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THE

GARDENS’ BULLETIN SINGAPORE

Volume XXVI 1972-1973

A periodical reflecting the interests and activities of the Botanic Gardens

Singapore

EDITOR: Chang Kiaw Lan

To be purchased at the Botanic Gardens Ciuny Road, Singapore 10

Published by Botanic Gardens Parks & Recreation Department Singapore

Printed by the Singapore National Printers (Pte) Ltd

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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 62 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

82

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

KP

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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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 - - . -

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THE LIBRARY GARDENS’ BULLETIN SINGAPORE Vol. XXVI, Part I | 30th December, 1972 CONTENTS

PAGE l— 48

49- 61 62 63- 83 85— 93 95-110 11 fiz

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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.

10

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

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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.

49

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).

56

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)

58

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 :

:

2

4

y)

6 4

8

;

in 4410 cm —

x3

x2

Fig. 3. Pentace grandiflora Kochummen (fruiting twig FRI 12013 flower FRI 9683, type). .

Notes on the Systematy of Malayan Phanerogams

6 8 in 4+10

Fig. 4. Schoutenia furfuracea Kochummen (FRI 8042, type); inset: fruit with ersistent bracts.

61

cm

Studies in Macaranga V: M. lowii

by

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.

62

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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

63

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

: : ] | : : } i 4

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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