr. B. C. Stone for help with the latin descriptions.

XVII. GUTTIFERAE

T. C. WHITMORE

Calophyllum

Calophyllum austrocoriaceum Whitmore sp. nov.

C. sp. 10 in Henderson & Wyatt-Smith Gdns’ Bull. Sing. 15 (1956) 313, plate VILL,

Fig. 3 E.

Arbor parva, ramulis crassis, laevibus, leviter complanatis, furfuraceis; gemmis

terminalibus crassis, lanceolatis, acutis, subtiliter tomentosis, usque 15 mm longis,

foliis valde coriaceis, oblongis vel anguste ovate-ellipticis (7 x 3) - ll x 3 - 16

x 5 cm, apice rotundato vel obtuso, base cuneato, leviter decurrenti, costa lato

* continued from Gdns’ Bull. Sing. 26: 49-61.

269

270 Gardens’ Bulletin — X XVI (1973)

pagina infera prominenti pagina supra prope basim interdum canaliculato, nervis

utrinque conspicuis subdistantibus, petiolo 2-3 cm crasso. Inflorescentiae axillares

pauciflorae. Flores ignoti. Fructus subglobosa, usque 23 mm longa apiculata,

pericarpio crasso duro siccitate.brunneo sublaevi, pedicello crasso 1 cm, pedunculo

crasso 15 mm.

JOHORE: G. Panti FRI 7846 (holotype KEP); FRI 7729, 13819, SFN 29969;

G. Belumut KEP 98038, FRI 7512, 7789, 8722, 8809.

Not seen: SFN 10723, cited by Henderson and Wyatt-Smith.

This is close to C. coriaceum of the mountains of the Main and east coast

ranges and replaces it on the southermost mountains of the Peninsula. The name

is chosen to reflect this affinity. It is also closely related to C. tahanense of G.

Tahan. From both of its relatives C. austrocoriaceum conspicuously differs in the

nerves being prominent and coarse on both surfaces, the short, few-flowered,

axillary racemes and the apiculate unbeaked fruit.

Calophyllum imophylloide var. singapurense Henderson et Wyatt-Smith Gdns”

Bull. Sing. 15 (1956) 316, plate IX, Fig. 4 D-H.

C. intramarginale Henderson et Wyatt-Smith loc. cit. 342, plate XXII, Fig. 13A, B-

syn. nov.

The latter species is said to differ in its smaller leaves with intramarginal vein,

shorter inflorescences with all the flowers apical on the peduncle, and its montane

habitat.

It appears that Henderson and Wyatt-Smith drafted the monograph when

only the type of C. intramarginale was known, a collection from a small tree

from 4000 ft on G. Padang; and that KEP 67753, from 1020 ft in Rasau Kerteh

F.R., was added later, but neither the key nor notes were amended to allow for

its lowland station.

I have since made a second collection from G. Padang (FRI 12644) in which

the intramarginal nerve is very difficult to see. Three other collections have recently

been made which have an intramarginal nerve on one or a few leaves (FRI 41/4,

8839, 10840) and these are all big trees which in leaf shape, bark, exudate colour

and the possession of buttresses resemble C. inophylloide var. singapurense. They

come from W. Kelantan, N. E. Johore and Trengganu respectively.

The flowers of C. inophylloide var. singapurense arise in a cluster from the

top of the peduncle and sometimes, on long racemes, there are also 1-2 lateral

pairs. In C. intramarginale these lateral pairs are always lacking.

I do not believe C. intramarginale can be maintained and therefore reduce it.

C. inophylloide var. singapurense differs mainly from the typical variety in

its smaller leaves.

Calophyllum rupicolum var. elatum Whitmore var. nov.

C. rupicolum var. in HENDERSON & WYATT-SMITH Gdns’ Bull. Sing. 15

(1956) 347, plate XXVII, Fig. 13 M, N.

A varietas typica elatiore usque 18 m, in foliis maioribus, latioribus, 8 x 2.7 — 11

x 3.5 cm, differt.

Systematy of Malayan Phanerogams XVIH-X XII 271

KELANTAN: FRI 2538 (holotype of var. at KEP). TRENGGANU: Ulu Brang

FRI 15256. G. Tebu F. R. FRI 2513, 2516. Bt. Rambai F. R. FRI 11401. KEDAH:

Ulu Muda F. R. FRI 6757. PAHANG: Rompin KEP 29648. PERAK: Dindings,

Telok Kopia F. R. FRI 3105. JOHORE: Kangka Sedili Kechil, Corner s.n. 17. VI.

34; Mawai, SFN 34706, 34747.

Henderson & Wyatt-Smith left this as an undescribed variety. More material

has now been collected and I feel confident in describing it, but as a variety of

the river-bed bush C. rupicolum and not as a full species. Apart from the relatively

broader leaves, and the habit and habitat it comes very close to C. rupicolum,

and there is some overlap in leaf.

All the collections are of small trees away from rivers except FRI 12526,

described as a riverside bush, yet with leaves 11 x 3.5 cm, the largest seen.

I have studied C. rupicolum on the Sg. Tahan near K. Tenor. There it grows

as a small, sinuous tree to 6 m tall on earth river banks at shingle shallows. Leaf

size was greatest at the top of the crown and tiny at the crown base, below the

level of the highest floods. The collection, FRI 15985, clearly shows the narrow

leaves of the typical variety, they are up to 8 cm long.

Garcinia

I have found it useful to divide Malayan Garcinia into six groups, named

A-F, based on the structure of the male flower. These groups are a refinement

of. the numbered sections proposed by King in the key to his account in the

Materials (J. As. Soc. Beng. 59, 1890, 148-72) and which were more or less copied

by Ridley in his Flora (1, 1922, 167-80). To some extent they are natural, though

groups B and C very probably are not (species of subgroups Bi and Ci are very

similar and easily confusable). A guide to the groups in the form of a key appears

in my account for the Tree Flora and this should also be consulted. It must be

emphasised that the groups are informal ones, and are based on a study only

of Peninsula material; they are published here because I believe they might per-

haps provide a useful starting point for other students of this big and intricate

genus, but they certainly should not be regarded as definitive. statements on the

taxonomy of the whole of Garcinia. The subgroups, where such are defined, are

species which are closely similar in one or several respects, i.e. ‘circles of affinity’,

and not all species fall into clear subgroups. Further notes on individual species,

including the subgroups, can be found in the Tree Flora.

In these studies I have not found helpful the groupings proposed by Anderson

(in Hk. f. Fl. Brit. Ind. 1, 1884, 259-70) or those proposed by Pierre (For. FI.

Cochinch. 1, 1883, i—xlii).

The groups are defined on the structure of the male flower. Other characters

common to the component species are also given in the descriptions below. There

is a final artificial group, G, of 3 incompletely known species.

Group A. Sepals and petals 4. Stamens numerous, on 4 fleshy processes, anther

cells 2 (4 in G. cuspidata, fide King), pistillode present.

Subgroup Ai. Leaves reminiscent of Eugenia, with fine, rather close, parallel,

secondary nerves arising nearly at right angles to the midrib. Flowers small, in

small inflorescences or clusters, axillary or behind the leaves.

272 Gardens’ Bulletin — X XVI (1973)

G. cuspidata King, G. eugeniaefolia Wall., G. merguensis Wight, G. rostrata

Wall., G. sp. Al.

A closely knit alliance for which I have only been able to give a provisional

account; species delimitation needs a critical reappraisal on a monographic basis.

See also the notes under G. rostrata below.

The following species, dissimilar from each other and from members of the

Subgroup Ai, are also in group A:

G. diversifolia King, G. holttumii Ridley, G. monantha Ridley, G. murtonii Whit-

more, G. sp. A2, G. sp. A3.

Group B. Sepals and petals 4. Stamens numerous, joined in a sometimes 4-lobed

ring, surrounding a central pistillode; anther cells 2. Flowers terminal, males

several, females solitary.

Subgroup Bi. Most easily defined as a close knit alliance of spp. extremely similar

to the well known mangosteen; virtually indistinguishable when sterile and differing

most noticeably in stigma.

G. hombroniana Pierre, G. malaccensis Hk. f., G. mangostana L.

Species of this alliance occur throughout the East Indies, the best taxonomic

treatment of the variation remains to be ascertained from a monographic base.

Subgroup Bii. G. costata Hemsley ex King, G. maingayi Hk. f. Two species

similar in their coarsely ribbed leaves.

Subgroup Biii. Male flowers in a short, terminal raceme, females solitary. Leaves

drying thin, green-black.

G. atroviridis Griff ex T. Anders. Very distinctive.

Group C. Flower structure just as in Group B but with no pistillode, so the

stamens as a central mass. Flower position various.

Subgroup Ci. Flower position as in Group B. Leaves drying with a distinct reddish

or orange tint.

G. murdochii Ridley, G. opaca King, G. penangiana Pierre, G. sp. Cl (?).

This Subgroup is very close to Bi above. It is discussed further below, under

G. penangiana.

Subgroup Cii. Flowers mostly behind the leaves, in clusters. Leaves leathery, drying

with a distinct reddish tint, or drying thin and green to green-black in G. forbesii,

G. griffithii.

G. bancana (Miq.) Miq., G. clusiaefolia Ridley, G. forbesii King, G. griffithii

King, G. sp. C2.

Subgroup Ciii. Leaves with rather open, reticulate nervation, drying brown-green.

Flowers minute, in axillary racemes.

G. minutiflora Ridley.

Not placed: G. montana Ridley.

Group D. Sepals and petals 4. Stamens numerous, as a central mass; no pistillode;

anther cells 4. Flowers small, in clusters in axils and behind leaves. Leaves drying

thin, commonly pinkish grey-brown and with a fine pattern of black dots and dashes.

Systematy of Malayan Phanerogams XVIII-X XII 273

G. cowa Roxb., G. nigrolineata Planch. ex T. Anders., G. parvifolia (Miq.)

Migq.

Corner (Gdns’ Bull. Str. Settl. 10, 1938, 36-8) sorted out the Malayan members.

of this small group.

Group E. Sepals and petals 4. Stamens not very numerous, connective peltate with

the anther cells marginal, often confluent, their dehiscence circumscissile; pistillode

absent. Fruits thinly woody. Leaves often drying greenish.

Subgroup Ei. Flowers and fruits clustered on small tubercles, axillary and behind

the leaves. Leaves usually thin. Often small shrubs.

G. dumosa King, G. scortechinii King, G. urophylla Scortechini ex King,

Golspy £3

This Subgroup is discussed below, under G. dumosa.

Subgroup Eii. Fruits solitary, mainly in leaf axils, some behind the leaves. Leaves:

usually leathery. Mainly mountains.

G. burkillii Whitmore, G. cantleyana Whitmore, G. hendersoniana Whitmore,

G. uniflora King, G: Sp. El (?), G. Sp. E2.

This Subgroup shows partly allopatric, partly sympatric distribution patterns

in the mountains which are discussed in the Tree Flora.

Group F. Sepals and petals 5. Stamens in 5 bundles around a small central pistil-

lode. Male and female (and hermaphrodite ?) flowers sometimes mixed. Fruits

usually drying shiny brown-black, intricately, sinuously wrinkled. Leaves commonly

drying with a greenish or yellowish tinge.

G. dulcis (Roxb.) Kurz, G. nervosa (Miq.) Miq., G. prainiana King, G. pyrifera

Ridley, G. xanthochymus Hk. f.

Long recognised, and at one time gave generic status as Xanthochymus Roxb.

Group G. Incompletely known species.

G. cataractalis Whitmore, G. sp. G1, G. sp. G2.

Garcinia burkillii Whitmore sp. nov.

Arbor parva, foliis subcoriaceis, ovatis 7.5 x 4.5 — 15 x 7 cm, acutis, base late

cuneato vel rotundato, minime decurrenti, siccitate fuscis, marginibus leviter recur-

vatis, nervis lateralibus tenuibus. Flores ignoti. Fructus axillaria solitaria ovoidea,

2 cm longa, usque ad 12 mm diam., siccitate castanea, laevia, tenuiter lignosa, cum

sepalis persistentibus ovatis 6 mm latis longisque, stigmatis integro tholiformi

3 mm diam. subsessili subtiliter papillato, axillari, solitario.

TRENGGANU: Sg. Loh, K. Datok, FRI 12003. PAHANG: Tahan River, Ridley

2239; G. Tapis FRI 10911 (holotype KEP).

In leaf this is close to G. uniflora King of the Main Range, to which I believe

it is related (though the flowers remain unknown). The fruiting stigma is, however,

quite different. It is also quite clearly a very close relative of G. hendersoniana

described below, and differs mainly in the fruit and its stigma.

Garcinia cantleyana Whitmore sp. nov.

Arbor parva, foliis parvis, crasso coriaceis, ovatis, 5.5 x 3 - 10 x 5 cm, apice

late rotundato interdum apiculato, base late cuneato, marginibus valde revolutis;

nervis lateralibus tenuissimis, petiolo crasso 14 mm. Flores (femineis ignotis) in

274 Gardens’ Bulletin — XXVI (1973)

axillas foliorum delapsorum poxiti, sessiles, sepalis 4 ovatis 4 x 4 mm, petalis 4

quam sepalis aliquantam minoribus, masculi cum staminibus numerosis androecium

globosum centralium formantibus, filamentis crassis 1 mm, antheris peltatis 1 mm

latis rima lata dehiscentibus, filamentis 1 mm longis spissibus, pistillodio nullo.

Fructus subglobosa 3 cm diam., siccitate brunnea, sessilia, calyce persistenti

lignoso insidens, stigmatis 3 mm diam. integro tholiformi grosse papilloso sessili.

PAHANG: Fraser’s Hill, FRI 5819 (holotype KEP). PERAK: G. Korbu, SFN

16301, KEP 45976; Cameron Highlands, SFN 20603.

G. cantleyana var. grandifolia Whitmore var. nov.

A varietate typica in foliis maioribus, 17 x 10 cm, fructibus interdum ovoideis

differt.

PAHANG/SELANGOR: crest of Main Range, road to G. Ulu Kali, FRI 12576

(holotype KEP), FRI 16185.

Garcinia cataractalis Whitmore sp. nov.

Frutex humilis, foliis lineari — lanceolatis 6 x 0.5 cm. Flores ignoti. Fructus

axillaria, carmina, subglobosa, 2—3 lobata, vivo 2.5 cm diam. (siccitate valde vietis,

grosse rugosa, 1.5 cm diam.), valde 8 mm rostrata, stigmate nigro 4 — lobato

undulato minute papilloso leviter procurrenti, seminibus valde complanatis in pulpa

aquosi translucenti inclusis.

TRENGGANU: Sg. Kerbat at Jeram Petang: FRI 20350 (holotype KEP); Sg.

Nipah at Jeram Gajah: SFN 2591/3.

Well known to the local boat men on the Sg. Kerbat as gelugor batu; a highly

distinctive rheophyte. |

/ Garcinia dumosa King

G. tenuifolia Ridley Kew Bull. (1928) 72 syn. nov.

Garcinia scortechinii King

G. gaudichaudii var. minor Ridley Flora 1 (1922) 177 syn. nov.

Garcinia urophylla King

All King’s three species were described in the Materials (J. As. Soc. Beng. 59,

1890, 166-8). They are only weakly differentiated by the descriptions. The only

syntype at Singapore is a sheet of Curtis 1249, G. scortechinii. I am able provision-

ally to maintain these species on the weak differential characters given in the

Tree Flora but they are undoubtedly close, with apparently little difference in

their reproductive parts. A monographer will have to reconsider this group together

with its Indo-Chinese relatives. I do not see that Ridley was justified in reducing

G. scortechinii to the Indo-Chinese G. gaudichaudii Planch. et Triana, as he did

not make the necessary critical comparative studies of this whole alliance, I there-

fore do not follow him and, further, I reduce his var. minor of G. gaudichaudii

to synonymy, having seen the Johore syntype. G. tenuifolia from the islands off

Pahang is simply superfluous, I see no difficulty in reducing this species to G.

dumosa.

Systematy of Malayan Phanerogams XVIII-X X11 275

Garcinia hendersoniana Whitmore sp. nov.

Arbuscula vel arbor parva, foliis percoriaceis, ovatis, 8 x 5.5 - 14 x 8.5 cm,

apice acuto, base late cuneato vel rotundato, marginibus revolutis, nervis lateralibus

tenuibus attamen manifestis, petiolo 1-2 cm longo. Flores (masculis ignotis) sessiles,

solitarii, in axillas foliorum delapsorum poxiti, sepalis 4 ovatis, exterioribus (2)

9 x 8 mm, interioribus (2) 5 x 8 mm, petalis 4 ovatis unusquisque 5 x 5 mm,

staminodiis peltatis circulatim dispositis, ovario cylindrico 4 mm longo x 3 mm

diam., stigmate sessili tenui integro circulari-discoideo paullo convexo grosse

papilloso paullo projectenti 4 mm diam. Fructus (? immaturi) ampulliformia vel

rotundata, 3 cm longa, siccitate nigro-cinerea laevia, tenuiter lignosa, sepalis

persistentibus.

TRENGGANU: G. Padang SFN 31065, 31824; FRI 12718; G. Mandi Angin FRI

10813 (holotype KEP), FRI 10826, 12103. PAHANG: G. Tapis FRI 10994.

PAHANG/SELANGOR: G. Ulu Kali FRI 16206.

Garcinia murtonii Whitmore sp. nov.

Frutex vel arbuscula, vel arbor parva, foliis glabris, coriaceis, ellipticis vel

ovato-ellipticis, 5 x 2 - 8 x 3.8 — (8.5 x 4.5) cm, apice acuto, base longe cuneato,

leviter decurrenti, nervis lateralibus approximatis (inter se 2 mm) parallelis rectis

prope marginem bifurcatis gracilis utrinque prominentibus, costa striato utrinque

prominenti. Flores (masculi) in racemis c. 1 cm longis congestibus, spissibus, axil-

laribus aggregatis, pedicellis 2 mm longis, sepalis 4 ovatis 2 mm latis caduceis,

petalis 4 simillimis, staminibus in 4 fasciculos dispositis, pistillodio fungiformi.

Flores feminei ignota. Fructus globosa, 2 cm diam., nigricans, laevia, stigmate

disciformi aliquantam quadrato integerrimo grosse papilloso 2 mm diam. promi-

nulenti, sepalis nec persistentibus caduceis.

JOHORE: G. Belumut, KEP 33830, KEP 33850 (holotype KEP), FRI 7511, 8730,

8737, 17825, SFN 10990. PERAK: G. Korbu, KEP 31486.

Garcinia nigrolineata Planch. ex T. Anders. Fl. Brit. Ind. 1 (1874) 263.

G. kunstleri King J. As. Soc. Beng. 59 (1890) 165 syn. nov.

King cites collections by Wray and Scortechini. Ridley (Flora 1, 1922, 177)

adds no others. I have seen at Singapore Wray 828, Scortechini 271, 1852 and

s.n., probably all the syntypes. The fruit is not grooved and ridged, the stigma is

2.5 — 3.5 mm wide, papillose and slightly raised, but not on an umbo; the female

flowers are 3 mm across on slender, 5 mm pedicels and contain a globose, very

young fruit. G. kunstleri has not been recollected. Apart from the very small

female flowers it fits well Corner’s diagnosis of G. nigrolineata in Gdns’ Bull. Str.

Settl. 10 (1938) 36-8, and I have little hesitation in suggesting that it is but a

weakly differentiated form of that species, best reduced to synonymy.

On the Scortechini sheets I have seen have no locality notes. I do not know

what led Ridley (loc. cit.) to say they came from the big limestone hill at Kuala

Dipang. There is no limestone near Tapah, the Wray locality.

The G. penangiana group (Group C)

Pierre (Fl. For. Cochinch. fasc. 6, 1883, 37) described G. penangiana from

Wallich’s collector (Porter) 4852D, a collection with male flowers in bud. He

pointed out that Hooker had already mentioned it as distinct from G. cornea L.

in a parenthetical addition to Anderson’s account of Garcinia in FI. Brit. Ind. |

276 Gardens’ Bulletin — X XVI (1973)

(1874) 260. King saw further collections and amended the description in his

account of Garcinia in the Materials (J. As. Soc. Beng. 59, 1890, 158). I follow

what I believe to be King’s circumscription, although he does not precisely state

which collections he saw, except for Kunstler 3853, which is not at Singapore.

I have examined at Bogor, Teysmann s.n. from Riouw, P. Bientang, the type

of G. caloneura Boerl. Cat. Hort. Bot. Bogor 68 (2) 1901. It looks very similar

to G. penangiana.

Dr. Kostermans has for some years now been labelling as G. celebica L. all

specimens of this species-group in the herbaria he has visited.

All these species are of very similar appearance and I see no prospects of

resolving their limits without monographic study.

There has been confusion between the G. penangiana group and the G.

hombroniana alliance, my Subgroup Bi; this might indicate close affinity, implying

that, as I indeed suspect, my species-groups, although convenient, are not entirely

natural aggregates.

G. bancana (Miq.) Miq. Fl. Ind. Bat. Suppl. (1861) 494. G. bancana var. curtisii

(Ridley) Whitmore stat. nov.

Basionym of variety: G. curtisii Ridley, Flora 1 (1922) 175, quoad syntypus

Curtis 691 = lectotypus.

G. curtisii Ridley differs from G. bancana, after which Ridley places it in his

Flora, consistently in the fruits with the stigma protruding, more or less lobed

and more or less coarsely papillose. The leaves tend to be narrower and are con-

sistently pointed. It is undoubtedly very closely related and I reduce it to a

variety. The fruits are exactly like those of G. penangiana which, however, always

has its flowers borne terminally; in this variety a few are terminal, but most are

in fascicles behind the leaves, as in G. bancana.

I have selected one of the syntypes as lectotype of my variety. The other,

Curtis 240, is typical G. penangiana.

Collections seen:

PENANG: Curtis s.n. May 1880, s.n. July 1890, 677; SFN 3034, 35322. JOHORE:

SFN 25854. SINGAPORE: Ridley 6194.

I provisionally also place FRI 4/50 from W. Kelantan here, though it differs

in its fruits drying less wrinkled and with sunken stigma.

Garcinia opaca King J. As. Soc. Beng. 59 (1890) 160.

Apparently a rare species. I have not seen the syntype(s) from which King

described the flowers; King’s Collector 5460 at Singapore, which I believe to be

a syntype, is in fruit.

G. opaca closely resembles G. hombroniana but (fide King) has no pistillode

in the male flower, thus it falls in my Group C, whereas G. hombroniana, with

pistillode, falls in Group B. As already noted I am not convinced B and C are

natural groups, although they are very convenient ones. Both these species have

the fruit wall thinly woody, and tending to fracture in the herbarium, this feature

is more marked in G. opaca and especially so in its var. dumosa described below.

Typically, G. hombroniana has a depressed globose fruit with the stigma on a

distinct apical umbo, but sometimes the fruit is slightly elongate. In what I have

Systematy of Malayan Phanerogams XVIII-X XII Zve

described as variety A (see Tree Flora) the fruit is much longer than broad as

in G. opaca, but I am not sure of the true affinity of this variety; its flowers have

yet to be found.

G. opaca var. dumosa Whitmore var. nov.

A varietas typica in habitu plerumque fruticoso, foliis minoribus, ellipticis,

9 x 3.5 - 13 x 5—17 x 7 cm, petiolo graciliore breviore, 8-10 mm, calyce in

fructu parvo demum cadenti, differt.

PAHANG: Bt. Terom, Ulu Keniyam Mohd. Shah 1560 (holotype SING); G.

Benom KEP 97842; Chini F. R. FRI 11066. JOHORE: Bt. Kuing SFN 28653;

S. Berassau, Corner s.n. 18.1V.32; G. Chabang Tiga Jumali & Heaslett 4114;

Mersing, Sg. Sarah working circle, KEP 77731. SELANGOR: Genting Sempah

KEP 1094].

KEP 97842 is a tree, all the others are shrubs, except that the sheet I have

seen (at SING) of KEP 1094] has no note on stature.

Garcinia pyrifera Ridley J. Str. Br. Roy. As. Soc. 73 (1916) 140 (sphalm

‘pyriferum’).

G. densiflora King J. As. Soc. Beng. 59 (1890) 171 (non G. densiflora Vieill. ex

Pierre Fl. For. Cochinch. 5 (1883) 25) syn. nov.

Ridley noted that his species was close to King’s except for certain foliage

differences which he specified. I have compared the type of G. densiflora (King’s

Collector 5933) with that of G. pyrifera (Curtis 3094) and all the subsequent

gatherings of the latter and can see no. good differences, though two of the three

leaves of the Singapore sheet are oblong-ovate, an unusual shape.

In Flora 1 (1922) 179 Ridley reduced King’s species to G. glomerata Vesque

(in DC. Monog. Phan. 8 (1893) 320), which predates his own G. pyrifera. I have

been unable to check this because there is no authentic material of G. glomerata

at Singapore or Bogor. Provisionally therefore I recognise G. pyrifera as the name

for this tree.

The flowers of G. pyrifera remain unknown. The description in the Tree Flora

is taken from King’s of G. densiflora flowers, so far as I could confirm it from

the poor specimen at Singapore.

G. pyrifera is a species with the Gestalt of the other Malayan species of the

Xanthochymus group (G. nervosa, G. dulcis, G. prainiana) and quite clearly dis-

tinct from them in detail.

Garcinia rostrata and its allies (Subgroup Aii)

/ Garcinia rostrata (Hassk.) Mig. Ann. Mus. Bot. Lugd. Bat. 1 (1864) 209.

G. wrayi King J. As. Soc. Beng. 59 (1890) 152, syn. nov.

Garcinia eugeniaefolia Wall. ex T. Anders. Fl. Brit. Ind. 1 (1874) 268.

I provisionally keep G. rostrata and G. eugeniaefolia apart, not having seen

the types of the former, on the stigma differences given by King, which are ‘flat

to concave, smooth’ and ‘flat to domed, papillose’ respectively. The two syntypes

of G. wrayi at Singapore (Wray 267, 362) show it clearly belongs to this aggregate.

Unfortunately all the flowers have fallen; King’s description of the stigma as

278 Gardens’ Bulletin — X XVI (1973)

‘hemispheric, smooth, entire’ (my italics) suggests it belongs with G. rostrata.

Ridley (in Flora 1, 1922, 169) reduced G. wrayi to G. merguensis, but I believe

that is a fairly distinct species (see key in the Tree Flora) on foliage as well as

flowers and that my placing of it is more probably correct. This whole complex,

my Subgroup Ai, needs careful monographic investigation.

Ca Garcinia merguensis Wight Illustrations of Indian Botany (1831) 122.

G. lanceolata Ridley Flora 1 (1922) 170, syn. nov.

I have examined the material at Kew, Kepong and Singapore and believe

these two to be the same.

Mesua

The Mesua assamica group

Mesua aff. assamica (King et Prain) Kosterm.

Kayea assamica King et Prain Ind. Forester 27 (1901) 62.

Symington first recorded this for Malaya when, at Kew in July 1939, he

matched KEP 4643 (sterile), KEP 5495 (fruiting) and possibly KEP 10600 to

H. G. Young s.n. December 1899 and 29 June 1900, two of the syntypes of K.

(M.) assamica. Symington (ms. notes preserved at Kepong) observed ‘“‘These, I

think, differ [from M. ferruginea]. They are not riparian trees and the fruits are

probably smaller than K. (M.) ferruginea. They more closely resemble K. (M.)

assamica K. et P. than anything else but the leaves are less cuneate and acuminate

and the veination [sic] is less conspicuously dryobalanoid”’.

I have placed a whole group of collections here which match KEP 4643 and

5495 in leaf, habit, habitat, and most especially in having the fruit enclosed in

two, strongly adpressed, enlarged sepals. The fruits are long-stalked and solitary

in 5495 but in open panicles in most of the new collections. King and Prain describe

K. assamica as possessing panicles 3-6 in. long. The size of the fruit varies, as

does the sepal surface texture and, slightly, the shape. I attach no importance to

variation in the degree of splitting of the 2 sepals which I believe to be develop-

mental.

I have not seen the types of K. (M.) assamica, and following the caution

expressed by Symington, name our material as aff. assamica. It is slightly hetero-

geneous but is held together by the nature of the fruiting calyx, and, though further

collections may reveal it to comprise more than one taxon, I have little doubt all

these collections are related.

FRI 12033 is an anomalous high altitude collection (780 m,G. Mandi Angin)

with possibly galled young fruits which, on present evidence, comes here; and the

sterile collection FRI 9347, from 960 m on G. Inas, Kedah matches it in leaf.

I have segregated from the above a number of sheets as Mesua sp. D. These

puzzle me not a little. They differ in having smaller leaves with more slender stalk,

and smaller fruits, also more slenderly stalked. Fuller descriptions are given in the

Tree Flora. At the extreme this taxon is rather distinct and KEP 110395, FRI 2617,

8106 approach $21745, 22469, 22507 from Sarawak named by Ashton (unpublished

ms.) as M. acuminatissima (Merrill) Kosterm., but it merges into typical Malayan

M. aff. assamica. Two collections, FRI 885], 10891, have the fruits solitary, the

other collections have panicles.

Systematy of Malayan Phanerogams XVIII-X XII 279

The problems of this species-group can only be finally resolved by a mono-

grapher who will, I suspect, need more collections than yet exist.

Collections seen:

M. aff. assamica

KEDAH: FRI 9347. KELANTAN: FRI 4054, 4099. TRENGGANU: Corner

am. ¢ Now. 35, 12 Nov, 35, FRI 12033. PAHANG: KEP 4643, 5495, 77710.

JOHORE: Kiah s.n. 19 Oct. 35, SFN 25973, 26140, 29272, Lake & Kelsall s.n.,

-28 Oct. 1892. re

M. sp. D. | ;

v 4 4

PAHANG: G. Tapis FRI 10071, 10115, 10891; FRI 8106. JOHORE: KEP

ie FRI 2617, 8851. alt

ih the fuer of flowers or fruits collections of the M. assamica group from

Malaya are very difficult to distinguish from M. ferruginea on morphology. On

the basis of size and habitat the following collections of fallen leaves are put

under the M. assamica group:

M. aff. assamica

TRENGGANU: KEP 81070, 93779. KEDAH: KEP 81329. NEGRI SEMBILAN:

KEP 105202. JOHORE: FRI 5088.

M. sp. D.

SELANGOR: KEP 76378.

Mesua sp. E.

Possibly this is another relative of M. assamica from which it differs con-

spicuously in the shiny leaves with very faint nervation and not or scarcely

reticulate. KEP 94924 has remarkable fruits enclosed in 2 big woody sepals plus,

alternating with them, 2 lanceolate ones. In other respects this collection seems

similar to KEP 76086 with fruits with the type of calyx found in M. aff. assamica.

FRI 8125 is again similar in leaf. Its flowers are larger than in the collections of

M. aff. assamica.

The taxon is known from the east coast as follows:

TRENGGANU: Bt. Bauk: KEP 76086.

PAHANG: Bt. Beserah: KEP 94924; Ulu Sg. Anak Endau: FRI 8125.

Mesua elegans (King) Kostermans Reinw. 7 (1969) 427.

Kayea elegans King J. As. Soc. Beng. 59 (1890) 183 basionym

Kayea caudata King loc. cit., syn. nov.

Mesua caudata (King) Kostermans loc. cit. 426, syn. nov.

The leaves of M. elegans, based on one flowering collection (King’s Collector

7346, K!), differ from those of M. caudata, based on one fruiting collection (King’s

Collector 7937, K!). They are lanceolate, tapering gradually into a pronounced

drip tip, the secondary nerves are faint, and a fine, faint reticulum can be made

280 Gardens’ Bulletin — X XVI (1973)

out with the aid of a lens on the undersurface. The leaves of the type collection

of M. caudata are much broader, taper suddenly into a drip tip, have the secondary

nerves prominent, and the reticulum is almost invisible. In both, the leaves dry

khaki-brown, the twigs are slender and tiny, and subulate ‘stipules’ and hypso-

phylls are present.

The two species are close, but on these collections they can be distinguished,

as indeed King does distinguish them in his key.

The numerous gatherings which have been made since 1885 show a complete

range of intermediates, and I do not believe that more than one species exists.

I have chosen to call it M. elegans because other species of Mesua in Malaya

also have caudate leaves.

M. elegans is in many ways a delicate version of M. kunstleri.

Collections seen (all at KEP and SING except the types):

PERAK: Batang Padang district: King’s Collector 7937 (type of K. caudata),

G. Bubu: King’s Collector 7346 “(type of K. elegans); Keladang Saiong F. R.:

FRI 5998; Parit F. R.: KEP 39492; Tapah: Ridley 14085. SELANGOR: Ulu

Langat: FRI 12203; Klang Gates: KEP 532, 33224, 47132, Ridley 13521, 13527.

PAHANG: Kemasul F. R.: KEP 10608, 10876. JOHORE: G. Panti: KEP

99232. SINGAPORE: Garden’s Jungle: Furtado s.n. 13 I 1931, Ahmad s.n. 10 III

1926.

Mesua kochummeniana Whitmore sp. nov.

Arbor magna, usque ad 36 m alta, 3 m diametro, ligno duro (ferreo); ramulis

crassis angulosis subsparse squamatis, foliis ovato-oblongis, c. 22 x 9 cm, base

rotundatis, apice rotundato-apiculatis, marginibus revolutis, laminis infra glaucis,

supra nitide castaneis, nervis lateralibus approximatis parallelis, ex costa subper-

pendicularibus, leviter curvatis, distaliter incurvato-approximatis arcuatis, petiolo

crasso rugoso c. 12 mm longo. Inflorescentia racemosa, racemis 3—10 terminalibus,

4-7 cm longis, 3-6 floriferis, floribus subapicalibus, pedicellis 5 mm _longis,

bracteolis 1 mm longis oppositis dettoideis; sepalis 4 ovatis 6 mm longis, petalis

caducis (non vidi), staminibus caespitosis exsertis, filamentis tenuibus 7 mm longis.

Fructus (probabiliter immaturi) c. 3 cm diametro, includentibus in sepalis duobus

globosis truncatis dilatatis sublignosisque in base connatis, in sicco rugosis, parte

basali styli crassi (2 mm longo) protrudento.

Type: Corner SFN 29430, 14 May 1935, Johore, 134 mile Mawai-Jemaluang

Road. Holotype in SING.

Also examined:

JOHORE: 13 mile Mawai-Jemaluang Road. Corner s.n. 31 Dec. 1934; 21 mile

Mersing-Kota Tinggi Road. KEP 70811; Jemaluang F. R. FRI 2773, KEP 69992,

KEP 71895. SOUTH PAHANG: G. Lesong F. R. KEP 83464, 86903.

Probably the huge woody fruits 7 cm across (at KEP), collected without

leaves as KEP 79193 from G. Lesong F. R. compt. 20, are this. All the collections

cited except the type are at KEP only and all are sterile twigs or fallen leaves

except KEP 69992 in flower.

I name this fine tree in honour of the Forest Botanist at Kepong Mr. K. M.

Kochummen. It is common in east Johore and the adjacent part of Pahang. Corner.

Systematy of Malayan Phanerogams XVIII-X XII 281

in making the collection I have chosen as type, noted ‘24 hrs of sheer cutting to fell

this tree ... wood exceedingly hard, like iron, as “‘penaga”’’ and that it is common

on hillocks and drier parts of swamp around Sg. Kayu Ara and Sg. Buloh Kasap.

Mesua kunstleri (King) Kostermans Reinw. 7 (1969) 427.

Kayea kunstleri King J. As. Soc. Beng. 59 (1890) 182; basionym.

Kayea rivulorum Ridley J. As. Soc. Str. Br. 54 (1909) 22. syn. nov.

Mesua rivulorum (Ridley) Kostermans loc. cit. 429, syn. nov.

‘King described his Kayea kunstleri in the Materials from four collections:

King’s Collector 3301, 6850, Curtis 1419 and Maingay 176. I have seen sheets of

the first three of these syntypes at Singapore.

I have no doubt that Ridley’s K. rivulorum belongs here, although it is known

only from two syntypes, both in fruit, whereas all the collections I have seen

of M. (K.) kunstleri are sterile or in flower. The argument for this conclusion is

deferred until after the emended description of M. kunstleri var. curtisii below.

In describing K. kunstleri in the Materials King noted ‘a shrubby form of this

occurs in Penang (Curtis, Nos. 805, 1418) and in Perak (King’s Collector, No. 1345)

in which all the parts are smaller and the leaves are less acuminate at the apex,

and rounded instead of much attenuated at the base’. I have seen (also at Singapore)

the two Curtis numbers and confirm King’s description.

In 1894 in another publication, King raised the Penang form to species status

as Kayea curtisii based on one of the sheets cited in the Materials, Curtis 805,

plus another collection Curtis 748, which I have also seen at Singapore, and which

is a good match. He made several serious mistakes in his description in firstly

saying ‘leaf . . . base acute’ whereas it is rounded, secondly ‘flowers . . . leaf

opposed’, whereas they are actually terminal, and thirdly ‘petals slightly smaller

than the sepals’, whereas actually they are twice as long. The earlier note in the

Materials and the syntypes clearly show however what was intended by K. curtisii.

Ridley in Flora 1 (1922) 191 sunk K. curtisii into K. kunstleri commenting that

Curtis 1418 (of King’s original note) is ‘obviously the same plant’ (? taxon) as

K. curtisti, yet retained under K. kunstleri by King.

Since 1890 numerous additional collections have been made, all from Penang

and Kedah. King’s original note is amply confirmed, that there are two distinguish-

able taxa, the extreme forms being very distinct. There are however few substantial

differences, and the placing of a few collections has to be rather arbitrary. Never-

theless I think it convenient and useful to give recognition to the ‘shrubby Penang

form’. I therefore accordingly reduce K. curtisii to varietal status. The necessary

formulation is set out below with a new latin diagnosis, necessary because of

King’s mistakes.

Mesua kunstleri (King) Kostermans var. curtisii (King) Whitmore stat. nov.

Kayea curtisii King Ann. Bot. Gard. Calcutta 5 (1896) 144, t. 174B basionym.

Mesua curtisii (King) Kostermans Reinw. 7 (1969) 426. syn. nov.

A varietas typica in foliis minoribus, basi rotundato, nervis lateralibus abaxialis

obscurioribus, differt.

Lectotype Curtis 805.

282 Gardens’ Bulletin— X XVI (1973)

Although, as noted above, I have seen no fruiting collections of M. kunstleri

I have seen several of M. kunstleri var. curtisii. These have the fruit wall thin

but not brittle, smooth, and irregularly covered with tiny, fawn, corky scales;

the fruits are mostly round, tipped by the 1-2 mm long style base, the largest is

3.3 cm across. The fruit is seated on the persistent sepals which are not enlarged.

Both syntypes of Kayea rivulorum Ridley are in fruit, young in Ridley 7349, and,

in Goodenough 1976, 4.5 cm long and ellipsoid, including a stout 1 cm apical

beak. The wall is wrinkled and bears the same tiny scales typical of M. kunstleri

var. curtisii. These fruits differ from those of M. kunstleri var. curtisii in drying

wrinkled and in their shape. King, in his description of M. kunstleri and later as

plate 173 in Ann. Roy. Bot. Gdn. Calcutta 5 (1896), depicts fruits of this shape,

and there is a single Curtis collection of M. kunstleri var. curtisii at Singapore

with neither number nor date which has them (this sheet I have labelled X).

I consider the resemblance in fruit and leaf adequate to justify reduction of K.

rivulorum. The types of K. rivulorum have the leaf base slightly rounded, whereas

in the types I have seen of K. kunsileri it is long cuneate (and is thus described

in the type description and depicted in plate 173 in Ann. Roy. Bot. Gard. Calcutta

5, 1896). There are 3 collections of Curtis 557] at Singapore, of these 2 have

cuneate and one slightly rounded leaf bases. All are dated March 1901 and all

bear flowers. I think in this case we can assume they come from a single gathering

of one tree, though Curtis often did place different gatherings under one number.

Amongst the other abundant material at Kepong and Singapore this character

breaks down. I do not think the shape of the leaf base is useful in this case to

distinguish two taxa.

M. kunstleri is in many respects a coarse kind of M. elegans with bigger

leaves and flowers. It differs from M. nervosa (in the circumscription of which |

have been assisted by manuscript notes made by Symington at Kew) in the lack

of harsh red tomentum. Monographic study is necessary to delimit more closely

these two species.

Mesua lepidota Anderson FI. Brit. Ind. 1 (1874) 278. KING J. As. Soc. Beng. 59

(1890) 185.

Anderson based M. lepidota on Griffith (K.D.) 845 from Malacca. King added

Scortechini 183b and King’s Collector 4551, 5881, recording that these differ in

smaller leaves, more slender twigs and slightly thinner pericarp. He went on to

note that these collections might be a second species, also possessing the very

remarkable woody reflexed fruiting calyx which had prompted Anderson to suggest

his species was ‘probably a new genus between Kayea and Mesua’.

The abundant gatherings which have now been made show there clearly are

two taxa, but a few collections are intermediate (KEP 32163, 71830, 76734, FRI

11321). It is convenient to give a name to the delicate form especially as it is

very similar in the sterile state to M. nuda; the two are common together in the

Ulu Gombak F. R. None of the cited material is represented in the Far East.

Symington however has matched KEP 12906 to Griffith 845 and KEP 37290 to the

slender variety.

In fact, I believe, from examination of Burn-Murdoch (=CF=FMS=KEP)

382, the single sheet extant of Kayea parviflora Ridley (kindly loaned to me at

Kepong by Kew) that this little known taxon is the same thing in young flower

bud, and it therefore provides the epithet for the variety, as follows:

Mesua lepidota var. parviflora (Ridley) Whitmore stat. nov.

Kayea parviflora Ridley J. As. Soc. Str. Br. 82 (1920) 170, basionym.

Systematy of Malayan Phanerogams XVIII-X XII 283

Mesua parviflora (Ridley) Kostermans Reinw. 7 (1969) 428 syn. nov.

Type: Burn-Murdoch 382-(at K).

Also examined (at KEP and SING):

PERAK: FRI 11937. TRENGGANU: Corner s.n. 9 XI 35, SFN 30480. SELAN-

GOR (all Ulu Gombak): KEP 10929, 13064, 12633, 12855, 37290, 93292, 94717,

94718, FRI 1154, Hume 9270. MALACCA: Derry 63, 902, collection o¢ (at SING,

no name or date). JOHORE, G. Blumut: SFN 10790.

In the Perak collection the fruit is young and has not developed a woody

calyx. The Johore collection has the leaves 5 X 1 cm and broadest near the base

with a long narrowed upper portion. These collections may represent slightly

divergent populations. The Trengganu collection, SFN 30480, is the only one

I have seen of the variety with the flowers in racemes.

The collections I have examined suggest the typical variety inhabits plains,

and var. parviflora is a tree of the hills. The four intermediate collections cited

all come from hilly places.

Mesua nivenii Whitmore sp. nov.

A M. wrayi valde simillime sed in floribus sessilibus et foliis infra conspicue

reticuloso-venulosis differt.

Arbor parva, foliis ovato-oblongis, c. 9 X 4 cm, basi late-cuneatis, apice acutis,

coriaceis, nitidis, nervis lateralibus subtus tenuibus prominulentibus; pagine infra

reticulato, nervis tertiariis valde approximatis, petiolo 1 cm longo ruguloso.

Inflorescentia racemoso-paniculata, racemis ramificatis, pedunculis 1 cm _ longis,

pedicellis obsoletis (floribus sessilibus) trifloriferis, sepalis ovatis rotundatis con-

vexis in sicco nigris rugosis 6 mm longis, petalis c. 10 x 6 mm obovatis rotun-

datis, staminibus exsertis caespitosis. Fructus (submaturus?) includentibus in sepalis

duobus paribus, sepalis maturitate incrassatis coriaceis rugosis turbinatis, parte

basalari stylo 2 mm protrudento.

Type: Whitmore FRI 12100, 12 July 1968. Trengganu, G. Mandi Angin, Summit

ridge north of birop (trig. point), 1425 m. Holotype in KEP, isotypes K, L, SING,

ARN. In flower.

Also examined: a,

TRENGGANU: leading E. ridge G. Mandi Angin: FR/ 1081/5; G. Padang SFN

31064, 33909 (in fruit). |

First collected by Moysey and Kiah on their pioneering journey up the Ulu

Brang to G. Padang in 1937. Found in full flower by myself, and a delightful

sight, on the summit ridge of G. Mandi Angin in 1968. Named for the part-time

Superintendent of the first Singapore Botanic Garden who ‘had a fine eye for

design’ (Purseglove).

Mesua nuda Kostermans ex Whitmore sp. nov.

Arbor, ramulis gracilibus teretibus plerumque griseis, foliis ellipticis (5 x 1.5)

— 6 x 2—8 x 3 cm, base cuneatis, apice cuspidatis, subcoriaceis, nervis laterali-

bus utrinque tenuissimis approximatis parallelis leviter arcuatis ex costa subper-

pendicularibus, petiolis gracilibus 3 mm longis. Flores (in alabastro) cum pedicellis

2 mm longis. Fructus solitarius, axillaris, pedunculo incrassato (a 5 mm), pedi-

cello 2 mm longo, maturitate oblongo-ovoideis 4 cm longis 2.5 cm latis, apice

interdum rostellatis, base rotundatis, paricarpio fructi tenuoso-fragile bubalino laeve

squamulis minutis suberosis obtectis, sepalis caducis. Semen unicum in fructu, testa

fusco nitido.

284 Gardens’ Bulletin — XXVI (1973)

Type: Whitmore FRI 3440, 24 March 1967. Pahang, G. Benom, Ulu Sg. Krau,

ridge crest 270 m. Holotype in KEP, isotypes K, L, SING, ARN.

Also examined:

KELANTAN: KEP 98807, FRI 2333, 2339, 7255. TRENGGANU: SFN 30390,

FRI 10682. PAHANG: KEP 4059, 29682, 100105, 108974, FRI 15333. Baloh

F. R.: KEP 3149, 97913, FRI 3798. KEDAH: KEP 105354. PENANG: Curtis

748. PERAK: KEP 29928; s.n. Wyatt-Smith 25 Jan. 1953. SELANGOR: Hashim

s.n. (SING), Burn Murdoch (KEP ?) 278 = 14248. Ulu Gombak F. R.: KEP

10942, 11220, 18163, 18165, 98401, FRI 2555. MALACCA: (illegible) 85/,

Alvins 1066, Derry 1003 no locus; Symington KEP 18163. NEGRI SEMBILAN:

KEP 104871, FRI 11347. JOHORE: KEP 69996.

Mesua nuda is a very common species, apparently endemic, which unfor-

tunately has never been found in full flower. It is highly distinctive in its brittle,

buff, naked fruits. The name is in wide usage but as far as I can discover Koster-

mans has never published it. Young fruits are sometimes globose, leaf size is

slightly variable as is the clarity of the secondary nerves, but I have little doubt

that all the collections cited belong to this species.

Mesua purseglovei Whitmore sp. nov.

Arbor excelsa, foliis coriaceis lanceolatis, 13 x 4 cm, lamina prope basin quam

apicem latiore, base rotundato, apice acuminatissimo; nervis lateralibus majoribus

ex costa subperpendicularibus parallelibus leviter curvatis distaliter incurvato con-

junctatis subtus conspicuis supra inconspicuis subdepressis, nervis lateralibus

secundariis minoribus (1-3 aggregatis cum majoribus alternantibus) subtus incon-

spicuis, nervis tertiariis (reticulationis) evidentibus. Flores ignoti. Fructus solitarius

(variter geminatus), axillaris, pedicello crasso 1 cm longo, pedunculo 4 cm longo,

in sepalem '+ inclusus (sepalis 4 maturitate valde dilatatis ovatis subcoriaceis

furfuraceis depresso globosi diametro 3-3.5 cm).

Type: Holttum SFN 31233, 14 May 1936, Pahang, Cameron Highlands, mile 47

Telom Rd. Holotype in SING.

This is a high mountain species and I think likely to prove to be endemic,

I therefore describe it, despite its being known from only one collection, in fruit.

M. purseglovei is similar to M. wrayi but conspicuously differs in leaf characters.

XIX. LECYTHIDACEAE

T. C. WHITMORE

Barringtonia payensiana Whitmore sp. nov.

Species nova, a speciebus ommibus generis Barringtoniae in fructibus pyrami-

dalibus differt.

Arbor parva, foliis obovatis acuminatis chartaceis vel subcoriaceis, c. 21 x 6.5

— 50 x 14.5 cm, base longe decurrente, marginibus leviter revolutis, apicem versus

leviter dissitiserratis, nervis lateralibus in paribus c. 26, petiolo 15 mm. /nflores-

centiae et flores ignotae. Fructus pyramidata usque ad 4.5 cm lata, c. 4.5 cm longa,

base omnino plano, plus minusve quadrata in sectione, superficie ruguloso, calyce

persistenti usque ad 15 mm diametro.

SELANGOR: Ulu Batang Kali FRI 4550 (holotype SING).

PERAK: Pangkor Island SFN 31658.

Named in honour of the recent monographer, Dr. J. P. D. W. Payens.

Systematy of Malayan Phanerogams XVIII-X XI] 285

XX. LEGUMINOSAE (d)

T. C. WHITMORE

Crudia viridiflora Whitmore sp. nov.

Arbor, foliis simplicibus subcoriaceis, glabris, oblonge-ovato, 15 X 6 — 22 X 8

cm, base cuneato, apice rotundato, apiculato, costa supra canaliculata, nervis

lateralibus c. 9 paribus, pagina inferiore valde prominentibus, pagina superiore

obscuris, nervis lateralibus secundariis reticulisve similiter, petiolo crasso rugoso

5 mm. Flores congesti, viridi, in racemos terminales, solitarios vel — 4 fasciculatos,

7-12 cm longi, rhachidi aureo pubescenti, bracteis minutis aureo tomentosis, pedi-

cello 3 mm aureo tomentoso, sepalis ovoideis 6 x 4 mm exteriore ad basim leviter

tomentosis, staminibus 8 exsertis, filamentis filiformibus 10 mm, ovario 5 mm

longo aureo lanato, stylo gracili 5 mm. Fructus ignoti.

PAHANG: (G.) Lesong F. R., Sg. Jekatih FRI 15949 (holotype KEP); Samsuri

540.

This distinctive new species was discovered too late for inclusion with the

Leguminosae in vol. | of the Tree Flora and will have to go into a supplement

later. It comes near to C. gracilis Prain apud King and C. subsimplicifolia Merr.

in De Wit’s monograph (Bull. Jard. Bot. Buitenz. sér 3, 18, 1950 407-34) but

differs from both. It can easily be distinguished from all the other Malayan species

from its simple leaves. I did not see any compound leaves on the limbs which

were cut for me from the crown of the tree from which the type collection was

made.

XXI. STROMBOSIA (OLACACEAE)

T. C. WHITMORE

Strombosia maingayi (Masters) Whitmore comb. nov.

Anacolosa maingayi Masters FI. Brit. Ind. 1 (1872) 580 basionym.

Strombosia rotundifolia King J. As. Soc. Beng. 64 (1895) 103 syn. nov.

Both names are based on the same type sheet, Maingay (Kew Distribution

No. 1019), and the former clearly has priority. When King transferred Masters’

species to Strombosia he incorrectly coined a new epithet for it. Ridley (Flora 1,

1922. 425) cites A. maingayi as a synonym of King’s species.

Strombosia multiflora King J. As. Soc. Beng. 64 (1895) 102: syntypes Perak, King’s

Collector 7824; Penang, Curtis 859.

King described this and his S. rotundifolia (see above) at the same time,

noting that they are closely related. The name S. multiflora has never entered

currency. The abundant material at Kepong and Singapore appears very homo-

geneous. I can see in it no significant difference in the three differential characters

given by King, namely the number of flowers per cymose fascicle (the pedicels,

it should be noted, are brittle and flowers easily break off), flower size (all have

tiny flowers only c. 3 mm long, on a 1.5 mm pedicel), or leaf shape and size

(King’s contrast of rotund, 2.5 x 2 in., versus 2.5 x 1.5 — 4 x 2 in. seems in any

case to be weak). There is variation between collections with sessile, round to

oblong, fruits and those with stalked, round fruits. The former group has small,

very woody fruits which might perhaps be galled.

286 Gardens’ Bulletin— X XVI (1973)

Mr. L. L. Forman has kindly made a very thorough examination of the Kew

material of S. maingayi (rotundifolia) and S. multiflora. Not one collection is in

fruit. He could see no leaf or twig differences and the material was too incomplete

to study the buds. He did however find a substantial difference in the flowers,

along the lines indicated by King; to quote ‘the Kew isotype of Str. multiflora

has dense bunches of flowers with petals 5 mm long and styles 3.5 mm. Srr.

rotundifolia on the other hand has petals only half that length and a very short

style less than 1 mm... These species are surely distinct’. He later reported that

in both species the flowers have 1.5 — 2.0 mm pedicels. The Singapore isotypes

which had been mislaid were subsequently rediscovered, examination of them

confirmed Forman’s careful observations.

The problem facing us in Malaya is thus a very tricky one. On the one

hand, the two species appear to differ substantially in flower but the types are

inadequate to discern any other differences. On the other hand, all the collections

im the East (many of them duplicated at Kew) appear to represent one species,

except Curtis 1371 from Pangkor Island, but they are mostly sterile or in fruit.

That some collections have sessile fruits does not help as both species have

pedicellate flowers, and in addition such fruits are suspected possibly to be galled.

Since King described S$. multiflora no further collections have been added to

the species, except Curtis 1371, just mentioned, which was misnamed ‘? Anacolosa

sp.’ Ridley in his Flora cites just the two syntypes. All the abundant material at

Kepong and Singapore has been named S. rotundifolia (=maingayi). The few

flowering collections amongst this abundance have the tiny flowers and short

stigmas of S. maingayi, and are a good match for the fruiting and sterile collections.

I conclude that the pragmatic solution is to name all these collections S.

maingayi. Although S. multiflora differs conspicuously in flower more collections

than have been assembled in the last 76 years will be needed to discover if these

two species can be differentiated in any other way. Thus, in summary, I propose

to follow the practice since 1895, in putting our material into S. rotundifolia, but

have had to revert to the earlier epithet for this taxon which becomes therefore

S. maingayi.

Curtis 1371 from Lumut, Pangkor (Perak) at Singapore might represent S.

multiflora in that the style is very long (4 mm) and thread-like, but the flowers

are only 2 mm long.

XXII. HELICIOPSIS (PROTEACEAE)

K. M. KOCHUMMEN

Three new species of this small genus, only differentiated from Helicia in

1955, have been discovered in preparing the account for the Tree Flora.

Heliciopsis cockburnii Kochummen sp. nov.

Arbor ad 24 m alta, 60 cm diam., ramulis striatis junioribus ferrugineis, foliis

(foliis pinnatisectis adhuc ignotis) ellipticis vel anguste obovatis, 9.5 x 2.8 — 21 x

7 cm, apice obtuso, base angustato decurrenti, nervis lateralibus 6-11 paribus

arcuatis, petiolo 1-4 cm longo base incrassato. Flores adhuc ignoti. Infructescentiae

10 cm longae rhachidi striata rubiginosa gracili 2 mm diam., bracteis linearibus

6 mm longis rubiginosis sub-persistentibus, bracteolis bracteis similibus 3 mm

Systematy of Malayan Phanerogams XVIII-X XII 287

longis, pedicellis 4 mm longis 2 mm crassis. Fructus oblonga nitida siccitate brun-

neola 2.5 cm longa 2 cm diam., stylo vestigio, exocarpio tenuissimo, mesocarpio

radiato fibroso, 3 mm crasso, endocarpio tenuissimo. Semina leaves, testa brunneola.

PAHANG: G. Tapis 800 m FRI 10934 (holotype KEP). PERAK: Keladang

Saiong F. R. KEP 2900, KEP 32181. UPPER PERAK: FRI 594.

Heliciopsis montana Symington ex Kochummen sp. nov.

Arbor 26 m alta, 60 cm diam., ramulis striatis cum cicatricibus foliorum

delapsorum notati, foliis simplicibus coriaceis, ellipticis vel anguste oblongis,

(9 x 3.5) — 15 x 6— 21 x 10 - (40 x 20) cm, apice obtuso, base cuneato raro

inaequilatero, nervis lateralibus 6 — 10 (- 14) paribus arcuatis, pagina supra nitida,

petiolo 2 — 4.5 cm longo, ad base gibboso, foliis pinnatisectis 48 x 24 — 54 x 27

cm, lobis 3-5 paribus oblongis 17 x 3 — 19 x 7 cm, apice acuto, nervis lateralibus

9-11 paribus arcuatis, petiolo 10-13 cm longo. Flores adhuc ignoti. [nfructescentiae

rhachidi crassa 5 mm diam. striata ferruginea, bracteis adhuc ignotis (? cadicis),

pedicellis crassis 6 mm longis 3 mm diam. Fructus siccitate purpurea vel brunneola

oblonga 2.8 — 3.8 cm longa x 2.5 — 3.5 cm diam., exocarpio coriaceo tenuissimo,

mesocarpio fibroso 3 mm crasso, endocarpio tenuissimo leavis.

PAHANG: Fraser’s Hill FRI 16172 (holotype KEP). Cameron Highlands KEP

27194, 30966, SFN 32871. PERAK: G. Korbu KEP 32248.

A species of mountain forests at 300 m and above. Endemic. Symington

named this Helicia montana in manuscript.

Heliciopsis whitmorei Kochummen sp. nov.

Arbor 18—21 m alta, trunco usque ad 40 cm. diam., ramulis striatis, junioribus

ferrugineis, foliis simplicibus coriaceis ellipticis vel oblongis 7.5 x 2.5 -— 21 x7

cm, siccitate viridiflavis, apice acuto, base acuto, nervis lateralibus 9-12 paribus,

arcuatis, utrinque inconspicuis, costa supra conspicua, petiolo 2-5 cm longo base

incrassato, foliis pinnatisectis 46 x 28 — 60 x 30 cm, lobis c. 11 paribus, oppositis,

12.5 x 2—15 x 3 cm, petiolo 9-14 cm longo. Flores adhuc ignoti. /nfructescentiae

15 cm longae, rhachidi striata glabra 3-4 mm diam. Fructus cylindrico-truncata

18-20 mm longa 8-11 mm lata, stylo persistenti 1 mm longo, exocarpio tenuissimo.

mesocarpio radiato fibroso 1 mm crasso, endocarpio 0.3 mm crasso.

TRENGGANU: G. Mandi Angin FRI 12062 (holotype KEP). Ulu Sg. Trengganu

900 ft FRI 10544. Ulu Bendong Kemaman, 700 ft SFN 30/93.

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A survey of the occurrence of epiterranean soil algae

in Singapore Island

ANNE JOHNSON

Department of Biology, Nanyang University

Singapore

Epiterranean soil algae found in the upper layers of soil are of considerable

importance in soil economy since they increase the organic content of the soil by

photosynthesis, disintegrate soil particles to increase the water-holding capacity of

the soil and secrete mucilaginous material binding surface particles together to

form a matrix; while heterocystous blue-green algae fix atmospheric nitrogen

increasing the nitrogen content of the soil (Johnson, 1962). Subterranean soil algae

which live in complete darkness and may lose their photosynthetic pigment (Tiffany,

1951) are less important to soil economy.

Few investigations have been made on the soil algae of tropical areas and

include those of Fritsch (1970) in Ceylon; Prasad (1949), Gonzales and Valavigi

(1960), Singh (1940-41), Sulaiman (1944) and Ghose (1923) in India; Bristol (1919)

- in Malaysia and Johnson (1962, 1969) in Malaysia and Singapore.

Over a large area of Singapore Island the soil is derived from sedimentary

parent rocks of the Triassic or Carboniferous period, with small areas derived

from igneous rocks of acid or basic composition, as well as alluvial soils in swamps

or valley bottoms. Most areas in the West of the Island the soil was derived from

argillaceous series including red iron-rich strongly lateritic soil, light coloured to

white iron-poor shale-derived soil, as well of soils mixed composition (Panton,

1955). Sedimentary soils of arenaceous series are commoner in the East of the

Island including both coarse and find-grained series. The red colour of laterized

soils is due to their high iron content since laterisation leads to an accumulation

of iron and aluminium in the upper layers (Becking, 1961). Most of the iron is

unavailable to the plants being in the form of insoluble iron-aluminium sequioxides

or ironstone concretions. Most soluble cations such as calcium, magnesium, potas-

sium and sodium have been leached away, and Singapore soils are extremely

deficient in cations. The soils are also low in available phosphate due to loss by

leaching and have a rather low pH. The rapid decomposition of organic matter

under tropical conditions and subsequent leaching make them typically moderately

low in nitrogen.

Singapore Island was originally covered with tropical lowland forest with

dipterocarps as the dominant type of trees. During the second half of the nineteenth

century considerable areas of the Island were cleared for the planting of gambier

and pepper (Jackson, 1968). The soil rapidly became exhausted leaving a very

infertile soil. When gambier and pepper cultivation was abandoned early this

century, the land was allowed to revert to grassland and scrubland (belukar). In

other areas the top soil has been removed in reclamation operations and exposed

289

290 Gardens’ Bulletin — XXVI (1973)

subsoil (B layer) remains. Forest is extremely limited in extent. Cultivated areas

support vegetable crops, and estates have rubber, coconuts, rambutans and other

fruit. Disturbed areas are city dumps, abandoned building sites, army camps etc.

Materials and Methods

Surface soil was collected from 100 habitats in the rural area of Singapore

to a depth of 2 cm. Two series of cultures were set up for each type of soil: —

(i) Plate culture, consisting of soil placed to a depth of 1 cm. in sterile

petri-dishes, moistened with sterile distilled water and

(ii) Liquid culture, consisting of small quantities of soil introduced into

sterile Knop’s solution in conical flasks under sterile conditions. Both

types of culture were exposed to 12 hours natural light per day. Observa-

tions on the algae appearing in the cultures continued for two months.

After setting up the cultures the remaining soil was air-dried and sieved

through a 2 mm. sieve before analysis. The soil type was characterised into clay,

clay-loam, loam, sandy-loam, loamy-sand, sand and sandy-clay according to the

method of Leamy and Panton (1966). The soil was analysed for percentage of

water, percentage of organic matter, ppm. nitrate, phosphate, calcium, ammonium

nitrite, magnesium, iron, aluminium, sulphate, chloride, potassium and carbonate

using the methods of Jackson, 1958 and those of the Edwards Laboratory, Ohio.

pH was determined by use of a pH meter on soil paste. Colour was determined

using standard Munsell Colour Charts to give hue, colour value and chroma on

both wet and dry soils.

Results

1. Soil type

The soil type characterised according to Leamy and Panton (1966) in different

habitats is seen in Table I. Surface soil consisted predominately of clay (24%) or

clay loam (39%). Sandy loam and loamy sand amounted to 10% and 13%

respectively. Sands (6%). sandy clays (2%) and loams (6%) are rare. Forest soil

is usually clay loam (66.7%) but clays and loams occur. Alluvial soil is usually

clay (57.2%) with some occurrence of clay loam, loam and sandy loam. Exposed

subsoil (B-layer) is usually clay or clay loam with rare occurrence of loamy sand

and sand. Grassland may be found on all types of soil. Scrubland (belukar) occurs

on all except sand and sandy clay. Estate soil (rubber, banana, rambutans etc.)

included no example of clay or sandy clay; while disturbed soil (dumps etc.) may

be found on clay, clay loam or sandy loam.

Note. In the tables the following abbreviations are used for different

habitats : —

F = forest soil; A = alluvial soil; E = exposed sub-soil (B-layer):

G = grassland: S = scrubland (belukar); C = cultivated soil:

Est. = estate soil; D = disturbed soil (city dumps etc.).

Samples were collected in the Jurong, Bukit Panjang, Serangoon and Katong

districts of Singapore.

Epiterranean soil algae in Singapore 291

TABLE |

Percentage samples with different types of soil

CLAY ss mae J «Sor JE de ee A 333 Bs 16.7 24

ool a EES CS 5 ae a

2. Soil Analysis

The percentage water in air-dried soil, percentage organic matter; and nitrate,

phosphate, calclum, ammonium, nitrite, magnesium, iron, aluminium, sulphate,

chloride and potassium in ppm. are given in Table 2. No soluble manganese was

detected by the methods used. The percentage of soils containing carbonate is

given in Table 3.

The water holding capacity of dry soil is closely correlated with the organic

content being high in forest soils and alluvial soils but low in all other soils.

Nitrate and calcium levels were normally at moderate to moderately low levels

especially in alluvial or cultivated soils but all other ions were at extremely low

levels. The values for phosphate, sulphate and potassium are critically low in most

souls.

Most soils are acid, forest soils having an average pH of 3.83 with the pH

somewhat higher in other types of soil especially in disturbed soil where an

average pH of 6.45 was observed. This is correlated with the higher levels of

potassium, calcium and carbonate in this type of soil.

Carbonate was detected in only 17% of the samples. It is completely absent

from alluvial and estate soil, but occurrs in 83.3% of disturbed soil (dumps etc.).

There are no calcareous rocks on Singapore Island.

292 Gardens’ Bulletin — XXVI (1973)

TABLE 2

Soil Analysis. Mean values for different soils

°% water (dry soil) .. | 6.29 |10.81 | 1.47] 1.54 | 233 | 192] 109] 145 2.53

% organic matter .. | 18.09 |12.62 | 4.95 | 7.66 | 6.29| 7.17| 5.19 | 647| 7.18 _

NO3 (ppm.)__.. g 8.6 | 14.16 | 6.00] 9.56 |11.00 |21.11 | 13.60 | 17.50 | 10.04

PO, x .. | 0.16 | 1.08 | 0.23 | 0.56 | 0.70| 1.33 | 1.50| 0.33 | 0.66

ca [1000/1166 |14.12 [1437 | 8.00 |13.30 12.70 /1330 13.08

NH4 2.33 | 0.67 | 0.70 | 1.87 | 2.40 | 3.50 | 3.60 | 233 | 1.95

pH 3.83 | 4.88 | 4.95] 5.80| 5.00] 5.67) 5.40, 6.45| 5.26

NO» EEE 0.09 | 0.17 0.08

Mg 0.66 | 4.83 | 1.88 | 2.31 1.80 | 2.66 | 3.70 | 3.00) 2.41

Fet* 1.66 | 1.00 | 1.65 | 0.65 | 0.80 | 0.89 | 1.80 | 0.33 1.02

Al = o4a7 oa; Pia ome

Mn Lee, H162 url lt arshly af tiuecSlelvmecia |satinal aanaeen nnn

SO, t, vol T8667) 333 [Poss Plo) oo go gee nae

Cl ‘ |... 3.33 | 1.76 | 003 | .. | O01) 400 ee

K pay 1.50 | 1.67. 5.00 | 0.60

Colour value (dry) is 80.1 53 167 (ee 1 oe Tae 6.1 63 | 5.9

(wet) | 33 | 40°R4N 42 | 43 | 42 |. 460) aOul ee

Chroma (dry) .. -- 28 | 43 | 49 | 36 | 26) | ae) ) aie 3.5

(wet) .. Serer ee eee ne me pe 4. | 3.2

TABLE 3

Percentage Samples Containing Carbonate

| of = f | | fF OC

+ CO3 Sa ee ey 7 11.8 9.4 | 30 35.3 is $3.3-], AE

—'CO3 ne .- | S30 | 100 | "66.2 | 506) 70 66.7 | 100 | 16.7 83

3. Soil Colour

The colour values in Table 2 reflect the somewhat lower values of alluvial

and forest soil, and the relatively higher values for exposed and estate soil. The

chroma is lowest in forest soils. The hue of wet soils all occurred on the YR

colour range of the Munsell colour charts (Table 4). In spite of the uniformity of

hue with 60% of the soils in the 10 YR range, Singapore soils are very variable

in colour (Table 5).

Epiterranean soil algae in Singapore

Hue

10 YR

5 YR

2.5 YR

TABLE 4

Distribution of hue in 100 samples (wet soil)

———— | fl | | | | EE

TABLE 5

293

Number of soils with different surface colour (Standard Munsell terminology-wet soil)

Strong brown

Dark brown

Brown

Pale brown

Very pale brown

Very dark greyish brown

Dark greyish brown

Greyish brown ..

Dark Reddish brown

Reddish brown ..

Light reddish brown

Dark yellowish brown

Yellowish brown

Pale yellowish brown

Black

Very dark grey ..

Dark grey

Red

Yellowish red

Pink

Reddish yellow ..

Brownish yellow

Light yellowish brown

ee Nee

2 1 Z

ibe to pdranfuegedeino>| 2 | 3 | 2

hartge Yo <0g Ipr0t g eajamege Tp og | voqeifrog | 4

—}—}—- +--+

————}—

mandrel treat siete dort ee bo le

per Oh Bent hops Hr aiibeoo Pay

(pS ogy Te ge gee a mela ge a yea ind

||}

eee Te incr sats athe Ud ieee dedinad

—- +--+ —

—}—|—} |_| — —- —

Ae a a

+ ——

sj}

meme iataiiih } |

||| +

oe ee ae a

——}—}——_ ——

| 2 ae ae ee

| | — —_-—__—_

com ee | Goan a

Total

—) pl

294 Gardens’ Bulletin — X XVI (1973)

4. Soil Algae

(a) constancy of occurrence. The appearance of algae in sterile culture indicates

the presence of these algae, at least as resting spores, in the soil. As has been

reported frequently, the frequency of algal species in culture does not necessarily

reflect their frequency in the soil since the conditions of culture may favour some

species more than others. Although frequency estimates of growth in culture were

made, these were considered to have no direct relationship with the actual frequency

in the soil and these results are not reported here. The constancy rather than the

frequency of occurrence of algal species was considered to be of greater significance

since it gives a measure of the number of soil samples in which the algal species

occurs.

80 species of algae were recorded in the cultures and they were distributed

with constancies shown in Table 6. The soil examined show a great diversity of

algal flora with very few constant species. The typical J-shaped curve of constancy /

no. of species is common in tropical floras (Johnson, 1966).

TABLE 6

Contancy and number of species

(Constancy = no. of samples; total no. of samples = 100)

Constancy ..| 85 | 29 | 19 | 17 7 6 5 4 a] 2 | 1

mi | | | LE LN TT EE

No. of species .. 1 1 1 1 5 1 1 5 7

Total no, of species = 80

Table 7 gives the habitat occurrence of species with a constancy of more than

5% (only 11 species have a constancy of more than 5%). Although the list includes

5 species of blue-green algae (with asterisk) none of these are capable of fixing

nitrogen under aerobic conditions.

TABLE 7

Number of Occurances of Algal species with a constancy of more than 5%

a | F A EB G Ss * Est. D Total

DESMOCOCCUS VULGARIS

Brand. 74 a 5 5 15 30 a 8 10 3 85

HORMIDIUM FLACCIDUM

(Kitz.) Br; 0 1 52 13 0 1 1 1 19

* OSCILLATORIA JASORVENIS

Vouk. ori antares per pa po ae

*OSCILLATORIA OBSCURA

PALMELLA MINATA Nag. .. 7 1 7 10 3 2 4 29

Briihl. et Biswas

Epiterranean soil algae in Singapore 295

TABLE 7 — continued.

Number of Occurances of Algal species with a constancy of more than 5%

Roo” E G S Cc Re om | Tent

*SYNECHOCOCCUS AERUGINOSUS | |

Nag. a2 ars 0 0 4 2 0 1 OOP rian 7

TROCHISCIA ASPERA (Reinsch)

Hansg. uO eh 1 0 0 1 ee 1 i 7

PINNULARIA sp. bis 0 2 0 3 0 0 2 0 7

NAVICULA sp. .. "a 0 0 2 1 0 1 1 a 7

* CHROOCOCCUS MINUTUS

(Kiitz.) Nag. & 0 ys Z 0 0 0 2 0 6

(Kiitz.) Nag. he 0 0 1

* CHROOCOCCUS TURGIDUS

(b) Blue-green algae. As in other tropical soils, blue-green algae (Cyanophy-

ceae) form a conspicuous part of the algal flora. In this investigation 41 species

of blue-green algae are reported from Singapore soils. These may be divided into

the non-heterocystous algae which do not normally fix nitrogen under aerobic

conditions, and the heterocystous algae which are capable of nitrogen fixation.

Of the 26 species of non-heterocystous blue-green algae (Table 8) none are

found in forest and only 3 in estates. They are characteristic of open areas such

as exposed sub-soil, grassland and scrubland where the light-intensity is high.

TABLE 8

Number of Occurrences of non-heterocystous blue-green algae

——- F A E G S . Est D Total

CHROOCOCCUS MINUTUS

(Kiitz.) Nag. 2% 2 Z | a 6

C. TURGIDUS (Kiitz.) Nag. .. 1 3 1 5

ee ee ee

C. SCHIZODERMATICUS West 1 | 1

CHROOCOCCUS sp. 3% 1 1 | 2

MICROCYSTIS AERUGINOSA |

Kiitz. ee “fe 1 E 2

CHAMAESIPHON FUCUS |

(Restaf.) Hangs. a 1 | | 1

OSCILLATORIA JASORVENSIS

Vouk. ae a 2 11 3 1 17

296 Gardens’ Bulletin — X XVI (1973)

TABLE 8 — continued.

Number of Occurrences of non-heterocystous blue-green algae

| |

or ae E G S C | Est. , D | Total

O. osscurRA Briihl. et Biswas 1 1 ‘a ah | | 1 7

| eee Se SS | |

O. ANNAEA Van Goor ae | 1 1 zm | | 4

Ipeut | reel os esd Sites cx at 220

O. CHALYBYA (Mertens) Gom. | | 1 3 | | ga

vr or ah

O. FREMYI De Toni 2 1 3

O. MINIATA (Zanard) Hauck 1 1 1 3

: a as oe i nn eM —_— es

O. TEREBRIFORMIS Ag. | 1 1 | )

ene ere 05 |

O. CHLORINA Kiitz. ). 1 1 2

| i ees ee | ee Sad

} |

O. OKENI Ag. .. a 1 1 | | | 2

O. Limosa Ag. .. on ] 1 | eek:

O. SUBTILLISSIMA Kitz. .. 1 | 1

O. GEITLERIANA Elenkin. .. 1 | | | 1

ideiaccunctisiriwe-satdedinaieiein waeeltealat ane

SYNECHOCOCCUS AERUGINOSUS |

Nag. re J Z | 1 | 7

ARTHROSPIRA TENUIS Stikenb. 1 | 1 | 2

ret ei |

SPIRULINA LABYRTHIFORMIS |

(Menegh.) Gom. | 1 | | 1

PHORMIDIUM TRUNICOLA Ghose | 1 | 1

PH. SUBINCRUSTATUM Fritsch

and Rich 4: fh 1 1

LYNGBYA POLYSIPHONAE a a 1 | 1

Frémy a |

| 1

L. BIRGEI Smith .. a 1 |

TOTAL NO. OF SPECIES 0 7 15 10 | >

Epiterranean soil algae in Singapore

297

Table 9 shows the occurrence of 16 heterocystous blue-green algae which are

capable of nitrogen-fixation. They occur in just over a quarter of the soil samples.

None are found in forest soil but at least one species of nitrogen-fixing algae has

been found in all other types of habitat. Their occurrence is sporadic in these

habitats and nearly three quarters of the samples showed no evidence of the

presence of nitrogen-fixing algae.

TABLE 9

The Occurrence of heterocystous blue-green algae

ANABAENA FERTILISSIMA Rao

ANABAENA SPIROIDES Kleb. ..

ANABAENA FUELLEBORNII

Schmidle

ANABAENA ORYZAEA Fritsch

ANABAENA ANOMALA Fritsch

ANABAENA SP.

NosTOC MICROSCOPICUM Carm.

N. MUSCORUM Ag.

N. | PUNCTIFORM

(Kiitz.)

Hariot ie id

TOLYPOTHRIX PHYLLOPHILA

West :

SCYTONEMA HOFMANNI Ag.

CYLINDROSPERMUM SPHAERICA

Prasad : -_

AULOSIRA AERIGMATICA Frémy

HAPALOSIPHON WELWITSCHII

West bs

NOSTOCOPSIS RADIANS

Bharadn.

WESTIELLOPSIS PROLIFICA

Janet :

TOTAL NO. OF SPECIES

F A | E G S c Est. D Total

1 1 1 3

1 1 2

1 1

1 7 ee

1 1

1 1 2

1 1

2 1 3

0 3 z 7 1 3 2 3 16

298

Gardens’ Bulletin — XXVI (1973)

ceae, Euglenophyceae) is seen in Table 10. The highly constant species Desmococ-

cus vulgaris, Palmella miniata and Hormidiun flaccidum are widely distributed, but

most of the 31 species are very sporadically distributed and include some species

as Gonium pectorale and Euglena elongata which are characteristic of freshwater

habitats.

TABLE 10

(Cholorphyceae, Xanthophyceae and Euglenophyceae).

DESMOCOCCUS VULGARIS Brand.

PALMELLA MINIATA Nag.

HORMIDIUM

FLACCIDUM

(Kiitz.) Br. my

TROCHISCIA ASPERA (Reinsch

Hansg. a *

MURIELLA TERRESTRIS

Petersen

EUGLENA ELONGATA

Schewiakoff ..

CHLAMYDOMONAS Sp.

SCENEDOSMUS FLEXUSOS

(Lemm.) Ahlstrom

CYLINDROCYSTIS sp.

BUMILLIERA EXILIS Kleb

SPONGIOCHLORIS

(Vischer) Starr

SPONGIOSA

MICROSPORA SP.

GONIUM PECTORALE Miill ..

OEDOCLADIUM Sp.

EUASTRUM TORUM

BINUCLEARIA TECTORUM

(Kiitz.) Beger

LEPTOSIROPSIS TORULOSA Jao

DICTYOCHLORIS FRAGRANS

Vischer oe

ZYGOGONIUM Sp.

TRENTEPOHLIN AUREA (L.)

Mart. by ji

STIGEOCLONIUM Sp.

ANKISTRODESMUS

FALCATUS

(Corda) Ralfs ;

eae es ae Re en

Peete Je 8 ee:

ia 278 Ldn dreabard

1 C20) fee

4 |

1 1-1—4

1 a

3 1 404.22 Pon.

ie nee ey es eee A

1 . 1 |

a ee

2 1

ated ee

Total

Epiterranean soil algae in Singapore 299

TABLE 10 — continued

(Cholorphyceae, Xanthophyceae and Euglenophyceae)

i zing ida a fs a a Ned a D | Total

CEDERCUREUTZIELLA SAVONEN-

sis Vischer .. of a 1 ie big de: ae i 1

DACTYLOCOCCUS INFUSIONUM

Nag. my i ‘ay 4 1 s mT oes ae = 1

MACROCHLORIS DISSECTA

Korch. 7 fy AES > a Fe a 1 ie) i. 1

PENIUM sp. ? 7 ” << sts od “<a - 1 eat 1

SCENEDESMUS OBLIQUs (Turp.)

Kiitz. e ? 1 1

ASTEROSIPHON TERRESTRE

Dangeard iy ge tae 1 am aa a v eee SE 1

PLEURASTRUM INSIGNE Chad. ey 1 ae sala - iu neg i: 1

SCENEDESMUS FALCATUS Chad. ee ] a = a - ‘% a8 1

CLOSTERIOPSIS sp. De 7 1 we ay x os ne 1

(d) Diatoms, Diatoms are surprisingly rare in Singapore soils. Owing to the

absence of a definite taxonomic work on terrestrial diatoms of this area, they

were difficult to identify. Nevertheless their rarity is evident from Table 11.

TABLE 11

Number of Occurrences of diatoms

$a | F A E G S Cc Est. D Total

PINNULARIA ‘Sp. 0} a 3 be 2 7

NAVICULA sp. .. aie ae Z 1 — 1 1 2 7

NAVICULA PYGMEA Kutz. .. =e Es Sel eee 1

NAVICULA CONFERVACEA Kiitz. tala 1 edi Aveltc ener. menaeee pod ofa: 1

NAVICULA HUDSTEDTH Krasske Rea} 2 | Se ee bate. Ap 2

GOMPHONEMA Sp. BL hd 1 ae j awoipyaens | 1

FRUSTULIA sp. .. ~ ee 1 ee ee 1

Total no. of species >. 1 | 3 | 3 | 3 | 0 | 1 | 2 | 1 7

300 Gardens’ Bulletin— X XVI (1973)

(e) Diversity of algal flora. Table 12 shows the occurrence of different kinds

of algae in the different habitats. Tree communities tend to lack diversity in algal

flora. In forest only 4 species are present, none of which is blue-green. Scrubland

and disturbed land have 15 and 16 species respectively. Algal flora is best deve-

loped in exposed sub-soil (29 species) and grassland (35 species). As mentioned

above nitrogen fixing blue-green algae may occur in all kinds of habitat except

forest.

TABLE 12

Summary Table for number of species of different groups of algae

= 2 Fy, | A’: |) Be ae Ss |) te | ee Total

a a bogeeay

Non-heterocystous blue-greens| 0 ‘4 15 10 9 7 2 | ot | 26

Heterocystous blue-greens .. 0 3 2 7 1 3 4 3 | 16

Green algae. te ap [i el Sa iy | | eee

Diatoms { SPs i se Ss i

Total no. of species = 5 23 2B) 35 15 18 | 16 | 10 | 80

Discussion

Although 80 species of algae have been found in the soil of Singapore Island

in this investigation, most of these show low constancy of occurrence in the Island

as a whole or in an individual type of habitat. The phenomenon of a large diversity

of species distributed with low frequency is familiar amongst tropical organisms

and occurs amongst such diverse groups as lowland tropical forest angiosperms,

beetles (Wallace, 1969) and tropical fish.

When the natural vegetation is completely cleared and the top soil removed

by man or erosion, an exposed sub-soil is devoid of any type of higher plant

vegetation. This is subject to excessive leaching by torrential rain, strong isolation

from the tropical sun and considerable temperature fluctuation day and night (Hill,

1966). Despite these severe conditions 29 species of soil algae were found some

of which may act as first colonisers of the bare ground before the establishment of

a higher plant flora. Non-heterocystous blue-green algae predominate in this

habitat.

When colonisation by higher plants commences, grassland becomes established.

The soil surface becomes protected from excessive leaching and there is an increase

in available ions for plant growth (Table 2.) Light intensity is still high but there

is less fluctuation in temperature day and night. 35 species of soil algae were found

including 7 species of heterocystous blue-green and 15 species of green algae.

Grassland progressively develops into scrubland (belukar) with colonisation by

woody species such as Adinandra dumosa Jack and Melastoma malabathricum

Linn. These species increase the shading of the ground layer. The light intensity

Epiterranean soil algae in Singapore 301

decreases and there is an increase in available nitrate. The algal flora loses some

of its diversity and only | species of heterocystous blue-green algae was found.

In the undisturbed forest the organic matter content is high but most cations

and anions remain low, since the ionic reserve in tropical forest is directly absorbed

by the shallow feeding roots of the trees and does not persist in the soil

(Jackson, 1968). The light intensity is very low being only about 0.1% of the

illumination outside the forest (Johnson, 1970). No blue-green algae are found

here and there are only four species of green algae and one diatom.

Cultivated soil in Singapore has been subjected to various manures and

organic fertilisers and usually has a relatively high nitrate level. The algal flora is

limited but heterocystous blue-green algae can occur. Estates are also subject to

disturbance by man. Since these are predominately tree crops (e.g. rubber, banana,

coconut, rambutans, durian, citrus etc.), the light intensity is usually lower than

in cultivated soil; but, since such estates are quite open, about 10% light penetrates.

The algal flora is limited but less so than in undisturbed forest.

Disturbed soil has been subject to all kinds of dumping and building operations

by man, including industrial pollution and general devastation. The occurrence of

such areas near a city of over two million population is to be expected. Chemical

analysis of the soil gives very erratic results often with high figures for one or

more ions. Ten species of algae were found in this kind of habitat.

Summary

The surface soil was collected in 100 localities in the rural area of Singapore.

The samples were cultured by liquid and plate culture to determine the algae

occurring, and the soil was subjected to physical and chemical analysis.

The surface soil consisted predominately of clay and clay-loam, with some

sandy loam and loamy sand, other types of soil being rare. The water-holding

capacity of dry soil is closely correlated with organic content being high in forest

and alluvial soils but low in all other types of soil. Nitrate level was normally at

moderate to moderately low levels. but the values for phosphate, sulphate and

potassium are critically low in most soils. Most soils are acid, forest soils having

an average pH of 3.83 with pH somewhat higher in other types of soil. Caroonate

was detected in only 17% of the samples. Although the soils were somewhat

uniform in hue with 60% soils in the 1OYR range, they were very variable in

colour.

80 species of soil algae were found in the soil. Tree communities lack

diversity in algal flora, which was best developed in exposed subsoil and grassland.

Most species show low constancy of occurrence in the Island as a whole or in

individual habitats.

302 Gardens’ Bulletin — XXVI (1973)

Literature cited

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Beijerinckia If Mineral nutrition and resistance to high levels of certain

elements in relation to soil type. Plant and Soil 14: 297-322.

Bristol, B.M. (1919) On a Malayan form of Chlorococcum humicola (Naeg.)

Rabenh. Jour. Linn. Soc., Bot., 44: 473-482.

Edwards Laboratory, Norwalk, Ohio: Soil Testing, in Technical Bulletin No. 162,

Michigan Agric. Exp. Station.

Fritsch, F.E. (1907) A general consideration of the sub-serial and freshwater algae

of Ceylon. Proc. Roy. Soc., Lond. B 79: 197-254.

Ghose, S.L. (1923) A systematic and ecological account of a collection of blue-

green algae from Lahore and Simla. Jour. Linn. Soc., Bot., Lond. 46: 333-346.

Gonzales, Ella A., and Valavigi, V.S. (1960) Algae in the rhizosphere of some

crop plants. Proc. Symp. Algol., Indian Council Agric. Research 333-342.

Hill, R.D. (1966) Microclimate observation at Bukit Timah Forest, Singapore.

Malayan Forrester 29: 78-89.

Jackson, H.L. (1958) Soil Chemical Analysis. Constable and Co., Ltd., London.

Jackson, J.C. (1968) Planters and Speculators. University of Malaya Press. Kuala

Lumpur.

Johnson, Anne (1962) Precursory studies on the epiterranean soil algae of Singapore

and Malaya. Gdns. Bull., Singapore, 19: 379-384.

——. (1966) B-values and the phytoplankton community. Verh. Internat.

Verein. Limmol. 16: 619-626.

. (1969) Blue-green algae of Malaysian rice-fields. Jour. Sing. Nat.

Acad. Sci. 1: 30-36.

—. (1970) Determinative factors in the distribution of lowland vegeta-

tion in Singapore and Malaysia. Nanyang Univ. J. 4: 165-174.

Leamy, M.L. and Panton, W.P. (1966) Soil Survey Manual for Malayan Conditions.

Division of Agric. Bull No. 19, Ministry of Agric. and Co-operatives, Kuala

Lumpur, Malaysia. |

Munsell Soil Colour Charts (1971) Munsell Colour Company Inc., Baltimore,

U.S.A.

Panton, W.P. (1955) Soil survey reports No. 2, A part of west Singapore Island.

Malayan Agric. J. 38: 1-26.

Prasad, S. (1949) Nitrogen recuperation by blue green algae in soils of Bihar and

their growth on different types of soil. J. Proc. Inst. Chemists (India) 21:

135-140.

Singh, R.N. (1940-41) The relation of algae to the nitrogen economy of the soil.

Chron. Bot. 6: 202-203.

Sulaiman, M. (1944) Effect of Algal growth of activity of Azotobacter in rice soils.

Ind. J. Agric. Sc. 14: 277-283.

Tiffany, L.H. (1951) in G.M. Smith, Manual of Phycology Ch. 15: 303-306.

Wallace, A.R. (1869) The Malay Archipelago, Macmillan and Co., London.

Pollination in Dioecious Figs

Pollination of Ficus fistulosa by Ceratosolen hewitti

J. GALIL

Department of Botany, Tel Aviv University, Israel

ABSTRACT

Pollination in the dioecious fig Ficus fistulosa was studied in Singapore. The pollinator,

Ceratosolen hewitti (Agaonidae), possesses closed thoracic pockets and coxal combs. Pocket

filling with pollen from detached anthers previously cut by the male wasps occurs in the

almost ripe male figs. Pollen is lifted from the anther to the underside of the thorax by

means of the arolia of the fore legs and then shoveled backwards into the pockets by the

combs. In the male figs, pollination of the stigmata occurs at the end of oviposition prior

to the withdrawal of the ovipositor. Pollen is taken directly from the pockets by the fore-

legs and is carried downwards onto the stigmata. The behaviour of the female wasps in the

female syconia is very similar except that the oviposition act does not culminate with actual

egg-laying. The significance of pollination in short-styled, gall-producing flowers and of

Ovipositor insertion into the styles of long-styled, seed-producing flowers is discussed.

Introduction

It is known that the symbiotic wasps (Agaonidae, Chalcidoldea) which serve

as pollen vectors in the genus Ficus carry out in the figs two types of activities

namely, pollination of the long-styled flowers and oviposition into the pistils of

the short-styled ones. Hence, the former produce seeds whereas the ovaries of the

latter develop into galls and give rise to the next generation of wasps. Both

activities are indispensable for the existence of fig symbiosis. There is no evidence

of the presence of apomixis in Ficus so that pollination, too, appears to be

essential for seed production. The question arises as to the occurrence of pollination

in the short-styled flowers and oviposition into the long-styled ones.

In the common fig (F. carica L.)., the most extensively studied Ficus sp.

(Grandi, 1929), the situation is comparatively simple. Here, pollen is passively

carried on the body of the female wasp Blastophaga psenes L. from one fig to

another so that pollination of both long- and short-styled flowers takes place

indiscriminately; on the other hand, the long styles, which so markedly differ in

structure from the short styles, do not appear to be suitable for oviposition.

The situation is not always as simple in other Ficus spp. There is considerable

variation in structure and behaviour of the pollinators of different species of Ficus,

in spite of an overall resemblance. Unique organs used by various spp. of wasps

as containers for transporting pollen from male figs (phase D*) to young receptive

female figs (phase B*) were recently discovered (Ramirez, 1969; Galil and

* According to Galil and Eisikowitch (1968) the developmental phases of figs were named

as follows:

Phase A (Pre-female): Young syconium prior to the opening of the ostiole.

Phase B (Female): Ostiolar scales loosen, female flowers ripen, sycophilous wasps

penetrate into the syconium and oviposit into the ovaries.

Phase C (Interfloral): Wasp larvae and fig embryos develop within their respective

ovaries. Ovaries occupied by the larvae are transformed into galls.

Phase D (Male): Male flowers mature, wasps reach the imago stage, fertilized

female wasps leave syconia via channels bored by the males.

Phase E (Post-floral): Both syconia and the seeds inside them ripen.

303

304 Gardens’ Bulletin — X XVI (1973)

Eisikowitch 1969; Galil and Snitzer-Pasternak, 1970; Galil et al., 1974). In these

figs, the pollination process consists of “deliberate” loading of the containers and,

upon entering the young figs, extracting the pollen and pollinating the female

flowers. Actual pollination takes place at the end of the oviposition act, before the

wasp withdraws her ovipositor from the pistil. Hitherto, pollination by wasps with

specialized pollen containers was exclusively studied in monoecious figs that

contain both long- and short-styled female flowers (the so-called seed- and gall-

flowers, respectively) in the same syconium. Because of the dense packing of the

fiowers within such figs, it is very difficult to observe the behaviour of the wasps

in regard to the different flower types. Nevertheless, there is some information

available. Johri and Konar (1956) discovered fig embryos up to the octant stage

in the short-styled ovaries of F. religiosa L. This is clear evidence of fertile

pollination of the short styled flowers. “Seed figs” in which both long- and short-

styled flowers bear normal seeds that are occasionally found in various species

(Galil and Eisikowitch, 1971), provide further evidence of normal pollination in

both types of flowers. On the other hand, the blackening of the upper part of the

style in long-styled flowers of F. religiosa, observed by the same authors, indicates

that the wasps insert their ovipositors into such pistils as well.

Direct observations of the behaviour of the wasps in long- and short-styled

flowers in regard to pollination and oviposition were hitherto carried out only in

the monoecious figs of F. sycomorus L. (Galil and Eisikowitch, 19769). However.

since the two flower types in that species are very similar and cannot be

distinguished from the above without destroying the fig, only a few sporadic

observations could be made.

Dioecious fig species, in which long- and short-styled female flowers develop

in different syconia, lend themselves more easily to detailed observations on wasp

behaviour with regard to the two flower types. For such observation Ficus spp.

whose pollinators are equipped with specialized pollen containers which are

actively loaded and emptied must be chosen.

The dioecious figs of F. fistulosa Reinw. ex Bl. (Fig. 5) which belong to the

Sycocarpus section of subgenus Ficus and are pollinated by Ceratosolen hewitti

Waterston, possessing thoracic pockets for pollen transport, prove suitable for the

present study.

Materials and Methods

The study was carried out in June-July, 1971 in Singapore. The chief object,

F. fistulosa, is a small tree, 3 to 4 m. high, that grows abundantly along road

edges in the suburbs of the town and on the borders of the jungle.

Generally, the study of pollination in dioecious figs is more complicated than

in monoecious species since it is necessary to find wasp-releasing D-phase male

figs and receptive B-phase figs of both sexes simultaneously in the same vicinity.

The month of June proved to be very suitable since it is a period of overlapping

of the two main crops of figs in F. fistulosa. Hence, all of the required develop-

mental phases of the figs were available (for Hong Kong, see also Hill, 1967).

During the study, no ripening female figs were found, but these were not indis-

pensable for carrying out the necessary observations.

The cauliflorous figs of F. fistulosa develop in great numbers, provide abundant

and easily accessible material. The timing of observations was adjusted to the

activities of the wasps in the male and female phases of the fig. Hence, the

Pollination of Ficus fistulosa 305:

observations on pocket filling were carried out in the early morning, between 3

and 5 a.m.

For observing the behaviour of the wasps in young figs during oviposition

and pollination, B-phase figs, male and female separately, were enclosed in plastic

vials covered with insect-proof netting together with wasp-releasing D-phase male

figs. The emerging wasps which penetrated the receptive B-phase figs provided good

material for observation of wasp behaviour during oviposition and pollination.

Observations

Syconia and pollinators

The figs develop on small lateral shoots on the trunks and branches (Fig. 6).

The receptive (phase B) figs of both sexes are pear-shaped with a short neck and

are borne on distinct peduncles that are up to 25 mm long (Fig. 7). The male figs.

are somewhat larger than female figs, 14 x 16 mm as compared to 11 x 13 mm.

The ostiole is depressed and is sealed by imbricate scales. The structural features

of the fig are especially prominent when bisected figs are viewed from the inner

side (Figs. 8, 9). The different components have different colours: the ostiolar

scales are orange and the pale stigmata stand out on the background of crimson

ovaries. The internal cavity of the fig is wide and distinct.

The male figs contain a large number of short-styled female flowers (“gall

flowers’) covering almost the entire inner surface of the fig and 30 to 35 ostiolar

male flowers arranged in one to two rows (Fig. 8). The male flowers are relatively

prominent and easily discernable, but are still closed and unripe. The “gall

flowers” are almost hidden within the fused perianth lobes (Fig. 1, IJ). The short

(about 0.3 mm) and relatively thick style and the funnel-shaped stigma at its end

project out of the perianth. The broad and dense stigmata form a continuous

surface around the fig cavity but do not coalesce into a single “synstigma”.

The long-styled female flowers (‘‘seed flowers”) which cover the entire inner

surface of the female figs are quite different from the short-styled ones. The

perianth is much reduced thus forming a narrow ring at the base of the ovary

(Fig. 1, IIL). Consequently, the pistils are naked and visible for their entire length

in bisected figs. The ovary is smaller and the style is narrower and much longer

(about 1 mm) than their counterparts in the short-styled flowers. The stigma is

cylindrical, thick and long (0.16 x 0.3 mm).

At the wasp-releasing stage (phase D), the male figs are almost globose (Fig.

10), somewhat broader than high (20 x 25 mm). The holes in the empty gall-ovaries

from which the wasps have already emerged have rounded lids which are shifted

to the side (Fig. 12).

Upon ripening of the male flowers, the filament of the single stamen elongates

so that the anther is pushed through the top of the fused perianth (Fig. 1, I). At

this stage, the males of the pollinating wasps approach the male flowers and cut

off some of the anthers causing them to be scattered within the cavity of the fig.

Thus, the emerging female wasps encounter cut anthers in addition to the intact

stamens remaining in situ around the ostiole (Fig. 11). At the end of phase D,

a large hole is cut through the scales in the centre of the ostiole by the male

wasps (Fig. 12). The pollen loaded females emerge from the fig through this hole.

306 Gardens’ Bulletin — XXVI (1973)

After the liberation of the wasps, the fig grows rapidly, and at ripening

(Phase E) it reaches about 30 mm in diameter. It becomes yellowish and attains

a characteristic smell which is typical for bat-dispersed fruit.

As noted above, no ripe female figs of F. fistulosa were found throughout the

period of the present study. Hill (1967) states that the ripe female figs are somewhat

smaller than the male figs, but are more succulent and their yellow colour is more

pronounced.

The pollinator, Ceratosolen hewitti Waterston (Wiebes, 1963, 1966; Grandi,

1963) is a small agaonid wasp about 1.5 mm long (Fig. 2). Only those

characteristics of the female wasp that pertain to pollination will be described

below.

Of the two types of specific pollen containers hitherto described in pollinators

of Ficus, namely “coxal corbiculae” and “thoracic pockets” (Ramirez, 1969; Galil

et al., 1974), the female of C. hewitti possesses the latter only. In whole mounts

of the ventrum of the body, the pockets appear on both sides of the mesothorax

as triangular depressions that are entirely covered by continuous flaps (Fig. 3).

Generally, the structure of the pockets is quite similar to that of the corresponding

organs described in C. arabicus Mayr, the pollinator of F. sycomorus L. in East

Africa (Galil and Eisikowitch, 1969). It appears that the pockets open and close

along their inner suture facing the median axis of the body through changing in

the position of the covering flaps.

Coxal corbiculae such as described by Ramirez (1969) and Galil et al. (1974)

were not found on the fore coxae of C. hewitti. There are no depressions along

the coxae: only the comb, namely the row of stiff bristles which usually fringe

the depression on its inner side, is found here. In C. hewitti, the comb is a distinct

organ formed from a straight row of about 18.38y-long bristles (Fig. 4).

The length of the ovipositor of the pollinating wasp is of decisive importance

in fig symbiosis. As seen from the side, the ovipositor of C. hewitti scarcely

projects beyond the apex of the body. In fact, the ovipositor is not short but

arises at some distance forwards, on the under side of the abdomen. Its real

length may be seen during egg-laying when its shaft is released from the protecting

valves and is directed towards the stigmata.

Pocket loading

Detailed observations of the filling of pockets of C. hewitti in D-phase male

figs of F. fistulosa are not easy. Figs with ostiolar male flowers are usually suitable

for such observations. However, in F. fistulosa the situation becomes more

complicated because of the early emergence of the female wasps from their gals

and the filling of the pockets prior to the tunnelling of the syconial wall by the

male-wasps. For successful observation of pocket loading, the figs must be cut

and studied during a comparatively short lapse of time when the anthers are still

attached but sufficiently ripe, and the fertilized females have not yet left their galls.

When the figs are cut at the appropriate time, the female wasps become active,

emerge from the galls, approach the male flowers and start filling their pockets

from the anthers which protrude out of the floral perianth.

Lifting of the pollen from the anthers to the side of the thorax is accomplished

by rapid alternating movements of the forelegs. Similar movements were observed

in the pollinators of monoecious figs previously studied. Clumps of pollen are

Pollination of Ficus fistulosa 307

Fig. 1. Ficus fistulosa, flowers: I— male flower,

Il — short-styled female flower (°);

III — long-styled female flower °.

Fig. 2 Ceratosolen hewitti (° and ¢).

Fig. 3. Mesothorax of Ceratosolen hewitti (from

below) showing pollen pockets.

Fig. 4. Fore leg of Ceratosolen hewitti, with

comb.

Pocket

308 Gardens’ Bulletin— XXVI (1973)

placed between the body wall and the fore coxae which move up and down when

the pollen is lifted by the bending legs. Shovelling movements of the fore coxae

and combs by which the pollen is brushed backwards from the underside of the

thorax to the thoracic pockets occur after each sequence of pollen liftings. The

coxal comb is clearly a unique organ and even in the absence of the coxal

depression (‘‘corbicula’’), it has a particular role in pocket filling. The shovelling

movements are rapid but can be clearly seen. The role of the combs in the

brushing of the pollen is evident from the pollen grains which are found on the

bristles. When the quantity of pollen in the anther does not suffice for the filling

of the pockets, the wasp moves to a second anther and begins a new series of

liftings and shovelling movements.

Pocket emptying: pollination in male figs

The observations on the behaviour of the female wasps during oviposition

and pollination were especially successful in the early morning when B-phase

receptive figs were enclosed in plastic vials together with wasp-releasing D-phase

male figs. As in the monoecious Ficus spp. previously studied (Galil and Eisiko-

witch, 1969; Galil et al, 1974), the impregnated and pollen-carrying female

‘wasp starts to oviposit and to pollinate the stigmata immediately after its entrance

into the receptive fig. In C. hewitti, as usual in the agaonid wasps, the ovipositor

shaft is released from the protecting valves at the beginning of oviposition act and

is supported and guided to the stigmata by the sternites that fold lengthwise to

form a triangular, gutter-shaped, body. The total length of the ovipositor is about

0.8 mm., i.e. its length is intermediate between that of the styles in long- and

short-styled flowers. After several probing movements for an appropriate oviposition

site, the ovipositor sinks into one of the funnel-shaped stigmata of the short-styled

flowers. It remains in the style only for 15 to 18 seconds which is a very short

time compared to the duration of the act in pollinators of other figs previously

studied, i.e. about 60 seconds in C. arabicus, and 3 to 3.5 minutes in Blastophaga

estherae Grnd., the pollinator of F. costaricana (Liebm.) Miq. (Galil et al., 1974)

Almost immediately after the insertion of the ovipositor into the pistil of the

fiower, the female wasp begins to perform typical pollination movements. The

forelegs fold upwards upon themselves one after the other intermittantly, thereby

bringing the arolia into contact with the pollen pockets on the under side of the

mesothorax.

The pollen grains are taken up by the legs directly from the pockets; in any

case, no shovelling of pollen from the pockets by means of the combs of the fore

coxae were observed. The pollen-carrying legs open downwards one ofter the

other and touch the stigmata beneath. It is apparent that normal pollination of

the short-styled flowers takes place systematically at the end of each oviposition

act. During oviposition and pollination, the head of the female wasp remains

motionless and no biting of the stigmata takes place, as observed in C. arabicus

(Galil and Eisikowitch, 1969).

Pocket emptying: pollination in female figs

In spite of the distinct differences in the structure of the pistils in the female

flowers of the male and female figs, the behaviour of the wasps was almost alike.

As in the male figs, the female wasps moved on the surface of the stigmata,

inserted the ovipositors into them one by one and performed the characteristic

pollination movements at the end of each insertion. The retention of the ovipositor

Plate 1.

Figs. 5-12 Ficus fistulosa

Fig. 5. Leafy branches. Fig. 6. Fruiting stem. Fig. 7. Short branch with male

figs at Phase B. x 0.7. Fig. 8. Cross section through male fig at phase B. (°)

short-styled female flowers. ¢ —vunripe male flowers . x 3.7. Fig. 9. Cross section

through female fig at phase B. ° — long-styled female flowers. Fig. 10. Male figs

at phase D. x 0.6. Fig. 11. Longitudinal section through male fig after release of wasps.

$ — attached anthers; A — detached anthers; x 2.3. Fig. 12. Cross section through male

a ey RE De gat) A ee) Ee eee ae > ee ar ey 2, a. me. |

Pollination of Ficus fistulosa 309

within the style was shorter than in the short-styled flowers and lasted barely 8

to 10 seconds. Here also pollen is transferred directly from the thoracic pockets

onto the stigmata of the long-styled female flowers by the arolia of the fore legs.

During these observations, several cases of unusual behaviour of the wasps

in the female figs were encountered. Exhausted female wasps, which have already

completed their usual activities, continued to move their fore legs in vain efforts

to extract pollen from the pockets and to deposit it on the stigmata.

In order to observe this phenomenon more closely, pollen loaded female

wasps were taped in a supine position on a glass slide and observed through the

dissecting microscope immediately after they emerged from the D-phase male figs.

Most of the wasps made vain attempts to get off the glass; however, in a few

cases, the wasps started to perform the characteristic pollination movements during

which they folded the fore legs onto the thorax and inserted the arolia into the

widely gaping pollen pockets. Pollen grains could be clearly seen on the arolia. No

such movements have hitherto been seen in the pollinators of the monoecious figs

in which pollination has been studied.

Discussion

In spite of numerous structural differences between the syconia of dioecious

F. fistulosa and monoecious F. sycomorus (Galil and Eisikowitch, 1969) which

belong to different subgenera of Ficus, namely subgenus Ficus and subgenus

Sycomorus respectively, the two have several biological features in common. Of

particular significance is the relatively small number of male flowers which reflects

the efficiency of the pollen vectors in pollen transfer. In both Ficus taxa, the

pollinating wasps are species of Ceratosolen possessing closed thoracic pollen

pockets of similar structure and coxal combs without corbiculae, and behave very

similarly in relation to the figs. In both species, the male wasps cut the stamens

and the females emerge from their galls, fill their pockets with pollen from the

detached anthers and gather in the fig cavity before the syconial wall is tunnelled

by the males. Also the behaviour of the female wasps during pocket filling and

emptying, i.e. pollination, is similar. Such likenesses indicate that physiological

conditions within the figs are probably also similar in both cases (Galil et al.,

1973).

In F. fistulosa, the behaviour of the female wasps in the short- and long-styled

female flowers (in different figs!) is almost identical in spite of the structural

differences between them. In the two types of flowers, both insertion of the

ovipositor into the style and deposition of pollen on the stigmas take place.

Essentially, it is the same process although eggs are not likely to be laid within

the long-styled pistils. In F. sycomorus, serial microtome sections through long-

styled pistils in pollinated figs failed to reveal any remnants of Ceratosolen

ova. The pistils of the long-styled flowers of F. fistulosa have not been similarly

investigated but there are some indirect indications that here too, no real

oviposition takes place. The relatively much shorter time that the ovipositor remains

in the long styles is significant. It appears that for effective oviposition, the stimulus

which the flower ovary exerts on the tip of the ovipositor is indispensable.

Consequently, when the style is too long and the ovary is beyond the reach of the

ovipositor, there is no egg-laying.

310 Gardens’ Bulletin — X XVI (1973)

The behaviour of the wasp in the male and female figs raises two questions

which may be of fundamental importance for the understanding of fig biology: 1.

What is the biological purpose of pollination of the short-styled pistils which are

in any case destined to develop into wasp-galls? and 2. Is there any significance

to the futile insertions of the ovipositor into the long-styled pistils? In F. religiosa,

where the first problem was studied more thoroughly, the selective advantage of

pollination of the gall-producing short-styled flowers is evident. In unpollinated

short-styled flowers, the biological equilibrium is destroyed because of paucity

of food, resulting from the failure of normal endosperm development, causes high

mortality in female wasp larvae (Galil and Eisikowitch, 1971). It is likely that in

F. fistulosa the situation is similar.

As yet, there is no satisfactory answer to the significance of the ovipositor

insertion (‘‘stinging”’) into the pistils of the long-styled flowers. Perhaps the

changes caused by wounding of the tissue has some impact on pollen germination

or pollen tube growth. It should be borne in mind that attraction of the wasps to

the female figs and the activities of the wasps within the female figs are of no

apparent immediate advantage to the insect. Identical chemo-attractants, produced

by both types of figs, could cause the wasps to mistake the female figs for the

male figs, which provide the necessary brooding place for the new generation.

Thus, it is also possible that insertion of the ovipositor into the long-styled pistils

has nothing to do with the biological processes in the female figs but represents

a link in an automatic, instinctive chain of activities, orignally concerned with

short-styled flowers in male figs.

It should be noted that a strict coordination between the pollination movements

and oviposition in both male and female figs is not always sustained. The

numerous pollen-extraction movements observed in dying female wasps and the

repeated pollen withdrawals from the pockets observed in taped wasps hint at a

possible disruption of such coordination. Consequently, normai intermittent

pollination which is regularly interrupted by search for new sites of oviposition

and insertions of the ovipositor into the stigmata may be replaced by a continuous

act, consisting of mere emptying of the pockets.

Acknowledgements

The author acknowledges with many thanks Professor E.J.H. Corner, Botany

School, Cambridge, and Professor J.T. Wiebes, Rijksuniversiteit, Leiden, for their

helpful information on the fig spp. and the fig wasps of Singapore. The author

is also indebted to Professor A.N. Rao, Head, Professor H. Keng and Dr. P.N.

Avadhani of the Department of Botany, University of Singapore and to Mr. A.G.

Alphonso, Director, and Dr. Chang Kiaw Lan of the Botanic Garden of Singapore

for their assistance and hospitality. Thanks are also due to S. Bullowa and Dr. A.

Horovitz for reviewing the manuscript, to S. Schaeffer for the illustrations, and

to J. Cohen and A. Shub for the photographic work.

Literature Cited

Corner, E.J.H. 1958. An introduction to distribution of Ficus. Reinwardtia 4:

15-45.

. 1965. Check-list of Ficus in Asia and Australia with keys to identification.

Gard. Bull. Sing. 21: 1-186.

Pollination of Ficus fistulosa 311

Galil, J. & D. Eisikowitch, 1968. On the pollination ecology of Ficus sycomorus

in East Africa. Ecology 49: 259-269.

& . 1969. Further studies on the pollination ecology of Ficus

sycomorus. Tijdschr. Entomol. 112: 1-13.

—— & . 1971. Studies on mutualistic symbiosis between syconia and

sycophilous wasps in monoecious figs. New Phytol. 70: 773-787.

, M. Zeroni & D. Bar Shalom, 1973. Carbon dioxide and ethylene effects

on the coordination between the pollinator Blastophaga quadraticeps and the

syconium in Ficus religiosa. New Phytol. 72 (5).

, W.B. Ramirez, & D. Eisikowitch, 1974 Pollination of Ficus costaricana

and F. hemsleyana by Blastophaga estherae and Blastophaga_ tonduzi

(Agaonidae) in Costa Rica. Tijdschr. Ent. In Press.

, & Y. Snitzer-Pasternak. 1970. Pollination in Ficus religiosa L. as

connected with the structure and mode of action of the pollen pockets of

Blastophaga quadraticeps Mayr. New Phytol. 69: 775-784.

Grandi, G. 1929. Studio morfologico e biologico dell Blastophaga psenes L. Boll.

Lab. Ent. Bologna. 2: 1-147.

. 1963. Catalogo regionato degli agaonidi de mondo. Boll. Ins. Ent. Univ.

Bologna 26: 319-373.

Hill, D.S. 1967. Figs (Ficus spp.) of Hong Kong. Hong Kong Univ. Press, Hong

Kong.

Johri, B.M. & R.N. Konar, 1956. The floral morphology and embryology of

Ficus religiosa Linn. Phytomorph. 6: 97-111.

Ramirez, W.B. 1969. Fig wasps. Mechanisms of pollen transfer. Science 163:

580-81.

— . 1970. Taxonomical and biological studies of neotropical fig — wasps

(Hymenoptera, Agaonidae) Univ. Kansas Sci. Bull. 49: 1-44.

Wiebes, J.T. 1963. Taxonomy and host preferences of Indo-Australian fig — wasps

of the genus Ceratosolen (Agaonidae). Tijdschr. Ent. 106: 1-112.

——— . 1966. Provisional host catalogue of fig wasps (Hymenoptera, Chalcido

idea). Zool. Verh. 83: 1-44.

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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.

Vol. 1 (1-5) January-May 1912 issued under title of Agricultural Bulletin

of the Straits & F.MLS.

Prices on application.

4. The Gardens’ Bulletin, Singapore (Series TV).

Vols. i2-25, 1949-1971. Vol. 26 Pt. II in print.

Price: Vol. 13 Pt. 1 (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.

Vol. 26 $36 per vol. (Pt. I $18; Pt. II $18).

Individual parts vary in price.

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.

Many from 1909 onward remain available.

Prices vary.

—~]

. (a) Malayan Orchid Hybrids by M. R. Henderson and G. H. Addison. $15.

(b) Malayan Orchid Hybrids, Supplement I by G. H. Addison. $21.

8. A Revised Flora 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 Director, Botanic Gardens, Cluny Road, Singapore

10.

Items 7-9 obtainable from Singapore National Printers (Pte) Ltd, Upper Serangoon

Road, Singapore 13.

Prices quoted are in Singapore Dollars

Overseas postage is extra