Pont hy m2 ~~ = Ene © tyne ww ‘ fv On Otel 4. Quem Wipe eee ~ 2 . . Tine ete Sant ae om - Mate are, Phelan, aan at PA TT Library Arnold Arboretum Harvard University Digitized by the Internet Archive in 2014 https://archive.org/details/gardensbulletins426unse 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 IVEX omaip'y eVeL-sVel a cxoroial =. gnitoslivy Ievibd © arabe sinsiof sdp lo. sibivi ¥ * ej : | F3 J fo a i , : Ai > t * q - of : s * ° a a 2 a * + ) ; ° 7 » anobiat) oir nae Hf lt 38 anil, . OP ercqpenr ni eect * ¢ *. he ’ + a oe 4 : = bh “a o % | i ansbvei aies 4 budailduy © : : en einai od aie Te hie aia i Vn ‘y . aes re a L ah 409 % 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 Bh-| a 1d- 2b cA ‘t8-t0 £0-22 OG} 1-28 ii Sif DEI-EN oti e£I-TS/ EQEREI B2t-Z2! BOLO? | Gis--2e1 Hs 4 Lot £—~REC VES-THE 1at-eR2ts #ah~E at T8S-@as SAE -OBo bik -< 08 " RSE~F te in ‘egal eit ae novnsbrs’ - to siden of} ao patieaty mp 4 | ah ihe: ay ae Se ne enere” lo Yiamisley? ot no etovl “atta att ae » ° ; : re A z 7 a. Siarol NM ¥ agamanol Es wd al st00M A pinprujistasnad ue paren" outs dent rik sOaNOE, rebi m i itil (het ‘abut, bia ont (noldes: 2 FV) innit isi ror et FF Good Hy denih 3 Xe 2aT9190 0 “or od 40 iar a ‘ seuincobars : ns “a er oval A ni I sddold sn oft to. plaice ge athe Rgesaindhas 0 of Horoaong 8 bas sieeatatt raont analy 2 gree a | Stine ie eynoaye « Ot es ; o ped 7 te, . estas to por eit" scigailid’ : ae iwa2 sl: of ant bas amis vioge — rusibiesd a at es bs wie ‘ ae wee rs atieks ak JM ery octane i> , , S20 coiingie hing sain Yorn sigolck oft reasons A A be ian 5 he ii itamue Sai nidamgise prow mot & a arorpnyi2 Yo atrushey hove lo teil .. | sale wi AGE ati aielndno® st omnes. ai ar irenntonaed brye Vitida é naifod sar sal t | a ba we ft Pes 7 >}, fea santa casein) inate allsidgneyd ewe) “ae woivest A bas OTC! wt Sawwme wa abo H, eunall ; , par srerealache te act ob ao 29101 simone Wt 7 ey ; Lees -e2ni2 at neil fiow nonunvatin bahia: viiat 3 wd DAS wa ani hos ae 2 ee ny. Ry wre ‘ Ay ot ee 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 OS ALS 6 Gt in ra No eclanw: ohient | jos a5! aL Ee ‘edi wien a2 ot a | ; ; mi’ wifihianien Sa here) nh vila % ee tel ‘ eferiexyin 4] Hidnts yaa? 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Cee oars Tt o2c T2E AhZo bes Tae ailiagV 20 giles Nee OTS giztobo e* [a ae ~ Th nsitiaaimg ati * TA gioozc1 ce ’ 7 00S ,ginofusea pllongvioV fA sithellaW Ob Pupsbionizeaqus sllsigiuehaW Iv iocnuld cinoenmnioW d08 -onull eageminen, fe / Lec PAID lg OS EOL murrisqeo mek £co¢ s¢ ; rand we: ees iy! VA Off gacieuae ace #2 «SS 20 Valet Ob miedo i5v : Of] stasdony BFC plot SEC) gucbrok BES gilolizgcim ” $€8 iixenoilig iay Bibom a £% pinot iy a eau FES eiyirsl 4 Mig ivciados J0e Oe val aoX m2. Rae dodigatSs cman ‘£89 nilgomea eusiqnysis | Ps SOL senongys 2: SOF ieee, hewn Sm / ' det VEY a ee . A Wie ites it S oe a ae — es COED PEOPLE OPE FE GFE a MAR 7 1973 P\ et i K. U.: The Lindsaeoid Ferns of the Old World VI te Continental Asia, - Japan and Taiwan . ! Forest Research Institute, Kepong, Loh Notes on the i of ss ee oa 1 Phanerogams XLXVII - - . - y DRANSFIELD J.: The Genus J ohannesteijsmannia H. E. Moore Jr. S x Hn R. D.: Soil Moisture under Forest, Bukit Timah Nature Reserve, Singapore (J ion RATNASABAPATHY M.: Algae from Gunong Jerai (Kedah Peak), Malaysia - p, A) b ee: Illustrated Guide to Tropical Plants (P.S. Ashton) - : : _ REVIEW: Pollen Flora of Taiwan (H. Keng) -, s ° : AY hs ~ STONE B. C.: Arthrodactylis and Pandanus: A brief comment on the ‘Characteres 4 x fis Generum Plantarum’~ - - = i : é z : Lim ‘SIEW-NGo: Cytogenetics and Taxonomy of the pelted Globba L. in Malaya _ — IV Distribution in relation to polyploidy’ - - OX : scan KENG: Two new Theaceous plants from Malaysia a Proposal to reduce A _ Tutcheria to a synonym of Pyrenaria - . a 5 - TIxter P.: Mount Maquiling Bryoflora’- - - - - - ‘ a SHIGEO KuraTA and MaSsaMi ToYOSHIMA: Philippine Species of Nepenthes : sscxmilamn E. J. H.: Studies in the basidium — spore spacing and the Boletus spore To be purchased at the Botanic Gardens, Singapore Price: S $18 Published oY Authority Printed ak Tan Cheong Hui, Acting Government Printer, Singapore ra . 1972 ‘ all PA med Fay 73 OS ’ 7 t £ 4 ¥ ® wed ‘ WHITMORE T. C.: Studies in Macaranga V : M. lowii_ - - - - 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 113-114 115-126 127-135 137-153 155-158 159-194 YR DLE EYER YO OIE OYE YEEIOIEEIEIIEOWPIOIPEO OE a> Kn? Wow VK ROS oe ERE ea < BOOK ANNOUNCEMENT Boletus in Malaysia. E. J. H. Corner — Boletus in Malaysia published by authority under the auspices of the Botanic Gardens, Singapore, is available for sale now. Pp. vili --- 263, 80 text-figs., 19 colour pls, 4 pls. Price $50.00 Singapore currency, hard cover. Orders should be placed with: The Government Printer, P.O. Box 485, Singapore. 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 , L | y Wty Y Sp LGD: Wy thoy YU Wyyg ™ COE f, # Yj YY Plate 4. JOHANNESTEIJSMANNIA ALTIFRONS Northern hill-top form. Bukit Koh, Kuala Kenyam Taman Negara, Pahang. WEE The Genus Johannesteijsmannia H. E. Moore Jr. 71 and Upper Montane forest (sensu Richards 1952, — non Robbins 1968), growing with Livistona tahanensis, Dipteris conjugata, Weinmannia blumei, Dacrydium elatum and Podocarpus neriifolius. Another high record for Jt. altifrons is on Gunong Mandi Angin on the borders of Kelantan, Trengganu and Pahang where it grows at 1.200 m (Whitmore, pers. comm.). In most of its localities, Jt. altifrons is found in Hill Dipterocarp forest; in the low-lying Johore localities it is found in Lowland Dipterocarp forest, and in Sarawak in Kerangas forest (heath forest) on deeply podsolized soils. In all localities visited, soils are well drained. However Miquel (1861) records Jt. altifrons in Sumatra as growing as a swamp-palm, in maritime swamps similar to the habitat of Nypa — this seems highly unlikely when the Malayan and Bornean habitats are considered. Palm and Jochems (1924) record Jt. altifrons as occurring abundantly on the higher ground, but being absent from swamps. Jt. altifrons is not confined to soils derived from any one rock type, being found on granite-, rhyolite-, shale-, and sandstone-derived soils. Its main requirements appears to be one of well- drained soils. Despite the abundance of apparently suitable habitats, Jt. altifrons is absent from large areas of Malaya. For example it is absent from more or less the whole of the Main Range of Malaya except for one record at Sungei Lalang, Ulu Semenyih, in Selangor. The two valley systems parallel to the Ulu Semenyih — Ulu Gombak and Ulu Langat — are well botanized and have large aborigine populations, and if Jt. altifrons grew there, it would have been recorded. Yet the three valleys are more or less identical in geology, vegetation, and aspect. The palm is such a conspicuous plant, and so well known to aborigines and Malaya that it is fairly safe to assume that absences represent real absences, and not lack of exploration. Similarly Jt. altifrons is absent from large areas of Taman Negara (the National Park), Pahang, being found only on Gunong Tahan, and on Bukit Koh near Kuala Kenyam. Bertam, Eugeissona tristis is similarly very local in the National Park, being known only from Bukit Jeram Panjang on the Tembeling River. TAXONOMIC NOTES Jt. altifrons varies considerably over its geographical range; depredations by aborigines and Malays for thatching, and forestry practices tend to obscure the pattern of variation in reducing the leaf size of the palm by leaf cutting and exposure to greater light intensities. Populations in the North of Malaya, in Pahang and Kelantan, and in Selangor on hill slopes and ridge-tops, usually have larger, more numerous leaves, and inflorescences with a larger number of branches than those of the populations of lowland Johore and Sarawak. It is sometimes possible to deduce the origin of a particular specimen if it is well collected. From the photograph in Palm and Jochems (1924) it appears that the Batang Serangan populations are most similar to those of lowland Johore and Sarawak. Plants at 1,000 m on Gunong Tahan approach in size those of lowland Johore but there is an observable gradation in size from large Hill Dipterocarp forest plants at 500 m to the small mountain forms at 1,000 m, as the mountain is ascended. Typical mature plants from Hill Dipterocarp forest at Sungei Lalang F. R., Ulu Semenyih, Selangor, had an average of 28 leaves per plant, leaves 5.5 m tall, lamina 3.5 m x 1.2 m: in contrast, typical plants from Sungei Kayu Aru, on the road between Kota Tinggi and Mersing, lowland Johore, had 20 leaves per plant, leaves 3.3 m tall, lamina 2.3 m x 1 m. Despite this variation in leaf size, floral structure appears to be constant. Owing to the scarcity of Gardens’ Bulletin, Singapore — XXVI (1972) 72 1Ocm JOHANNESTEIJSMANNIA PERAKENSIS Fig. 3. inflorescence, B: flower A: The Genus Johannesteijsmannia H. E. Moore Jr. 73 flowering material it has not been possible to pursue this problem further. It is possible that some of the variation in size is environmentally induced; on Gunong Tahan the soils at 1000 m where the palm occurs are extremely poor podsols derived from weathering quartzite, whereas lower down the soils are derived from sandstones and are not podsolized. Some of the variation, however, is probably genetically controlled; many of the populations are isolated and the chances of cross pollination with other populations are minute, so circumstances are ideal for divergence of populations. It is only possible to speculate on the significance of the variation: unwieldy plants such as Jt. altifrons present too many difficulties for experimental work. Johannesteijsmannia perakensis J. Dransfield, sp. nov. DIAGNOSIS Jt. altifrons multo affinis, sed differt caule ascendenti trunci instar, inflore- scentiae ramis in ordinibus quattuor divaricatis, floribus fragrantibus alabastro rotundatis, fructu maiore verrucis pluribus praedito. DESCRIPTION Palma solitaria, aetate caule ascendenti robusto ad 4 m alto et 15 cm diametro, annulato-cicatricoso. Folia erecta grandia ad 4 m longa. Petiolus ad 1 m longus, 2 cm latus. Lamina ad 3 m longa, ad 1.6 lata, squamis furfuraceis fuscis intra secus costam et in lobis brevibus marginalibus tecta. Plicae numerosae ad 20 utrimque, inter plicas nervulis lateralis anastomosantibus. Margo follii inferior spinis brevibus (ad 1 mm) armatus atque petiolus. Petiolus duobus striis luteis notatus. Vagina folii fibrosa marcescens reticulum fuscum formans. Inflorescentia axillaris corona foliorum arcuata. Spathae fugaceo-furfuraceae, primo cremeae dein fuscentes, 5 —6 numero, 20 cm longae et 8 cm latae, basi tubulosae superne inflatae, apicibus acutis, imo latere fissae. Pedunculus tomen- tosus ad 50 cm longus et 2 cm latus, ordinibus quattuor divaricatis ramosus. Rami floriferi 50— 100, ad 2.5 mm diametro, ad 10 cm longi, viriduli, alboto- mentosi, floridi. Flores albi glabri, alabastro rotundati, solitarii vel bini vel terni ageregati, in tuberculis prominentibus insidentes, unusquisque bracteola minuta subtentus. Calyx brevis ad 1 mm altus, carnosus, glaber, paullo 3 — lobatus. Petala carnosa alba, ad 3 mm longa, late triangularis basi in tubo breve connata. Annulus staminalis minute epipetalus, in filamenta 6 aequalia superne abrupte subulata. Antherae ovales. Grana pollinis alba. Ovarium glabrum carpellis 3 basi liberis, apice in stylo communi 0.8 mm longo connatis. Fructus ad 5 cms diametro, plerumque a carpello uno evolutus, raro a 2 —3 carpellis, c. 60 verrucis suberosis fuscis sordidis ad 9 mm altis, tectus. Endocarpium lignosum ad 1 mm crassum. Endospermium osseum ad 2.5 cm diametro, basi integumentorum contextu suberoso penetratus. Embryo lateralis. Malay Peninsula: Perak, in monte Gunong Kledang in Hill Dipterocarp Forest. J. Dransfield 871, Holotypus, June 1968; (K) et in monte Gunong Bubu, Whitmore FRI 0673. Very similar to Jt. altifrons Reichb. f. et Zoll. but differs from this in the ascending stem, forming a trunk, inflorescence with four orders of divaricate branching, sweet-smelling flowers, rounded bud, and the larger fruit with more humerous warts. 74 Gardens’ Bulletin, Singapore — X XVI (1972) Solitary palm, in age with a robust ascending stem to 4 m high and 15 cm in diameter, marked with annular scars. Leaves large, erect to 4 m high. Petiole to 1 m long, 2 cm wide. Lamina to 3 m long, to 1.6 m wide, clothed with brown scurfy scales along the underside of the costa, and at the short marginal lobes. Plicae numerous, to 20 or more on each side of the costa, with small anastomo- sing lateral nerves between the plicae. Leaf margin armed with short spines (to 1 mm) as is the petiole. Petiole marked with two yellow lines. Leaf base fibrous, withering to form a brown network of fibres. Inflorescence: axillary, arching out of the crown. Spathes covered in fu- gaceous scurfy scales, at first cream than becoming brown, 5—6 in number, 20 cm long and 8 cm wide, tubular at the base, inflated above, split on one side, with acute apices. Peduncle tomentose to 50 cm long and 2 cm wide, with four orders of divaricate branches. Floriferous branches 50 — 100, to 2.5 mm in diameter, to 10 cm long, greenish, covered in white tomentum, densely covered in flowers. Flowers glabrous, white, rounded in bud, solitary or grouped in 2’s or 3’s, borne on prominent tubercles, each subtended by a minute bracteole. Calyx short to 1 mm high, fleshy and glabrous, shallowly three-lobed. Petals fleshy, white, to 3 mm long, broadly triangular, connate in a short tube at the base. Staminal ring minutely epipetalous, abruptly contracted above into six filaments. Anthers oval. Pollen grains white. Ovary glabrous with three carpels free at the base, at the apex, connate into a common style 0.8 mm long. Fruit to 5 cm in diameter, — usually developing from one carpel, rarely from 2—3 carpels, covered in circa | 60 corky warts, dirty-brown in colour and 9 mm high. Endocarp woody to 1 mm thick. Endosperm bony to 2.5 cm in diameter, penetrated at the base by corky integumental tissue. Embrvo lateral. Malay Peninsula: Perak; on the hills of Gunong Kledang in Hill Dipterocarp forest. J. Dransfield 871, Holotype, June 1968: (K) and on Gunong Bubu, Whitmore ~ FRI 0673. COLLECTIONS EXAMINED 1 — H.N. Riprey (1898) Bukit Keledang, Perak (SING). 2 — C. CurtTIs 3350 (1898) Kledang, Kinta, Perak (SING). 3 — R. Lonc (1909) Kledang Hills, Perak (K). 4 — ex Penang Botanic Gardens (1909) source unknown (K). 5 — Mb. HANIFF and Mb. Nor, Sing. F.N. 6958 (1920) Sungei Siput, Perak (K, SING). 6 — Macuapo s.n. & s.a. Kamuning, Perak (SING). 7 — J. DRANSFIELD 871] (1968) Kledang Saiong Forest Reserve, Perak. (K) 8 — T. C. Wuitrmore, FRI 0673, Gunong Bubu, Perak (SING, KEP). HABITAT In the Kledang Saiong Forest Reserve near Ipoh, Perak, Jt. perakensis is an — abundant palm, growing gregariously on hill slopes and ridge-tops from about 175 m to 850 m. At the southern end of the range near Parit, it is equally abundant on hill slopes and ridge-tops; in the Virgin Jungle Reserve it is con- — fined to slopes and is absent from ridge-tops. Soils are well drained and derived — from granite. Much of the Forest Reserve has been selectively-logged, but Jt. © perakensis has survived the logging in many places. On Gubong Bubu, Whitmore — records the palm as locally very common on ridge-tops at 800 m. | Plate 5. JOHANNESTEIJSMANNIA PERAKENSIS Kledang-Saiong Forest Reserve, q Uy; Ws % % y Plate 6. JOHANNESTEIJSMANNIA MAGNIFICA (left) AND Jt. LANCEOLATA (right) S. Lalang Forest Reserve, Selangor. (Photo Courtesy of HO Sai Yuen, Kepong). The Genus Johannesteijsmannia H. E. Moore Jr. 7s TAXONOMIC NOTES Jt. perakensis and Jt. altifrons are more closely related than the other species of the genus. The separation of Jt. perakensis as a distinct species is justified on the basis of the characters in the diagnosis (see above) and the great isolation of the populations. The nearest populations of Ji. altifrons are in (1) West Kelantan separated from the Perak Hills by 100 km and the intervening Main Range rising to 2,400 m and nowhere in this region below 1,000 m, (2) Sungei Lalang in Selangor 200 km to the South, and (3) the Batang Serangan in East Sumatra 300 km to the West and separated by the Straits of Malacca. Isolation was probably even more effective during the Pleistocene at the maximum sea level, when, with sea level circa 80 m above present levels (Walker 1954), the Bubu massif and the Kledang-Saiong massif were probably isolated as islands off the main Malayan landmass. In badly collected specimens it is usually difficult to distinguish Jt. perakensis from Jt. altifrons. Infructescences are usually represented, and these always have divaricating floriferous branches. Johannesteijsmannia magnifica J. Dransfield, sp. nov. DIAGNOSIS Differt a speciebus ceteris foliis latissimis, lamina infra indumento lacteo tecta, inflorescentia ordinibus 5—6 ramosa, petalis angustis reflexis. DESCRIPTION Palma solitaria acaulescens, caule subterraneo procumbente ad 15 cm diametro. Folia erecta grandia latissima ad 5 m longa, lamina ad 3 m longa et 2 m lata, superne glabra, infra indumento lacteo vestita, costa marginibusque lobatis nudis. Plicae numerosae ad 20 utrinque inter plicas nervulis lateralis anastomosantibus. Margo inferior folii spinis brevibus (ad 15 mm) armatus atque petiolus. Petiolus duobus striis conspicuis notatus. Vagina folii fibrosa ad 25 cm longa marcescens reticulum fuscum formans. Inflorescentia axillaris e cumulo foliorum mortuorum et fragmentorum plantorum aliorum in corona collectante arcuata. Spathae primo cremeae, dein fuscentes, 5—6, 26 cm longae et 7 cm latae, basi tubulosae superne inflatae, apicibus acutis, uno latere fissae indumento albo dense tectae. Pedunculus tomen- tosus ad 80 cm longus, 2 cm diametro, ordinibus 5— 6 ramosus. Rami floriferi 500 — 1,000, ad 1.5 mm diametro ad 7 cm longi albi glabri, floridique. Flores glabri, alabastro acuti, solitarii vel bini vel terni aggregati, in tuberculi prominenti- bus insidentes, unusquisque bracteola minuta subtentus. Calyx brevis ad 0.5 mm altus, carnosus glaberque, paullo 3 —lobatus. Petala ad 15 mm longa, anguste- triangularia, 3— plo longiora quam latiora, carnosa alba, reflexa, basi in tubo breve connata, intra rugosa. Annulus staminalis minute epipetalus, in filamenta 6 aequalis superne abrupte subulata. Antherae ovales. Grana pollinis alba. Ovarium glabrum carpellis 3, basi liberis, apice in stylo communi 0.4 mm _ longo connatis. Fructus ad 4 cm diametro, plerumque a carpello uno evolutus aliquando a 2—3 carpellis, c. 150 verrucis humilis suberosis fusco-rufescentibus, ad 2.5 mm altis, tectus. Endocarpium lignosum ad 1 mm crassum. Endospermium Osseum ad 2.5 cm diametro, contextu integumentorum suberoso basi penetratus. Embryo lateralis. Gardens’ Bulletin, Singapore — XXVI (1972) 76 Fig. 4. JOHANNESTEIJSMANNIA MAGNIFICA A: inflorescence, B: flower The Genus Johannesteijsmannia H. E. Moore Jr. ao Malay Peninsula: Selangor, in valle humido, Virgin Jungle Reserve Sungei Lalang Forest Reserve, Semenyih, Kajang, in Hill Dipterocarp Forest. J. Dransfield 862, Holotypus, May 1968 (K); et Negri Sembilan in monte Bukit Tangga, Seremban, Napier 1903 (SING) et Dransfield (1967); et in monte Gunong Angsi, Corner. Differing from the other species by the very broad leaves with the lamina covered with white indumentum below, the inflorescence with 5—6 orders of branching, and the flowers with narrow reflexed petals. Solitary acaulescent palm with procumbent underground stem to 15 cm in diameter. Leaves erect, large and very broad, to 5 m tall, with lamina to 3 m long by 2 m broad, glabrous above, on the lower surface covered with white indumentum except along the costa and at the marginal lobes. Plicae numerous to 20 or more pairs on either side of the costa, with small anatomosing lateral nerves between the plicae. Lower leaf margin armed with short spines (to 1.5 mm) as is the petiole, Petiole with two conspicuous yellow lines. Leaf base fibrous, to 25 cm long, withering to form a brown network of fibres. Inflorescence axillary, arching out of the pile of dead leaves and other plant debris accumulating in the crown of leaves, Spathes cream at first then becoming brown, 5— 6 in number, 25 cm long and 7 cm wide, tubular at the base, inflated above, split down one side, with acute apices, and densely covered in white indumen- tum. Peduncle tomentose to 80 cm long, 2 cm in diameter, with 5 — 6 orders of branching. Floriferous branches 500 — 1,000 in number, to 1.5 mm in diameter and 7 cm long, white, glabrous, and densely covered in flowers. Flowers white and glabrous, acute in bud, solitary or grouped in 2’s and 3’s, borne on prominent tubercles, each subtended by a minute bract. Calyx short, to 0.5 mm high, fleshy and glabrous, shallowly three-lobed. Petals to 1.5 mm long, narrowly triangular, three times as long as broad, white and fleshy, reflexed, connate at the base into a short tube, and rugose within. Staminal ring minutely epipetalous, abruptly narrowed above into six equal filaments. Anthers oval. Pollen grains white. Ovary glabrous with three carpels, free at the base, connate at the apex into a common style 0.4 mm long. Fruit to 4 cm in diameter, usually developing from one carpel, occasionally from 2— 3 carpels, covered by c.150 low, reddish-brown corky warts, to 2.5 mm high. Endocarp woody to 1 mm thick. Endosperm bony to 2.5 cm in diameter, with corky integumental tissue penetrating at the base. Embryo lateral. Malay Peninsula: Selangor, in humid valley, Virgin Jungle Reserve, Sungei Lalang Forest Reserve, Semenyih, Kajang, in Hill Dipterocarp Forest. J. Dransfield 862, Holotype, May, 1968 (K); and Negri Sembilan on Bukit Tangga, Seremban, Mapier 1903 (SING) and J. Dransfield (1967); and on Gunong Angsi, Corner. COLLECTIONS EXAMINED 1 — H. J. Napier (1903) Bukit Tangga, Negri Sembilan (SING). 2 — J. DRANSFIELD (1967) Bukit Tangga, Berembun Forest Reserve, Negri Sembilan. 3 — WonGc YEW Kwan 95088 (1961) Cpt. 24, Sungei Lalang Forest Reserve, Selangor (KEP). 4 — J. DRANSFIELD (1967) Cpt. 3, Sungei Lalang Forest Reserve, Selangor. 5—J. DRANSFIELD 862, (1968) Virgin Jungle Reserve, Sungei Lalang Forest Reserve, Selangor. 78 Gardens’ Bulletin, Singapore — X XVI (1972) HABITAT In all localities visited, Jt. magnifica is a plant of steep slopes and ridge-tops except in the Virgin Jungle Reserve, Sungei Lalang Forest Reserve where it also grows at the side of a small river in a very humid valley bottom, alongside Jt. lanceolata; in this last locality Jt. magnifica occurs in great abundance in the untouched forest, from the valley bottom at 150 m to the ridge-top at 500 m. At Bukit Tangga in Berembun Forest Reserve in Negri Sembilan, Jt. magnifica grows in much disturbed, logged forest. TAXONOMIC NOTES As Jt. magnifica and Jt. lanceolata grow side by side in the Virgin Jungle Reserve (see plate 6) it is possible that hybridization between the two species could take place. As any hybridization experiments would be extremely prolonged, difficult to perform, and dependent on simultaneous flowering, it has only been possible to use indirect evidence to assess the possibility of hybridization taking place. There are no intermediates observable in mature plants; in young plants the presence of white indumentum on the lower leaf surface is always correlated with laminae about 14 times as long as broad. Narrower leaves are always giabrous except for scurfy brown scales along the midrib. Narrow leaved adults are always glabrous and always have the inflorescence and fruit characters of Jt. lanceolata. These facts suggest that hybridization does not occur. Johannesteijsmannia lanceolata J. Dransfield sp. nov. DIAGNOSIS Differt a speciebus ceteris foliis angustis, inflorescentia ramis 3— 6, crassis, petalis papillatis. DESCRIPTION Palma solitaria acaulescens, caule subterraneo procumbente ad 15 cm dia- metro. Folia erecta, plus minusve lanceolata ad 3.5 m alta. Lamina ad 2.4 m longa et 30 cm lata, squamis furfuraceis fuscis infra secus costam et nervos primarios tecta. Plicae numerosae ad 25 utroque costae latere, inter plicas nervulis lateralis anastomosantibus. Margo inferior folii spinis brevibus (ad 1 mm) armatus atque petiolus. Petiolus duobus striis luteis conspicuis notatus. Vagina folii fibrosa ad 25 cm longa marcescens, reticulum fuscum formans. Inflorescentia axillaris inter petiolos infra cumulum foliorum mortuorum et fragmentorum plantorum aliorum infossa. Spathae primo cremeae dein fuscentes, caducae, 3 — 4 indumento fusco sparsim tectae, ad 20 cm longae et 2 cm latae, basi tubulosae, non inflatae. apicibus acutis, uno latere fissae. Pedunculus tomentosus ad 25 cm longus, 10 mm diametro, ordine uno ramosus. Rami floriferi 3 — 6, crassi ad 10 mm diametro et 16 cm longa, pubescentes, arcuati rective. Flores sessiles albi papillosique, alabastro obtusi, solitarii vel bini vel terni aggregati, unusquisque bracteola minuta subtentus, spiraliter dispositi. Calyx brevis, 0.3 mm altus, carnosus glaberque, paullo 3—lobatus. Petala ad 1 mm longa, late-triangularis, carnosa crassissima, basi in tube breve connata, extus papillosa, intra apices aliquantum verruculosa. Annulus staminalis minute epipetalus, in filamenta 6 aequalis superne abrupte subulata. Antherae ovales. Grana pollinis alba. Ovarium glabrum carpellis 3, basi liberis, apice in stylo communi 0.3 mm longo connatis. Fructus ad 3.4 cm diametro, plerumque a carpello uno evolutantes, saepe a 2 — 3 carpellis, c. 90 verrucis humilibus suberosis fusco- rufescentibus, ad 2.5 mm altis, tectus. Endocarpium lignosum ad 1 mm crassum. | q 79 The Genus Johannesteijsmannia H. E. Moore Jr. Fig. 5. JOHANNESTEIJSMANNIA LANCEOLATA inflorescence, B: flower A: 80 Gardens’ Bulletin, Singapore — X XVI (1972) Endospermium osseum ad 2.5 cm diametro, contextu integumentorum suberoso basi penetratus. Embryo lateralis. Malay Peninsula: Selangor, in valle humido prope flumen, Virgin Jungle Reserve, Sungei Lalang Forest Reserve, Semenyih, Kajang, in Hill Dipterocarp Forest. J. Dransfield 861, Holotypus, May 1968; (K) et Pahang, in Ulu Sungei Tekal Besar, Temerloh, Henderson March 1923 (SING). Differs from the other species by the narrow leaves, the inflorescence with 3 —6 thick branches and the flowers with papillate petals. Solitary acaulescent palm, with subterranean procumbent stem to 15 cm diameter. Leaves erect, + lanceolate, to 3.5 m tall. Lamina to 2.4 m long and 30 cm wide covered with brown scurfy scales on the underside, along the midrib and the primary nerves. Plicae numerous to 25 along each side of the costa, with small anastomosing veins between the plicae. Lower leaf margin with short spines (to 1 mm) as on the petiole. Petiole with two conspicuous yellow stripes. Leaf sheath fibrous to 25 cm long, withering to form a brown network of fibres. Inflorescence axillary buried between the petioles under a pile of dead leaves and other plant debris. Spathes at first cream, then turning brown, caducous, 3 — 4 in number, sparsely clothed in brown indumentum, to 20 cm long and 2 cm wide, tubular at the base, not inflated, acute at the apex, split down one side. Peduncle tomentose to 25 cm long, 10 mm in diameter, with one order of branching. Floriferous branches 3 — 6 in number, thick, to 16 cm long and 10 mm in diameter, pubescent, curved or straight. Flowers sessile, white and papillate, obtuse in bud, solitary or grouped in 2’s and 3’s, each subtended by a minute bracteole, arranged in a spiral on the branch. Calyx short 0.3 mm high, glabrous and fleshy, shallowly three-lobed. Petals to 1 mm long, broadly triangular, fleshy, and very thick, connate into a short tube at the base, papillose without, somewhat verrucose within towards the apex. Staminal ring minutely epipetalous, abruptly contracted into six equal filaments. Anthers oval. Pollen grains white. Ovary glabrous, carpels three, free at the base, connate at the apex to form a common style 0.3 mm long. Fruit to 3.4 cm in diameter, usually developing from one carpel, often developing from two or three, covered with c.90 low corky warts, reddish-brown in colour, to 2.5 mm high. Endocarp woody to 1 mm thick. Endo- sperm bony to 2.5 cm in diameter, penetrated at the base by corky integumental tissue. Embryo lateral. Malay Peninsula: Selangor in humid valley by the river, Virgin Jungle Reserve, Sungei Lalang Forest Reserve, Semenyih, Kajang, in Hill Dipterocarp Forest. J. Dransfield 861, Holotype, May 1968: (K) and Pahang in Ulu Sungei Tekal Besar, Temerloh; Henderson, March 1923 (SING). COLLECTIONS EXAMINED 1 — Henderson (1923) Ulu Sungei Tekal Besar, Temerloh, Pahang (SING). 2 — J. Dransfield 861 (1968) Virgin Jungle Reserve, Sungei Lalang Forest Reserve, Selangor (K). HABITAT In Sungei Lalang Virgin Jungle Reserve, Jt. lanceolata is known from the steep banks of a forest river in a deep humid valley; growing with Jt. magnifica. Despite the proximity of the river, Jt. lanceolata grows on well drained soils and not in the low swampy ground at the bends in the river. The population of Jt. lanceolata was probably once greater—further down the valley scattered individuals occur growing in logged forest near the river. The Genus Johannesteijsmannia H. E. Moore Jr. 81 The Geography of Johannesteijsmannia Johannesteijsmannia is confined to North Sumatra, the Malay Peninsula, and West Borneo, with one widespread species Jt. altifrons, and three species endemic to the Malay Peninsula. The concentration of species in the Malay Peninsula may be a reflection of the more intensive fieldwork carried out in - Malaya; there are, for example, as far as in known, no post 1940 specimens of Jt. altifrons from Sumatra, and the possibility of new species of the genus occurring in Sumatra cannot be ruled out. Jt. altifrons, however, in Sarawak appears to be genuinely rare, and Forest Staff and Ibans in Brunei did not know of the existence of this very conspicuous and useful palm. It seems most reasonable, however, to suppose that the centre of diversity of the genus coincides with present day centre of distribution —1i.e. the uplands of Selangor, Negri Sembilan and Pahang, where three of the four species grow together. In late Mesozoic time this area was somewhere near the centre of the Sundaland mass (van Bemmelen 1949, Umbgrove 1949), and this area together with West Sarawak, East Sumatra, Banka and Billiton, has remained above sea level until the Pleistocene marine incursions (Umbgrove l.c.) when it seems probable (extrapolation from Malaya) that much of Banka and Billiton and of lowland East Sumatra would have been submerged during the maximum sea level of 80 m (Walker 1954). It is the area of Sundaland now represented by Malaya and Borneo which probably remained as land through the Pleistocene, that harbours the greatest concentration of species in the palm genera Licuala, Calamus, Daemonorops, and Korthalsia. (the number of species of Licuala on New Guinea exceeds the number in Malaya and Borneo individually, but if Malaya/Borneo is treated as one land mass, i.e. part of Sundaland, then the total exceeds that of the land mass of New Guinea and its outlaying islands). The limited distribution of the genus can be explained in two main ways: (1) the genus is an ancient one and survives as a relatively unsuccessful form in the area of its diversity as a relic of Coryphoid evolution, (2) the genus has relatively recently evolved and has not yet spread to areas outside its area of origin. That Johannesteijsmannia has many characters in common with Licuala but is in many ways more primitive than Licuala (e.g. leaf and inflorescence structure) suggests it is probably more ancient than the successful widespread genus Licuala. It therefore seems more likely that Johannesteijsmannia is an ancient genus surviving in Sundaland. Speculating further I would suggest an origin during the Mesozoic from primitive Coryphoid stock. If a Mesozoic origin of the genus in the Malay Peninsula is assumed then the spread away from the centre could have occurred any time during the late Mesozoic and the Tertiary. Any pre-Miocene invasion of Sumatra would probably have been destroyed by marine incursions, for much of Sumatra was submerged during the Miocene (Umbgrove 1949). Only one species at the present day, Jt. altifrons, has spread out of the proposed centre of diversity, and is therefore to be regarded as the most successful and adaptable species. Jt. altifrons probably reached Sumatra during the Pliocene, and was then cut off from the Malayan populations by the appearance of the Straits of Malacca. Jt. altifrons could have reached Sarawak any time before Borneo was separated from the Malay Peninsula during the Pleistocene, but the very local occurrence of the species in West Sarawak suggests 82 Gardens’ Bulletin, Singapore — X XVI (1972) that the invasion of Borneo is relatively recent, (e.g. in the Pliocene corresponding with the proposed time of spread into Sumatra). Jt. altifrons, although it has reached Sarawak, is absent from Pulau Tioman (Pahang), Penang and the Anamba and Natuna Islands; the absence may be due to chance extinctions if the palm ever colonized these islands or else it was unable for some reason to reach the islands. The distribution of Jt. altifrons within the Malay Peninsula is disjunct and it seems reasonable to suppose that Jt. altifrons was at one time widespread over the whole area; this may have coincided with the proposed period of active expansion which resulted in the colonization of Sumatra and Borneo. The disjunction of the range of distribution has resulted in slight vicarous evolution; the isolated popu- lations have diverged slightly from each other. In Perak, populations on the Bubu Range and Gunong Kledang have diverged further and reached a degree of divergence where it is convenient to regard them as a distinct species, Jt. perakensis. The evolution and distribution of Jt. lanceolata and Jt. magnifica can- not be explained by vicariism (as Croizat 1968 would explain the speciation of Ficus — refuted by Corner 1969); the two species occur alongside Jt. altifrons in the one Forest Reserve in Selangor, apparently with the same ecological require- ments. It is most reasonable to regard Jt. lanceolata and Jt. magnifica as rela- tively unsuccessful early products of form making within the genus which have survived near the centre of diversity without having spread. (Jt. lanceolata has reached central Pahang from the proposed centre of diversity). The dispersal powers of Jt. altifrons appears to be very limited — seedlings are rare, — and the distribution of the whole genus suggests plants of very limited dispersal powers at the present day, surviving as relics in the centre of massing of the Asiatic coryphoid palms, as do Liberbaileya gracilis and Maxburretia rupicola (Furtado 1941). Acknowledgements This study was carried out during the tenure of a Royal Society Leverhulme Scholarship and a Science Research Council NATO studentship, and represents part of a thesis submitted in Cambridge for the degree of Doctor of Philosophy. I should like to thank Professor E. J. H. Corner for supervising my work, Dr. T. C. Whitmore for an inestimable amount of help in Malaya and England, and all the people, too numerous to mention, who helped me in my field work. Bibliography Beccari, O. (1931): Asiatic Palms — Corypheae. Revised and edited by U. Martelli. Ann. R. bot. Gard., Calcutta. 13. Bemmelen, R. W. van (1949): The Geology of Indonesia. Government Printing Office, the Hague. Burkill, I. H. ((1935) 1966): A Dictionary of the Economic Products of the Malay Peninsula Vols. 1 and 2 Government of Malaysia and Singapore. : The Genus Johannesteijsmannia H. E. Moore Jr. 83 Corner, E. J. H. (1969): The complex of Ficus deltoidea; a recent invasion of the Sunda Shelf. Ficus sect. Adenosperma. Phil. Trans. R. Soc., 256, 281 —- 355. Croizat, L. (1968): The biogeography of the tropical lands and islands east of Suez-Madagascar: with particular reference to the dispersal and form-making of Ficus and different other vegetal and animal groups. Atti Ist. bot. Univ. Lab. crittogam. Pavia (Ser 6), 4, 3 — 400 Faegri, K, and Pijl, L. van der (1966): The Principles of Pollination Ecology. Pergamon Press, Oxford and London. Furtado, C. X. (1941): Palmae Malesicae IX. Two new Coryphaceous genera in Malaya. Gdns’ Bull. Straits Settl., 77, 236 — 43. Gatin, C. L. (1906): Recherches Anatomiques et Chimigues sur la Germination des Palmiers. Annls Sci. nat. (Botanique) ser. 9.3, 191 — 314. Miquel, F. A. W. (1857): Nova genera Apocynearum Indicarum. Versl. med K. Akad. Wetens. Amsterdam, 6, 191 — 194. (1861): Revue des Palmiers de Vile de Sumatra. J. Bot. Néerlm. 7, 1-29. Miquel, F. A. W. (1868): Teysmannia Zolling. Palmarum genus. Ann. Mus. Bot. Lugduno — Batavum. 4, 89 — 90. Moore, H. E. Jr. (1961): Johannesteijsmannia—a new name for the Palm genus Teysmannia. Principles, 5, 116. Palm, B. T. and Jochems, S.CJ. (1924): De sang-palm van Sumatra Trop. Natuur, 13, 9— 12. Richards, P. W. (1952): The Tropical Rain Forest. Cambridge. Ridley, H. N. (1925): Flora of the Malay Peninsula Vol. 5. L. Reeve, London. Robbins, R. Q. (1968): The Biogeography of Tropical Rain Forest in S.E. Asia. Aoc. Symp. Recent Adv. Trop. Ecol. 521 —535. Teysmann, J. F. (1957): Dag verhaal eener Botanische Reis over de Westkust van Sumatra. Natuurk. Tijdsch. Ned. Indié 14, 320 — 22. Tomlinson, P. B. (1960): Essays on the Morphology of Palms. 1. Germination and the seedling. Principes 4, 56 — 6l. Umbgrove, J. H. F. (1949): Structural History of the East Indies. Cambridge. Walker, D. (1954): Studies in the Quaternary of the Malay Peninsula (1) Alluvial deposits of Perak and Changes in the relative levels of land and sea Fed. Mus. 1. i +. 2, 19 — 39. Zollinger, H. (1856) (1858): Uber ein neues Palmengeschlecht von der Insel Sumatra. Linnaea 28, 657 — 60. ; pf nize iain higact & nr ot FAG Seat ‘ I BR at 7 on Py i. > eae i? oe i Dy ) > wart ga Teas bes niet: onthe sme Metals fies MN vglgnee, dnceetter hth. “anieGnedn iewere’ tint a Ubtosk , ’ % oniaal | okra angled andaipelgg 5 A@OR Don bli LOL. Re eae (salen es vag pee 4 : Anh re Si ) ; aT ats -* ' viited oi i Lee tie Bus oe 0 a (eat) =. r. i is "i TUTORS soetndtadt aa 7 a ? = Pi Pally ST Gat oe alle nea ip, L raeteurte? aly tai ob sient nb ne'¥s god Bho - | iw i ' mic’ .oetht) sitatires i ry Re a : VA La yh ry) ) a ari \ i] : mw) 1) 7 | ‘ = 4 ui be wh. ‘ rd etwas lacing arg clepitiod cA Anadis : ; P e 7 . pape: (yon: peupiae Sy yt ic-2AF id | Bee rT ates its te +. 3 ) hon wv Aseoyt Heowerm) - «ole. Wo sea ct oils. uo Nati ae ; ibn bt a ee ‘ - mika ehisduu ade yp tveity Jppcwonilh, Ww may Rcvhts ni ’ Weads wae > "fe eS iw 4 =) a laivollA Th} ido of (Gln ve ‘on ian (amend. a ai ee A carob beaut Waele ih iprisdlian ila sath Sickness one 7 Bo) si eae Re 7 a) a aby Lari reerngy, i eli a ' ee : Poodunila ’ ay 4 Soil moisture under forest, Bukit Timah Nature Reserve, Singapore By R. D. HILyi University of Singapore Abstract The results of a study of moisture of three forested hill-slope soils are presented and compared with the results of a similar study in Malaya. Soil moisture levels were found in general to be rather higher than under other forms of vegetation. Statistical analysis indicated that change of moisture content was consistent within each soil but less consistent between soils. A series of statistical experiments suggested that except for the A soil horizon, soil moisture (as determined by drying) correlated reasonably well with number of raindays and that the introduction of the factor e (evapotranspiration) resulted only in slightly improved correlations except for the A soil horizon where the use of e gave much better correlations. The study of soil moisture in the region is one that is strangely neglected, a search of available literature failing to reveal more than two studies, both concerned with soil moisture as related to rubber. Belgrave (1930, 493-4) made observations from three levels, 2-6 in., 6-12 in. and 12-18 in., of a Serdang series soil having two types of cover, Centrosema, Mikania, and under bare soil. Watson, Wong and Narayanan (1963) made eighteen observations at monthly intervals from two levels, 0-6 in. and 12-18 in. of a Rengam series coarse sandy clay loam with a vegetation of two and three year old rubber and cover crops of legumes, grass, adventives and also under bare soil. Soil Characteristics The present study reports the results of weekly observations made from mid-February until mid-June 1970 at three points down a surveyed transect line off Jungle Fall path, Bukit Timah Nature Reserve which has a somewhat open lowland Dipterocarp-forest. The upper sampling point, 1, (172 m a.s.l.) was some 25 m from the crest of a gently-rounded interfluve and about 4 m below it with a local slope of almost 15°. The middle sampling point, 2, (141 m a.s.l.) was about 87 m downslope from the first sampling point, 1, and it had a local slope of 24°. Both soils belong to the Rengam series (Leamy & Panton 1966, 105). The lowest sampling point, 3, (126 m a.s.l.) was at the foot of the same slope, 33 m from 2, and located on a small terrace 1.5 m above the level of a small perennial stream. The soil materials were derived from colluvium and alluvium with the former predominating. This soil cannot be assigned to any described series. 85 86 Gardens’ Bulletin, Singapore — XXVI (1972) The moisture-holding capacity of soil is significantly influenced by grain-size and organic material present. These data are presented for four levels in each soil, 0-10 cm (04 in.), 10-20 cm (4-8 in.), 30-45 cm (12-18 in.) and 60-75 cm (24-30 in.), in Table 1. Detailed analysis of grain size was made using the A.S.T.M. standard method (American Society for Testing and Materials, 1963). Only a summary of the data is presented here. The data for loss on ignition (at 480°C) do not accurately measure total organic matter because the initial oven-dry samples still retain tightly-bound water of hydration. Total organic matter is thus slightly lower than indicated in Table 1. Table 1. Grain Size (%) according to International System and Loss on Ignition Sample Depth Coarse Sand Fine Sand Silt and Clay Loss on : Textural name Ignition % Point (cm) —_(2000,-250.) (250-25) (less than 25,,) pecan 1 0-10 44.6 16.7 38.7 Coarse sandy loam 7 10-20 39.3 12.9 47.8 Coarse sandy loam* 8.8 30-45 38.2 13.3 48.5 Coarse sandy loam* is 60-75 34.1 22 DAGL Coarse sandy loam 7.0 2 0-10 40.3 21.9 37.8 Coarse sandy loam 10.3 10—20 Bd nl i338 47.0 Coarse sandy loamt 5.8 30-45 36.2 18.1 45.7 Coarse sandy loam+ 4.5 60-75 33.5 20.6 45.9 Sandy clay loam a7 3 0-10 60.0 5.9 34.1 Not namedt 52 10-20 57.6 20.8 21 & Coarse sandy loam 5.1 30-45 56.7 16.1 26.3 Coarse sandy loam 4.4 60-75 38.5 17.5 440 Coarse sandy loamt fe * Almost sandy clay + Almost sandy clay loam t Suggested name: coarse sandy clay Collecting Methods Prior observation established that the maximum variability amongst samples taken from the same level at the same point on the slope was + one per cent provided that augered samples were taken at points where there were no large roots and which were some distance away from the stems of nearby trees. Successive weekly samplings thus required only one sample from each level at each observation point. Several samples taken at point 3 were notably coarser than the majority and values for moisture are therefore less reliable. Samples of about 300 gm were augered from each level and immediately placed in polythene bags and sealed. On arrival at the laboratory samples were well-shaken or stirred to incorporate moisture condensing on the inside of the bag and to break down lumps. Samples of about 100 gm were then spooned into dishes for drying at 105°C for 24 hours followed by weighing at 40°C. Prior experiment showed that differences arising from more protracted drying and from cooling in a desiccator to room temperature were much smaller than the differences arising from variations in the exact point sampled. It was however, necessary to weigh dried samples very rapidly taking them one by one from the oven. LS _eset eee Soil Moisture under Forest, BT Nature Reserve, S’ pore 87 Weather The period of observation began with the dry spell which usually occurs towards the end of the north-east monsoon, and extended through a wetter inter-monsoon period to the early part of the south-west monsoon. Rainfall data were collected daily from a rainfall station some three hundred metres from the soil sampling points. Open pan evaporimeter data were obtained for Paya Lebar Airport some 14 km from Bukit Timah. Soil samples were taken shortly after the daily rainfall measurement was made. Rainfall data are presented at the top of each diagram in Figures 1-3. Results The results are presented in Figures 1-3. The diagrams suggest a fairly clear relationship between rainfall and soil moisture, the general trend of low values during dry spells being clear. In the two hill-slope soils points 1 and 2, soil moisture conditions were fairly stable below 30 cm with increasing variability in the upper portions of the soil. This was more marked in the 0-10 cm zone which corresponded with the A, soil horizon. Rather large fluctuations, sometimes even during fairly dry periods reflect large but short-lived rises of soil moisture following showers. The mid-slope soil was slightly drier than the upper-slope soil during the main dry spell but slightly wetter during the remainder of the period. This suggests that greater slope angle and hence more rapid soil drainage becomes influential only during dry spells whereas at other times receipt of upslope water results in slightly moister conditions. Differences of soil texture would seem to be too minor to account for these differences. Furthermore the mid-slope soil contained rather less organic matter than the upper-slope soil and this should have resulted in lower moisture values for the former soil had this been an influential factor. The slope-foot soil shows an almost complete reversal of the pattern of the other two soils. In it variation was greatest at depth, undoubtedly because of fluctuations in the position of the water-table. The upper zones of this soil were a good deal sandier than those of the hill-slope soils, organic matter was much less and rapid infiltration of rainfall and overland flow from upslope occurred. The range of moisture is summarised in Table 2. Discussion Only the study by Watson, Wong and Narayanan (1963) presents data adequate for some comparisons to be drawn. Expectably the soil moisture values are considerably higher under forest than those under legumes, grass, adventives and bare soil reported in that study. The lowest values at Bukit Timah are roughly equivalent to the mean values at Serdang for the upper 20 cm of soil. The data of Watson et al (1964, 165) also show much greater variation of moisture in the 0-15 cm zone than in the 30-45 cm zone. They report moisture values for the 30-45 cm zone which correspond quite well with those of the present study. co oe) Gardens’ Bulletin, Singapore — XXVI (1972) 0090 > 15 05 # 30 1-0 2 1-5 z 2-0 60 2 25 75 = 305 90 2 seltantr isle bhnials oO] 4” (0 -10¢cm.) 30 mi CU iets See 4- 8" (10-20cm.) : 12-18" (30- 45cm.) DAILY RAINFALL (MM.) 5 oa 24-30" (60-—75cm.) 25 LOSS OF WEIGHT (*%.) ceca eaieamam) Soammaieien®| ‘nearer Giese Tiers (Alia ||" ieee [ieee en Dees |? ima: aanann aaa aM! (Pitan Eels ep See 4 11 18 25 4 1113 18 24 1 7 15 22 29 6 13 20 27 - 10 “f FEBRUARY MARCH APRIL MAY JUNE Fig. 1 Soil moisture (loss of weight on drying) for the soil near the top of the slope (1) and daily rainfall. Table 2. Range of Soil Moisture (%) Sample Depth Date Date Ban. ie Highest (1970) Lowest (1970) Range ] 0-10 30.2 18.-2 2h3 4-3 8.9 10-20 26.7 22-4 19.9 114 6.8 30-45 24.7 13-5 19.5 11-3 5.2 60-75 23.5 13--5 20.2 24-3 3.3 2 0-10 31.6 aah 20.0 113 11.6 10-20 218 10-6 17.9 11-3 99 30-45 23,5 13-5 18.6 4-3 49 60-75 26.4 22-4 20.1 11-3 6.5 3 0-10 23.4 22-4 18.5 4-3 4.9 10-20 22.9 22-4 20.2 25-2 ay 30-45 PAS Be 22-4 10.1 13-5 6.7 60-75 30.4 22-4 19.7 1-4 10.7 Statistical Analysis Two sets of statistical analyses of the hill-slope soils (1 and 2) were made. The first aimed to establish the extent to which the rate of change of soil moisture between successive observations at each level within the soil were correlated. The results are summarised in Table 3 which shows that in general there is a greater degree of association between moisture change at one Jevel within the soil and moisture change at the level below it than between moisture change at the same level of different soils. In other words rates of change within a soil were internally consistent with the notable exception of the soils at 60-75 cm where poor or no correlations with the soils at 30-45 cm were found. Soil Moisture under Forest, BT Nature Reserve, S’ pore 89 2 00 9 = 15 0-5 e e 30 1:0 > 45 re z z 20 > é 60 25° 7 75 3-05 & 90 35 2 > 30 . Oe ee O=- 4" (O0-10cm.) : eo oqyoccccce 4- 8" (10-20cm.) : E 12-18" (30-—45cm,) : Ww ——$——_— 24-30" (60-75¢m.) : TS ee ore 3 25 é a ny i 4 : re) Ww yn } 20 a ee eae fee a eee ee i . ahit«l: i. Tr 1 4 11 18 25 4 1113 18 24 1 Aen 15 22 29 6 13 20 27 4 10 17 “FEBRUARY MARCH APRIL MAY JUNE Fig. 2 Soil moisture (loss of weight on drying) for soil on the middle of the slope (2) and daily rainfall. DAILY RAINFALL (MM.) wo ~ oa > w ak oO o oO oo o oO Ww WwW RH A _ — o Oo omovwvoWws#owso CSNI) WIW4NIv¥ Ativa 2 Ree O- 4” (0-10cm.) ee ee 2 es 4- 8” (10-20cm.) 12-18" (30-—45cm.) 24 -30" (60-75 cm.) (*/e) OBSERVATIONS DISCONTINUED LOSS OF WEIGHT [oF SA ae EEL et el a ee Tis Tepe ea cia iets eae mea area Bees lee - 11 18 25 & 11:13 18 24 1 7 15 22 29 6 13 20 27 . 10 17 FEBRUARY MARCH APRIL MAY JUNE Fig. 3 Soil moisture (loss of weight on drying) for the slope-foot soil (3) and daily rainfall. 90 Gardens’ Bulletin, Singapore — X XVI (1972) The second set of statistical analyses attempted to find a simple parameter with which soil moisture was correlated. Since soil moisture is affected by a large number of factors including endogenous factors such as grain size and structure and exogenous factors such as rainfall and evapotranspiration, it was hardly to be expected that with only 19 successive observations statistically significant correlations between soil moisture and rainfall alone would be found. Such proved to be the case. Soil moisture was compared with the total rainfall over varying numbers of days prior to each observation. The best, but still poor correlations were first, for moisture in soils 1 & 2, (10-20 cm), with total rainfall over the fourth, third and second days before observation; r = + 0.47, and second, for moisture in soils, 1 & 2 (60--75 cm), with total rainfall over the fifth, fourth and third days before observation; r = + 0.47, both values for r being significant at the 5 per cent level with 24 degrees of freedom. Student’s ¢ test was used to test significance in each case. Table 3. Correlations of Rates of Change of Soil Moisture at Various Levels in Soils 1 and 2 Variables Degrees of Significance ¥ y freedom / (better than) 1 (0-10 cm) 2 (0-10 cm) 16 +0.76 i% 1 (10-20 cm) 2 (10-20 cm) 18 +(0).49 5% 1 (30-45 cm) 2 (30-45 cm) 18 + 0.57 hy 1 (60-75 cm) 2 (60-75 cm) 18 + 0.56 1% 1 & 2 (0-10 cm) 1 & 2 (10-20 cm) 34 +0.71 0.1% 1&2(10-20cm) 1&2 (30-40 cm) 36 +0.70 0.1% 1&2(3040cm) 1&2 (60-75 cm) 36 + 0.46 19, 1 (0-10 cm) 1 (10-20 cm) 18 +(0.71 0.1% 1 (10-20 cm) 1 (30-45 cm) 18 +0.76 0.195 1 (30-45 cm) 1 (60-75 cm) 18 + 0.68 1% 2 (0-10 cm) 2 (10-20 cm) 16 + (0.73 1% 2 (10-20 cm) 2 (30-45 cm) 18 + 0.60 1% 2 (30-45 cm) 2 (60-75 cm) 18 + 0.38 None A similar comparison of rate of soil moisture change with rainfall was even less satisfactory A comparison between soil moisture and number of raindays (> 0.1 in. or 2.5 mm per day) in the week preceding measurement gave rather better results and these are shown in Table 4 together with correlations of rate of moisture change with number of rain-days in the preceding week. Soil Moisture under Forest, BT Nature Reserve, S’ pore 9] Table 4. Correlation of Soil Moisture and Rates of Soil Moisture at Various Levels with Number of Rain-days in Preceding Week Soil Degrees of Significance x y freedom is (better than) Soil m, 1 & 2, (0-10 cm) t/days 38 + 0.44 jibes Soil m, 1 & 2,(10-20cm) ___r/days 38 + 0.56 0:1:% Soil m, 1 & 2, (30-45 cm) r/days 38 +0.53 0.1% Soil m, 1 & 2, (60-75 cm) r/days 38 +0.55 Oke, Rate of s.m. change, 1 & 2, (0-10 cm) t/days 34 + 0.56 0.1% Rate of s.m. change, 1 & 2, (10-20 cm) r/days 36 +().49 1% Rate of s.m. change, 1 & 2, (30-45 cm) r/days 36 +(0).47 oe Rate of s.m. change, 1 & 2, (60-75 cm) r/days 36 + 0.47 1% In an attempt to gain rather better correlations the factor for evapotranspira- tion (e) was introduced. In this, e¢ is evaporation, multiplied by the empirical constant 0.9. The actual formula used was R — e, where R = daily rainfall and e the daily evapotranspiration. Values of R—e were summed for varying numbers of days prior to each determination of soil moisture. The selected best results are given in Table S. ; Table 5. Selected Correlations of Rates of Change of Soil Moisture with R — e over Various Days before Moisture Determination si Bnei wcleses tine tro > teen in 1 & 2, (0-10 cm) 3,4204 34 +0.57 Pes 1 & 2, (0-10 cm) I, say 34 + 0.60 1%, 1 & 2, (0-10 cm) dei44 3 34 +0.45 3% 1 & 2, (10-20 cm) 4, 3,2 36 + 0.44 3% 1 & 2, (10-20 cm) 5, 4, 3 36 +0.31 Nene 1 & 2, (30-45 cm) 3, 4.3 36 +-0.37 None 1 & 2, (60-75 cm) >, 4, 3 36 + 0.42 5% It will be seen from Table 5 that the introduction of the factor e which might have been expected to have added precision to the correlations failed to do so except in the case of the uppermost layer of soil from 0-10 cm. In order that higher values for r might be obtained the empirical formula Rw — e was used. It was felt desirable to retain the factor e since it represents a continuous process which results in loss of soil moisture. It was also desirable to weight the value R since rain falling seven days before a soil moisture deter- mination would have much less effect upon the soil moisture level than rain falling 92 Gardens’ Bulletin, Singapore — X XVI (1972) in the day before the measurement. Rainfall was weighted with the following values, corresponding to 1, 2, 3... 7 days before measurement; 1.0, 0.93, 0.85, 0.74, 0.60, 0.40, 0.10 which approximate to an exponential curve. The values for R weighted —e were then summed for varying periods prior to each moisture determination and separately correlated with soil moisture and rate of change of soil moisture. Selected results, giving highest values for r, are given in Tables 6 and 7. Table 6. Selected Correlations of Soil Moisture with R weighted—e, over Various Days before Moisture Determination Degrees of Significance Soil Days freedom r (better than) 1, (0-10 cm) : ee 19 + 0.46 5% 1, (10-20 cm) S. «ett 19 + (0.67 1s 1, (30-45 cm) | 19 + 0.66 1% 1, (60-75 cm) Dieneue 19 +0.60 1% 2, (0-10 cm) SS¢ 1 19 + 0.67 hy 3 2, (10-20 cm) ys l 19 +0.57 1% 2, (30-45 cm) fF 3 2 19 +0.57 1% 2, (60-75 cm) (lke = 19 + 0.62 1% 1 & 2, (0-10 cm) a 37 + 0.56 = 1 & 2, (10-20 cm) a as 38 +0.59 1% 1 & 2, (30-45 cm) ( pre 38 +0.59 1% 1 & 2, (60-75 cm) A eucasd 38 +0.57 1% Table 7. Selected Correlations of Rate of Soil Moisture Change with R weighted — e over Various Days before Moisture Determination Degrees of . Significance Soil ae freedom : (better than) 1, (0-10 cm) ie l 18 + 0.65 1% 1, (10-20 cm) aie | 18 +0.61 1% 1, (30-45 cm) Ry 2 18 + 0.63 1% 1, (60-75 cm) yee 2 18 + 0.40 None 2, (0-10 cm) Ds | 18 + 0.67 1% 2, (10-20 cm) a l 19 +-0.57 hy 4 2, (30-45 cm) aa 2. 19 +0.57 1% 2, (60-75 cm) Pi 2 19 +0.62 1% 1 & 2, (0-10 cm) ~ Sy l i +0.56 r% 1 & 2, (10-20 cm) Deak l 38 +0.59 0.1% 1 & 2, (30-45 cm) 7? 3 2 38 +0.59 0.1% 1 & 2, (60-75 cm) mu, 2 38 + 0.57 0.1% Soil Moisture under Forest, BT Nature Reserve, S’ pore 93 These data suggest that while correlation with this empirical formula gives slightly higher values for r than correlations with number of rain days, especially for the upper layer of the soil, the introduction of the factor e is hardly necessary. The weighting of R would also seem to be barely worth the labour of computa- tion involved. The conclusion is therefore, that with the exception of the upper layer of the soil where the introduction of e is desirable, the number of rain days in the period (here a week) before the measurement of soil moisture provides a simple parameter from amongst the meteorological factors affecting soil moisture. Since the method of determining soil moisture is by no means highly accurate, the coupling with it of such a ‘rough-and-ready’ measure as raindays seems justifiable, whereas the use of more sophisticated meteorological parameters is scarcely worth the labour involved. Acknowledgments I am grateful to the Director of the Botanic Gardens (Chairman, Nature Reserves Board), Singapore for permission to use the study area and for providing rainfall data. Evaporation data were kindly supplied by the Director, Singapore Meteorological Service. This paper is one of a projected series of studies of forest ecology in part of the Bukit Timah Nature Reserve. It is my pleasure to acknowledge fruitful discussions and the assistance of Mr. D. H. Murphy, Department of Zoology, University of Singapore who also commented upon a draft of this paper. Literature American Society for Testing and Materials, 1963, Standard Method for Grain-size Analysis of Soils, D422-63, Washington D.C. BELGRAVE, W. N. C., 1930, ‘The effect of cover crops on soil moisture’, Malay. agric. J., 18, 492-496. LeaMy, M. L. and Panton, W. P., 1966, Soil Survey Manual for Malayan Condi- tions, Ministry of Agriculture and Cooperatives, Bull. 119, Kuala Lumpur. Watson, G. A., Wong Phui Weng and Narayanan, R., 1964, ‘Effects of cover plants on soil nutrient status and on growth of Hevea V. Loss of nitrate — nitrogen and of cations under bare soil conditions. A progress report on results from a small-scale trial’, J. Rubb. Res. Inst. Malaya, 18, 161-174. le Ux WO: abi rie > inthcitipeh- Abid . / A90} ri pd bo pe ky te fom: SK . Jo iuade! dettrtinereyiduathachagis t eomoncs ah | ‘30 galeceniiell oan mo ft<8i wouloacs ret’ Tish Todt tr ei ube unk oat dort ficerod rhe Terry uf i} mh Chae 0 2 por poe ynd9 yolaroatnen sel) egos mont TOHoTLayae pal ge i green fie geal jciveratobe }: bea dita aan 3 ‘TEL | Me STASE rk A slot den, te, dig sath ' aioe pei jo smu odt aay nal osvibvai > nogal rls Sere are th sat ra siapeaghsl wugdo A r)) ansiyini) 2 —_ af) lortoienid 4 yute ofl oau 7 GdiapirNIaR, ior S19 = rl beilaque yvibatt saa Righ notimoqgevs ru fle bis ao ‘a soiree kl th Horo 4, ~~ reqorny we het o ehbuin ip soieie haiesioig « ‘Yo ont a e | luotinal4 wots i ciuassi veo ef OE eee utgktedeentt ae . ay be ate Pex SOCIO | Lats / ijqms?4 as 4 ears 4 | W.. one | i iM : De is Sorry —_ i : ’ ; os Pt 109 Parti be yt > 6 gneve. ind H jo diwmg ao ban eM usa, 1abi tH an f { o_ " a - ¢ 114 el Mii * e ¢ — — J* ». oi ‘m4 Algae from Gunong Jerai (Kedah Peak), Malaysia by M. RATNASABAPATHY School of Biological Sciences, University of Malaya, Kuala Lumpur Abstract The author’s collection of algae on Gunong Jerai is listed here with brief notes on habitat and existing records in The Malay Peninsula. Structural features of several species are also described. The total collection consists of 32 numbers, the 102 species and varieties are listed under 57 genera. Introduction Gunong Jerai or Kedah Peak is a mountain largely composed of sandstone, quartzite, quartz and some haematite (Robinson and Boden Kloss, 1916), and acidic intrusive igneous rocks (Alexander 1962). It rises to nearly 1219 metres (4000 feet) above mean sea level and is a prominent landmark of Kedah state. The vast flat lowlands more or less surrounding the foothills are mainly planted with rice. The sea to the west is only about 8 km (5 miles) away and in the mornings or when the moisture- and possibly salt-laden westerly winds blow, one could notice a thick mist moving across the peak towards the east. Ridley (1916), amongst other workers, has suggested that Gunong Jerai was once an island separated from the Malay Peninsula as in the case of Mt. Ophir, Johore and showed that there is much resemblance between the floras of these two mountains. The physical isolation of Gunong Jerai from the main range of the Peninsula and the absence of neighbouring mountains of any comparable altitude is phyto- geographically significant, and the lack of a true Thai element in the flora (see Ridley, 1916) is noteworthy. Though the angiosperms, gymnosperms and especially ferns have been abundantly collected and well documented, no data on algae have been published. Material and Method This paper contains the results based on the study of 32 numbers of collections. The algae were collected by the writer at altitudes of about 823-1189 metres (2700-3900 feet) above mean sea level between May 12 and 20, 1969 (see figure 1). The two main sites of collection were (1) a stream near the Government bungalow and (2) the only concrete reservoir situated on the left side of Peak Road one-third the distance from bungalow to peak. On a clear day, temperature readings of the stream between 10.30 a.m. and 12.30 p.m. were found to vary between 21.8 and 22.2°C and pH from 8.0 to 8.3: at the reservoir the pH was 6.2 at 22.5°C. The latter were determined in the field with a Lovibond comparator. Specimens were first examined fresh, then preserved dry or in solutions of formol acetic alcohol (Smith, 1950, p.34) or 5% formaldehyde. Of the nineteen species of diatoms collected eighteen were obtained as empty frustules from muds, bottom sediments and detrital material. The author consulted Johansen’s ‘‘Botanical microtechnique” for staining methods and simple microchemical tests. 95 96 Gardens’ Bulletin, Singapore — X XVI (1972) Species determination is incomplete due to inadequate reference material and absence of reproductive stages in some samples. For the nomenclature and sequence of major taxa (Divisions and Families), the system of Silva (1962) is largely adopted and to a lesser extent those of Papenfuss (1955) and Smith (1950). Genera are arranged according to systems of the following authors: Desikachary (1959, Cyanophyta), Prowse (1959, Bacillariophyta), Huber-Pestolozzi (1955, Euglenophyta), Randhawa (1959, Zygnemataceae), Scott and Prescott (1961, Desmidiaceae). Numbers in the list refer to the author’s collection. A complete set (101-133) is deposited at the Botany Division, School of Biological Sciences, University of Malaya, Kuala Lumpur; and 102, 104, 106, 107, 109-130, 132 and 133 are also at the Tropical Fish Culture Research Institute, Malacca. List of Species of Algae CYANOPHYTA CHROOCOCCACEAE Chroococcus turgidus Nag. Nos 100 & 111. Collected from washings of Sphagnum and sedge growing in reservoir and of moss scraped from submerged concrete sides of reservoir. Chroococcus ? minutus (Kiitz.) Nag. No 109. Collected from bottom muds, reservoir. Gloeocapsa sanguinea (Ag.) Kitz. Nos 112, 113, 115. Collected from surface of moist rock. Gloeotheca ? samoensis Wille No 109. Colonies of 2 to 8 ovoid cells with rounded ends, daughter colonies with individual sheaths, embedded in clear mucilage of parent envelope. From bottom muds of reservoir. Synechococcus ? aeruginosus Kitz. No 114. Planktonic in reservoir and stream. Dactylococcopsis acicularis Lemm. Nos 109, 110. Collected from submerged moss-covered sides of reservoir. OSCILLATORIACEAE Phormidium inundatum Kitz. ex Gomont Nos 109, 116, 117. Bright blue-green sheets; hormogonia abundant. From surface of concrete roadside drain with water trickling over it. NOSTOCACEAE Anabaena sp. Nos 116, 118. Filaments short, some up to about 500 » long; heterocysts rectangular, about 14 times longer than vegetative cells, intercalary, with two polar nodules; akinetes absent. Planktonic in pool, roadside drain. Algae from Gunong Jerai 97 SCY TONEMATACEAE Scytonema sp. Nos 115, 124. In mass appears as black somewhat shiny patches on wet or dripping rock; abundant. Tolypothrix sp. No 128. On submerged rock or mixed in with other algae and vegetation in stream. STIGONEMATACEAE Hapalosiphon hibernicus W. et G.S. West Nos 124, 125. Main axes usually uniseriate, occasionally biseriate, cells shorter or a little longer than broad, sheaths yellowish; branches uniseriate, mostly unilateral, sheaths non-lamellated, colourless, cells 2-6 times longer than broad, apex of terminal cells broadly rounded. Collected from torrential stream amongst other algae. Stigonema ocellatum (Dillw.) Thuret No 131. Scraped from surface of rock in stream. Reported from Templer Park (Hirano, 1967). Stigonema panniforme (Ag.) Bornet et Flahault No 129. Collected from torrential stream. RHODOPHYTA BATRACHOSPERMACEAE Batrachospermum ? moniliforme Ralfs Nos 101-107, 108, 124. — Plates 1, 2A-B Thalli macroscopic, 2-10 (-20) cm, moniliform, dark or pale blue-green or purplish brown, mucilaginous, corticated, freely branched. Nodes prominent. Internodes clear; corticating cells 2-6 times longer than broad, with short or long uniseriate filaments at the base; internodal filaments bear globose to obovate antheridial cells. Nodal filaments longer, branched; terminal cell ends in a long hair with dilated base, when fertile may bear clusters of antheridial cells at distal end. Cells of nodal branch citriform to oval, distinctly moniliform; apical cell dome shaped, obscured by upgrowing branches; branches may develop from the fourth axial cell and on the sixth, branches save as many as eight cells; young carpogonia clavate, stalked, develop at end of short but many-celled nodal branch; cystocarp rounded, more deeply pigmented, 1-3, more or less at periphery of nodal branch. The plants collected showed considerable variation in such features as colour, size, length of internodes, degree of branching, structure of cortical and other cells, and, it is possible that two forms or may be even species are involved here. “Chantransia’’-like thalli were not uncommon; these were bluish green, uniseriate, branched, the cells many times longer than broad, without terminal hairs, and vegetative. Some of these thalli resemble Audouinella. Found attached to rocks or gravel in stream; few specimens found on wet soil of stream bank where seepage water overflowed, very few small plants in the reservoir. Distribution patchy, sometimes gregarious. 98 Gardens Bulletin, Singapore — XXVI (1972) PYRRHOPHYTA — PHYTODINIACEAE Dinococcus bicornis (Woloszynska) Fott Nos 123-125 — Plates 2C, 3A-B Cells solitary, dorsal margin in front view convex, sometimes slightly depressed in the middle, showing at each end a short pointed spine; ventral view somewhat angular; vertical view ellipsoid; end view ovate. Stalk short, stout, colourless. Chromatophores obscured by dense accumulation of food reserve including a more or less central area of oil. Epiphytic on Oedogonium, other filamentous algae, and colonial diatoms e.g. Eunotia in stream; frequently gregarious. BACILLARIOPHYTA EUNOTIACEAE Eunotia ? camelus Ehrenb. Nos 122, 124. Collected from stream muds. Eunotia diodon Ehrenb. var. minor Grun. Nos 122, 124. In stream muds and epiphytic on other algae. Recorded in Cameron Highlands and Malacca River (Prowse, 1962a). Eunotia lunaris (Ehrenb.) Grun. No 123. In stream muds and epiphytic on other algae. Recorded in Cameron Highlands (Prowse, 1962a). Eunotia lunaris (Ehrenb.) Grun. var. capitata Grun. No 122. Collected from stream muds. Recorded in Malacca River (Prowse, 1962a). Eunotia monodon Ehrenb. var. tropica Hustedt Nos 122, 123. Among bryophytes and other vegetation in stream. Three varieties were recorded from stream, river and fish pond elsewhere in West Malaysia (Prowse, 1962a). Eunotia pectinalis (Kiitz.) Rabenh. var ventralis (Ehrenb.) Hustedt No 120. Collected from stream muds. The species was recorded at Cameron Highlands (Prowse, 1962a). Var. minus (Kiitz.) Rabenhorst has been recorded from Perak (Patrick, 1936). Eunotia ? pseudoflexuosa Hustedt No 123. Collected from stream muds. Eunotia robusta Ralfs Nos 118, 120. From roadside pool with flowing water and stream sediments. Recorded in Malacca river (Prowse, 1962a). Desmogonium rabenhorstianum Grun. Nos 120, 122, 123. Epiphytic on other algae, bryophytes and in sediments of stream. Prowse (1962a) reported this species from various habitats elsewhere in Malaysia, including swamps and coastal streams, at Cameron Highlands in West Malaysia. Algae from Gunong Jerai 99 ACHNANTHACEAE Achnanthes sp. Nos 118, 123. Collected from roadside pool with flowing water; epiphytic on liverwort in reservoir. NAVICULACEAE Frustulia rhomboides (Ehrenb.) De Toni Nos 106, 115, 120-123, 126. Distributed widely and abundantly in various habitats; its common occurrence suggests ecological significance and further studies are warranted. Recorded from most states of West Malaysia, including Cameron Highlands and Fraser’s Hill (Prowse, 1962a). Very abundant in algal samples collected by the writer in July 1970 at Tasek Bera, Pahang. Navicula spp. Nos 104, 118, 120. Many small species, especially common in bottom muds of streams and roadside pools. Pinnularia biceps Gregory Nos 112, 113. Collected from bottom sediments and submerged concrete sides of reservoir. Recorded from padi-fields, Beranang (Johnson, 1970). Var minor (Boye Petersen) A. Cleve has been recorded in Malacca river and fish ponds (Prowse, 1962a). Pinnularia legumen Ehrenb. No 111. Collected from stream and reservoir sediments. Recorded from Perak (Patrick, 1936); Taiping lakes (Prowse, 1962a) and from Templer Park (Hirano, 1967). Pinnularia maior (Kiitz.) Cleve Nos 122, 123. Collected from stream muds and pools. Reported from Perak (Patrick, 1936). Var. linearis was recorded in Templer Park (Hirano, 1967). CYMBELLACEAE Cymbella ? ventricosa Kitz. Nos 112, 113. Collected from bottom sediments of reservoir. Recorded at Cameron Highlands (Prowse, 1962a). NITZSCHIACEAE Hantzschia amphioxys (Ehrenb.) Grun. ? var. capitata O. Muller No 126. Collected from stream mud. Recorded from stream, Klang Gates (Prowse, 1962a). 100 Gardens’ Bulletin, Singapore — XXVI (1972) SURIRELLACEAE Surirella linearis W. Smith Nos 123, 126. Collected from stream sediments. Reported from Perak (Patrick, 1936); from fish ponds at Penang and Malacca river (Prowse, 1962a). Surirella spiralis Kutz. Nos 101, 114, 120. Collected from stream sediments. Rare. Reported from various habitats in Malacca (Prowse, 1962a). The writer has collected the species at various times during 1968 to 1970 in the Gombak river, Selangor and in July, 1970 at Tasek Bera, Pahang. Surirella ? tenuissima Hustedt Nos 123-125. From sediments on bed-rock and liverwort in stream. Recorded in Cameron Highlands (Prowse 1962a). CHRYSOPHYTA MALLOMONADACEAE Several species of Mallomonas were encountered in the stream sediments, reservoir and amongst other algae and bryophytes. Cells were mostly empty and spines usually lacking. The following species appear to be present: Mallomonas ? elliptica (Kisselew) Conrad Nos 110, 111. Mallomonas producta (Zacharias) Iwanoff Nos 112, 113. Mallomonas ? teilingioides Prowse Nos 112, 113. DINOBRYACEAE Dinobryon sertularia Ehrenb. No 111. Mostly only empty envelopes were found; solitary or in colonies of up to six cells. Colonies few-celled epiphytic on filamentous algae, rare in stream; mostly empty envelopes, common in sediments and waters of reservoir. Recorded from many acid swamps and fish ponds in many states of West Malaysia (Prowse, 1962b). RHIZOCHRYSIDACEAE Lagynion scherffeltii Pascher Nos. 108, 109. Epiphytic on filamentous algae in stream, sometimes gregarious, fairly common. Lagynion ? macrotrachelum (Stokes) Pascher No 124. Epiphytic on filamentous algae in stream. XANTHOPHYTA STIPITOCOCCACEAE Stipitococcus crassistipitatus Prescott Nos 122, 123. Epiphytic on filamentous algae in stream. Algae from Gunong Jerai 101 CHARACIOPSIDACEAE Characiopsis sp. No 128. Epiphytic on liverwort in stream. TRIBONEMATACEAE Tribonema sp. No 124. Cells 14-3 times as long as broad, slightly swollen in middle and narrowing at cross walls, ‘H’-pieces distinct but rare, chloroplasts many. This species is related to T. viride Pascher. Tychoplanktonic and in bottom muds of stream; often entangled with other filamentous algae. Neonema sp. No 124. Similar in habitat to that of Tribonema; rare. EUGLENOPHYTA EUGLENACEAE Euglena 2? mutabilis Schmitz No 120. Collected from stream muds. Recorded in Kedah padi fields (Sands, 1934), in carp ponds and stagnant water elsewhere in Malaysia (Prowse, 1957b, 1958) and in Templer Park (Hirano, 1967). The writer has collected it several times between 1968 and 1970 in the Gombak river, Selangor. Var. minuta Fritsch has been reported from many stagnant waters in West Malaysia (Prowse, 1958). Trachelomonas volvocinopsis Swirenko No 120. Collected from stream muds. Recorded from standing water and fish-ponds in various states of West Malaysia (Prowse, 1962b). Lepocinclis ? marsonni Lemm. emend Conrad No 120. Collected from stream muds. Common. Recorded from Taiping Lakes (Prowse and Ratnasabapathy, 1970). CHARACIACEAE Characium sp. Nos 128, 129. Cells solitary, fusiform, straight or slightly arcuate, poles acute and produced as long fine spines, each spine about half or more cell length; one spine terminating in a small rounded colourless attachment disc; chloroplast single, parietal, almost filling cell. The specimens were growing attached to the leaves of aquatic liverworts in a torrential stream. This species of Characium closely resembles Schroederia setigera, which however is reported to be always planktonic (see Smith, 1950). The presence of an eyespot and the disc-like ending of the spine in some species of Schroederia was reported. These species may well be attached forms that have become subsequently detached by accident and incorporated into the plankton. Further work on the ecology and taxonomic status of Schroederia with its described species is needed. 102 Gardens’ Bulletin, Singapore — X XVI (1972) ULOTRICHACEAE Stichococcus ? scopulinus Hazen Nos 128, 131. Filaments uniseriate, short, 1 to 4 or rarely 5 cells; cells cylindrical, sometimes up to 9 times longer than broad but usually shorter, ends rounded; chloroplast single, parietal, laminate, encircling about half the cell and usually not extending to the ends, apple-green coloured; pyrenoids lacking. Epiphytic on leaves of aquatic bryophyte in torrential stream; tends to be gregarious. Ulothrix sp. Nos 120, 132. Collected from a small pool of Government bungalow; in slow-flowing stretches of stream near bungalow. Rare. TRENTEPOHLIACEAE Trentepohlia iolithus (L.) Wallr. Nos 114, 133. Orange coloured, forming felted patches; filaments much-branched, unilateral at times, cells usually 2-5 times longer than broad, length up to about 90 42, chloroplast indistinct, reticulate; haematochrome abundant; cell wall fairly thick, lamellated in older cells; sporangia borne terminally or laterally on short or many-celled branch, usually single; spherical, diameter up to 25 yp, stalk cell inflated at base and bent distally. Epilithic, aerial. OEDOGONIACEAE Bulbochaete sp. Nos 111, 120. Thallus mostly uniseriately branched, cells 3-7 times as long as broad, length of cells 25-45 »; cell apex slightly inflated, with laterally placed seta; setae swollen at the base and very long, about 250 ». No reproductive stages found. Attached to other vegetation and twigs in stream and reservoir. Oedogonium sp. Nos 112, 113, 120. Only vegetative filaments were encountered, and probably more than one species are involved. Attached to aquatic vegetation, twigs, etc. in reservoir and in streams. ZYGNEMATACEAE Mougeotia ? elegantula Wittr. No 120. Collected from quieter stretches and pools of streams. Common. Several specimens resembling Mougeotia and Debarya (No. 115-119) could not be assigned accurately as only the vegetative stages were encountered. Zygnema sterile Transeau No 118. Cells up to two times longer than broad; cell wall thick, lamellated; chloroplast two, more or less stellate, each with one pyrenoid. Collected from stream, roadside pools and drains, often mixed in with other filamentous algae. Common. Several unidentified species of Zygnema or Zygnemopsis (Nos 115, 117, 120) were collected from the stream; some of these had cells many times as long as broad; reproductive stages were absent. Algae from Gunong Jerai 103 Spirogyra spp. Nos 120, 127. At last 3 species of Spirogyra were collected from the streams and one species from the reservoir; they had one to four chloroplasts. One of these (No. 127) was found in abundance in almost a pure stand in small cracks of stream bed-rock with static or gently flowing water, showing recently completed conjugation and young zygospores. Specific determination was not possible because the sculpture was not clear. Sirogonium sp. No 120. A species with four chloroplasts, each making half to three quarters of a turn; all filaments were in vegetative condition. Collected from quieter edges of stream. MESOTAENIACEAE Mesotaenium caldariorum (Lagerh.) Hansgirg No 133. Cells cylindrical, ends rounded, sides slightly to more or less parallel, length up to two times longer than broad; chloroplast single, axile, laminate, with or without a pyrenoid; mucilage surrounding cells clear, enveloping several cells. Collected from moist rock of roadside associated with Trentepohlia. Cylindrocystis brebissonii Menegh. No 115. Amongst Stigonema, Scytonema and Cosmarium on moist rock. The writer has collected the species in April 1970 in Tasek Bera, Pahang. Netrium digitus (Ehrenb.) Itzigs. et Rothe Nos 120, 122, 124 — Plate 3C Collected from slow-flowing stretches of stream. Common. Recorded in Taiping Lakes (Prowse and Ratnasabapathy, 1970). DESMIDIACEAE Actinotaenium cucurbita (Bréb.) Teil. No 118, 119. Collected from pool with flowing water, dripping wall of small iron storage tank, roadside of hill. Actinotaenium cucurbitinum (Biss.) Teil. No 119. Collected from dripping wall of small iron storage tank, roadside of hill. Actinotaenium elongatum (Racib.) Teil. No 119. Collected from dripping wall of small iron storage tank, roadside of hill. Actinotaenium grande (Delp.) Teil. No 112. Collected from small impoundment to reservoir. Actinotaenium subglobosum (Nordst.) Teil. No 119. Collected from dripping wall of small iron storage tank, roadside of hill. Actinotaenium truncatum (Bréb.) Teil. Nos 112, 118. Collected from pool with flowing water, roadside of hill and small impound- ment to reservoir. Actinotaenium wollei (Gronbl.) Teil. No 119. Collected from dripping side of small iron storage tank, roadside of hill. Closterium cynthia de Not Nos 112, 113, 122. Collected from bottom muds, reservoir. Recorded in pond, Ampang (Biswas, 1929). 104 Gardens’ Bulletin, Singapore — XXVI (1972) Closterium libellula Focke var. intermedium (Roy et Biss.) G. S. West Nos 111, 112 — Plate 3D Collected from bottom muds of reservoir. The species was recorded from the Taiping Lakes (Prowse and Ratnasaba- pathy, 1970). Closterium libellula Focke var. interruptum (West et West) Donat Nos 120, 126. Collected from stream mud. Closterium parvulum Nag. No 120. Collected from stream muds. Reported from Johore (Bernard, 1909); Setapak Lake (Biswas, 1929); padi- fields, Beranang (Johnson, 1970); elsewhere in Malaysia (Prowse, 1957a). Closterium navicula (Bréb.) Liitkem. No 115. Scraped from moist rock, grows mixed in with Cosmarium spp., Cylindro- cystis and Cyanophyceae. Recorded from Taiping Lakes (Prowse and Ratnasa- bapathy, 1970). Closterium striolatum Ehbrenb. Nos 109-111. Collected from muds and vegetation in reservoir. Recorded in Templer Park (Hirano, 1967). Closterium tumidum Johnson Nos 104, 123. Collected from muds overlying bottom rock of stream. Recorded in pond, Setapak (Biswas, 1929). Species of Closterium were found in practically all the samples of streams, rock pools, roadside drains and the reservoir. As an adventitious element in the plankton of both flowing and still waters they were not conspicuous. Multiplication by transverse division of cells was evident and appears to be the general method of maintaining the populations. Zygospore formation was lacking. The wide- spread occurrence, the ability to colonize a variety of ecological niches and fair abundance suggest ecological and economic importance. Pleurotaenium minutum (Ralfs) Delp. var. minus (Racib.) Krieg. Nos 104, 119, 125: From rock scraping and mud of stream; dripping side of small iron storage tank, roadside of hill. Var. excavatum was recorded from Taiping Lakes (Prowse, and Ratnasa- bapathy, 1970). Pleurotaenium trabecula (Ehrenb.) Nag. No 120, 122-124. Collected from stream muds, mixed in with other algae and occasionally in plankton. Fairly common. Tetmemorus laevis (Kiitz.) Ralfs Nos 120, 124, 126 — Plate 4C Cells up to 130 » long and eight times the breadth, sinus open and con- spicuous; semi cell slightly inflated just above base, apex deeply incised with the lobes somewhat rounded and asymmetrical; cell wall finely punctate, four punctae radially arranged at base of apical incision; chloroplast ridged with about six pyrenoids arranged longitudinally. Collected from stream muds. Fairly common. Recorded elsewhere in Malaysia (Prowse, 1957a). Algae from Gunong Jerai 105 Euastrum binale (Turp.) Ehrenb. var. brevius (Bernard) Hirano No 125. A small species fairly frequently found in waters of streams. Reported from Templer Park (Hirano, 1967). Euastrum dubium Wolle No 113. Collected from small impoundment to reservoir. Euastrum sinuosum Lenorm. var. dideltoides Krieg. Nos 109-111 — Plate 4A A larger species, up to 75 p long. | Amongst vegetation and bottom muds, reservoir. The species was recorded from Taiping Lakes (Prowse and Ratnasabapathy, 1970). Euastrum gnathophorum W. et G.S. West Nos 106, 107, 120, 124. Collected from stream muds. Recorded from Taiping Lakes (Prowse and Ratnasabapathy, 1970). Euastrum turgidum Wolle No 119. Collected from dripping side of small iron storage tank, roadside of hill. Cosmarium contractum Kirchn. Nos 120, 124. Collected from stream. Var. Jacobsenii was recorded in a pond, Ampang (Biswas, 1929). Cosmarium decoratum W. et G.S. West No 18. Collected from stream muds. Cosmarium hians Borge forma No 113. Collected from small impoundment to reservoir. Cosmarium lunatum Turn. Nos 111, 118, 120. Collected from pool with flowing water, hill roadside; bottom muds, stream; bottom muds, reservoir. Common. Cosmarium quadrifarium Lund. No 118. Collected from bottom muds, reservoir. Cosmarium taxichondrum Lund. No 118. Collected from pool, roadside drain with flowing waters. Rare. Cosmarium sp. No 106 — Plate 4B Several other species of Cosmarium specifically unidentifiable are not listed. Staurastrum sp. No 114. Planktonic in reservoir. Spondylosium pulchellum Arch. No 127. Short chains of cells epiphytic on aquatic vegetation in stream. Hyalotheca dissiliens (J.E. Smith) Bréb. Nos 107, 120, 124, 127 — Plate 4D Collected from stream, common in samples of other filamentous algae. Recorded in pond, Setapak (Biswas, 1929); from elsewhere in Malaysia (Prowse, 1957a). 106 Gardens’ Bulletin, Singapore — X XVI (1972) Analysis of Results Among the algae collected, 57 genera and 102 species including varieties were identified. About 25 species were attached forms, and 72 species occurred in the bottom muds or as tychoplankton. The true planktonic element appeared to be lacking and the few encountered such as Frustulia rhomboides and Hyalo- theca dissiliens were largely adventitious, stirred up from the bottom and suspended by the flowing waters. This was inferred from their abundant presence in bottom samples, apart from their features of habit and morphology. There was an abundant population and often a rich variety of species, especially desmids and diatoms, in the shallower, slow-flowing stretches of streams with aquatic phanero- phytes, bryophytes and decaying vegetation and where the canopy of trees overhead was less dense. The reservoir, which is a comparatively small habitat, being only about twenty metres square, had some 25 species, though few of these also occurred in the stream. The number of species in the Divisions of algae represented in these collections were as follows: Cyanophyta 13, Rhodophyta 1, Pyrrophyta 1, Bacillariophya 19, Chrysophyta 6, Xanthophyta 4, Euglenophyta 5, and Chlorophyta 51. The Cyanophyta, Bacillariophyta and Chlorophyta were also numerous in number of individuals and may be considered as dominant. Among the Chlorophyta, the Zygnematales formed about 80% of which the Desmidiaceae alone comprised two-thirds. Several of the species of algae are known to occur outside Malaysia, some have been recorded from elsewhere in Malaysia, and some are being recorded for the first time here. The occurrence of the temperate species Dinococcus bicornis Fott here extends its known range of geographical distribution. The presence of Dinobryon sertularia and species of the related flagellate Mallomonas in the reservoir may be undesirable with respect to the quality of a water source. Population outbursts of Dinobryon and Mallomonas have been reported to impart an obnoxious odour and a fishy taste to the waters of Hoover Reservoir, Ohio and Lake Michigan thus necessitating the treatment of the water supplies (Palmer, 1964). There are many problems concerning the freshwater algae of Gunong Jerai worthy of further investigation. Species of Closterium, Cosmarium, and Frustulia rhomboides were common and widely distributed; Batrachospermum, though rarer, showed an interesting sporadic occurrence with Audouinella-like thalli which may turn out to be the juvenile stages of the species. Studies on such aspects as their distribution, morphological variation, taxonomy, reproduction and life history should be rewarding. A comparative study of the Gunong Jerai algal flora with similar areas in other parts of Malaysia is not possible owing to the lack of published records. It may be mentioned, however, that about seven of the nineteen species of diatoms listed have been previously recorded at Cameron Highlands (see Prowse, 1962a). There is a wide and exciting field for future investigation of the Malaysian mountain algal floras. Acknowledgements I am indebted to Dr. Gerald Prowse, lately Director of The Tropical Fish Culture Research Institute in Malacca for advice and for checking identifications. I also wish to thank Dr. Ralph A. Lewin, Professor, Scripps Institution of Ocean- ography, University of California, La Jolla for critical comments on the manu- script and Dr. B. C. Stone, Reader at The School of Biological Sciences, University of Malaya, for introducing me to the background literature on Gunong Jerai. Plate 1. Batrachospermum ? moniliforme Ralfs A: Apex of thallus stained with Phloxine B B: Whorls of lateral branches of limited growth C: Cystocarps at periphery of whor! of lateral branches yy y & ] \ % Sys j YY ee 30 Plate 2. A: Batrachospermum ? moniliforme Ralfs, cystocarp terminal on short lateral branch, stained with Phloxine B B: Idem, spermatia on lateral branches of limited growth C: Dinococcus bicornis Fott, epiphytic on colonial Desmogonium, front view ) S0u > Dinococcus bicornis Fott, epiphytic on filamentous Xanthophyceae, front view Idem, end view Netrium digitus (Ehrenb.) Itzigs. et Rothe - Closterium libellula Focke var. intermedium (Roy et Biss.) G. S. West Euastrum sinuosum Lenorm. var. dideltoides Krieg. Cosmarium sp. Tetemorus laevis (Kiitz.) Ralfs Hyalotheca dissiliens (Smith) Bréb. stained with methylene blue (oil immersion) ad Iwuif SONONNS JO dV¥W L 38NSIS > 1S'LZ e001 . 18% 0008 1S'2Z 0H SAYSS3SY LSAYOS IVHar SNONND TN AANK . O67 A Nudd quawYyoie5 \ UBA | S'd70S < rs) re) < | < = we oO weisy jyods w N 0605 _ < Gosyno yr0d 3 HID = ~~ Tp) : a1 JaAly poo Amo Sam 6u1}59/109 yvaadN3H) uno Algae from Gunong Je 108 Alexander, J. B. 1962 Bernard, Ch. 1909 Biswas, K. 1929 Desikachary, T.V. 1959 Fott, "B. 1959 Fritsch, F. E. 1935 Gronblad, R., Prowse, G. A. and Scott, A.M 1959 Hirano, M. 1967 Huber-Pestalozzi, G. 1955 Johansen, D. A. 1940 Johnson, A. 1970 Migula, W. 1907 Palmer, C.M. 1964 Papenfuss, G.F. 1955 Gardens’ Bulletin, Singapore — XXVI (1972) Bibliography A brief summary of the geology of Malaya. Malayan Nat. J. le ¢ 30-35. Algues unicellulaires d’eau douce récoltées dans le dom- aine Malais. Dept. de TAgric. aux Indes Néerl., Buitenzorg. 44 pp., 6 plates. Papers on Malayan aquatic biology IV. Freshwater algae (with addendum). J. Fed. Malay States Mus. 14 (3-4) : 404-435, 479-481, plates 9-13. Cyanophyta. I.C.A.R. Monogr., New Delhi. Algenkunde. Veb Gustav Fischer Verlag, Jena. The structure and reproduction of the algae. Vol. 1. Camb. Univ. Press. Sudanese desmids. Acta Bot. Fenn. 58 : 1-82. Freshwater algae collected by the joint Thai-Japanese biological expedition to Southeast Asia 1961-2. Nature and Life in Southeast Asia. Vol. V, pp. 1-71, 16 plates. Das Phytoplankton des Susswassers. Teil 4. Euglenophy- ceen; E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart. Botanical microtechnique. McGraw Hill Book Co., Inc., N. Y. Blue-green algae in Malaysian rice-fields. J. Sing. natn. Acad. Sci. 1 (3) : 30-36 Kryptogamen-Flora von Deutschland, Deutsch-Osterreich und der Schweiz 6, Kryptogamen 2, Algen 1 : Cyanophy- ceae, Diatomaceae, Chlorophyceae. Text pp. 1-918, plates 1-43. Algae in water supplies of the United States. IN Algae and Man, ed. by Jackson, D. F. Plenum Press, N. Y., pp. 239-261. Classification of the algae. IN A century of progress in the natural sciences. Calif. Acad. Sci., San Francisco. | Algae from Gunong Jerai 109 Pascher, A. 1925 ————. 1937-39 Patrick, R. 1936 Patrick, R. and Reimer, C. W. 1966 Prescott, G. W. 1962 Prowse, G. A. 1957a ——. 1957b ———. 1958 ———. 1960 . 1962a . 1962b Prowse, G. A. and Ratnasabapathy, M. 1970 Randhawa, M. S. 1959 Ridley, H. N. 1916 Robinson, H. C. and Boden Kloss, C. 1916 Sands, W. N. 1934 Scott, A. M., Gronblad, R. and Croasdale, H. 1965 Die Susswasserflora Deutschlands, Osterreichs und der Schweiz. Vol. 11, pp. 1-118. Heterokonten. In L. Rabenhorst, Kryptogamen — Flora Deutschland, Osterreich und der Schweiz. Vol. 11, 1092 Pp. A taxonomic and distributional study of some diatoms from Siam and the Federated Malay States. Proc. Acad. Nat. Sci. Philad. 88 : 367-470. The diatoms of the United States. A. N. S. P. Monogr. 13, Vol. I. Philad. Algae of the Great Lakes area. Cranbrook Inst. Sci. Bull. 31 : xiii + 1-977. An introduction to the desmids of Malaya. Malayan Nat. J. 11 (3): 42-58. Fish and food chains. Malayan Nat. J. 12 (2) : 66-71. The Eugleninae of Malaya. Gard. Bull. Sing. 16 136-204. Vegetation changes in experimental ponds. In Proc. Centenary and Bicentenary Congr. Biol. Sing. pp. 288-291. Diatoms of Malayan freshwaters. Gard. Bull. Sing. 19 (1) : 1-104. Further Malayan freshwater Flagellata. Gard. Bull. Sing. 19 (1) : 105-145. A species list of freshwater algae from the Taiping Lakes, Perak. Gard. Bull. Sing. 25 (2) : 179-187. Zygnemaceae. 1. C. A. R. Monogr., New Delhi. The natural history of Kedah Peak V. Botany. J. Fed. Malay States Mus. 7 (2) : 39-58. The natural history of Kedah Peak I. Introduction. J. Fed. Malay States Mus. 6 (4) : 253-256. The coloured scums of padi fields. Malay. Agri. J. 22 (10) : 484. Desmids from the Amazon Basin, Brazil. Acta Bot. Fenn. 69 : 3-94. 110 Gardens’ Bulletin, Singapore — XXVI (1972) Scott, A. M. and Indonesian desmids. Hydrobiologia 17 (1-2) : 1-132, with Prescott, G. W. 63 plates. 1961 Silva, P. C. 1962 Classification of algae. In Physiology and biochemistry of algae. Edited by Ralph A. Lewin. Academic Press, N. Y. Skuja, H. 1949 Zur Siisswasseralgen — Flora Burmas. Nova Acta R. Soc. Scient. upsal. Series IV, 14 (5) : 1-188, 37 plates. Uppsala. Smith, G. M. 1950 The freshwater algae of the United States. 2nd ed., N. Y. Teiling, E. 1954 Actinotaenium genus Desmidiacearum resuscitatum. Bot. Notiser. 4 : 376-426. Tiffany, L. H. and The algae of Illinois. Chicago. Britton, M. E. 1952 Wildeman, E. 1900 Flore de Buitenzorg III. Algues. Leiden. Review E. J. H. CORNER AND K. WATANABE, //lustrated Guide to Tropical Plants. Hirokawa, Tokyo, 1969. 1147 pp., 29 colour pl., c.2000 line drawings. The title of this large and ambitious book is deceptive: It is based on a collection of line drawings of phanerogams, made by Professor Watanabe during a 3-year stay in Singapore and Malaya; the species selected are therefore largely unrepresentative of the tropics as a whole, both on taxonomic, ecological and utilitarian grounds. Besides most of the commoner cultivated plants of Malaya, many of which are exotics and indeed widely grown in the tropics, they include a somewhat arbitrary selection of native species, weeds, and many plants existing in that region only as individuals in botanic gardens, of which some are by no means common in their countries of origin. The drawings are arranged by families according to Engler’s system; each is captioned by its family name. Beneath the illustration is given the scientific, Malayan (if known), Japanese, and sometimes Indian and English names; a short but useful descriptive note indicating distribution, habit, leaf arrangement and flower and fruit colour by Professor Corner; and a note on uses in Japanese. The usefulness of each plant is then summarised in a curious annotated point-grid system, which appears to have been intended as a ready-reference abbreviation but is in fact the very reverse, being difficult to follow and space-demanding; finally, the position in Hutchinson’s classification is indicated. Some 700 of the more than 2,000 line drawings were originally published in two volumes by the Japanese administration of the Singapore Botanic Gardens during the second world war, entitled “Illustrations of Useful Plants of the Southern Regions: Section 1: Medicinal,” and “Section II: Edible Plants.” These books are now rare, but many of the same plates were used again by Professor H. B. Gilliland for his book “Common Malaysian Plants” (University of Malaya Press, 1958). This book cannot be used for identifying plants, as the illustrations and descriptions make no attempt to be diagnostic, and no keys are given. Its purpose would appear to be a reference book for laymen in S.E. Asia, especially foreign agriculturists, foresters and businessmen who, on being shown a named plant to which certain useful properties have been ascribed, wish to check the veracity of this information. The book is'a monument to Professor Watanabe’s energy, but it is a pity that the illustrations, the originals of which must generally have been of high quality, have had to be reduced to a scale where the details, especially of floral morphology, have often been lost. P. S. ASHTON Institute of South-East Asia Botany University of Aberdeen 111 REVIEW TSENG-CHIENG HUANG. Pollen Flora of Taiwan National Taiwan University, Botany Dept. Press, Taipeh. 1972; pp. v+ 276, 177 photographic plates and index. Price U.S. $27.00 (postage and handling included). This is one of the very few local pollen floras that have appeared in the past decades. The author was probably inspired by M. Ikuse’s “‘Pollen Grains of Japan” published in 1956. Part one of this book is an introduction, including an illustrated account on pollen morphology. Part two constitutes the main body of the book — the systema- tic treatment. It contains keys to the families (divided into 25 classes) and descriptions of 197 families of seed plants (Gymnosperms-9, Dicotyledons-157, and Monocotyledons-31). The families are arranged alphabetically within these three taxa. Each family consists of a general description of the pollens and a bracketed key to the genera (if more than 2). The descriptions of genera and species are in general very concise. Numbers of voucher herbarium specimens are also cited. Two most valuable features of this book are probably the comprehensive and useful keys based on pollen characters for identifying families and genera, and the excellent quality of photographic plates. The latter is attributable to the facilities of fine equipment in the Palynological Laboratory at the National Taiwan University and the skilled technique in preparing slides and photographs. This book will be very useful to plant taxonomists and palaeobotanists and I have no hesitation in strongly recommending it. H. KENG Department of Botany University of Singapore 112 Athrodactylis and Pandanus: A brief comment on the ‘*Characteres Generum Plantarum’ of J. R. and G. Forster by BENJAMIN C. STONE School of Biological Sciences, University of Malaya, Kuala Lumpur Although it is well known that, in the work “Characteres Generum Plantarum” of 1776 by Johan Reinhold and Georg Forster, these authors boldly and improperly utilized information they had acquired through their relationship with Sir Joseph Banks in working on the materials which Banks and Solander had amassed during the first of Captain Cook’s voyages. in somewhat the same way that had been earlier done by the brother (Parkinson) of the expedition artist, it is interesting to discover that they did not confine their borrowings to any one source. While working briefly in the Herbarium of St. Xavier’s College in Bombay I was able to use originals of the Forsters’ work, as well as a number of other ancient and precious volumes. One of these was the ‘‘Horti Indici Malabarici” of _ Henricum van Reede tot Draakestein (published between 1678 and 1703). Studying in particular the references to Pandanus I then turned to the Forsters’ work, and was immediately struck by the resemblance virtually to the point of identity between the drawing of a fruit (phalange) in fig. 6 of Reede’s ‘‘pars secunda” with the name “‘Kaida Taddi” and the illustration (fig. 75) of the Forsters’ ““Athrodactylis spinosa” (i.e. Pandanus). The Forsters’ illustration (titled “‘Appendix Palmae’’) shows (a) one stamen, (4) one phalange, and (c) one seed. The last two are almost _ exact copies from Reede’s illustration of ““Kaida Taddi”’. This copying, for such I am sure it was, tallies logically with the Forsters’ _ descriptive text, where (p. 150) they give the diagnosis of ‘‘spinosa. 1. ATHRO- _ DACTYLIS.” in the Linnaean manner, with the synonyms “Bromelia Sylvestris. Lin. Flor. Zeyl. Burm. Ind. 79.; Keura Forskal Arab.; Kaida Rheed. Hort. Malab. _p. IL. t. 1-8. Pandanus Rumph.” This is somewhat confusing, but it seems to _ suggest that the Forsters considered all these names synonymous with their species _ “Athrodactylis spinosa,” although Reede’s figures show clearly (to the modern _ reader) about four distinct species of Pandanus. The Forsters’ list of synonyms _ beneath the ‘‘species” rather than beneath the “‘genus” diagnosis (although, strictly _ speaking, there is no separate botanical diagnosis of the species; only an explanation of the generic name, followed by the ‘“‘Explicatio” for Tab. LXXV.) seems to a modern reader a mixture of generic and specific synonyms. It is difficult to come to a conclusion as to whether the Forsters thought that all the synonyms they cite really referred to a single species or to a single genus. The only post-Linnaean - name cited is Keura Forskal. This was monotypic, and thus by citing it in synonymy _ we would expect the epithet of the species to be taken up if we apply the current - nomenclatural Code. The full name of Forskal’s plant was Keura odorifera. This _ name is based on material which Forskal purchased in a small village in south- western Arabia, where the staminate inflorescences were on sale as a perfume. 113 114 Gardens’ Bulletin, Singapore — X XVI (1972) To this day in both India and in adjacent regions the practice persists, and in 1968 in the villages on Madh Islet in Bombay (for example) one can still see these inflorescences for sale; children and women carry and wear them, “harvesting” them from the trees. The plants are as common in cultivation as wild, especially along the coast. Forskal’s genus Keura is certainly synonymous with Pandanus Stickm. of 1754; the publication of Forskal’s ‘Flora Aegypto-Arabica” was in 1775. The generic name Athrodactylis J. R. & G. Forst. is thus clearly a synonym since Forskal’s publication was cited. Unfortunately the type material of Keura no longer exists. It is, however, very likely to be the plant which is still used in the way mentioned — as perfume — along the coasts of India and is traded for such in adjacent areas. This is clearly identical with Pandanus odoratissimus L. f. (the type of which is from Ceylon). At any rate, it is clear that: (1) Athrodactylis, as a genus, is equivalent to Pandanus; (2) Athrodactylis as a name is not nomencla- turally usable, being published with a valid name in synonymy (i.e. Keura); ‘3) the epithet “‘spinosa”’ is also illegitimate since the epithet “‘odorifera’’ of Forskal should have been taken up; (4) except for the reference to Pandanus Rumph. the remaining synonyms all refer to Indian or Ceylon plants; thus it is plain that A. spinosa is none other than Pandanus odoratissimus Lf. This is borne out convincingly by the fact that to illustrate A. spinosa, the Forsters copied Reede tot Draakestein’s illustration of “‘Kaida Taddi” which is unquestionably P. odoratis- simus Lf. As a final point it is interesting to note that G. Forster in his later work on Pacific Island Plants (Prodr. 69. 1786) used the binomial Pandanus odoratissima (sic). This certainly means that he then considered Pandanus as the correct generic name, and by taking up Linnaeus filius’ specific epithet also (though altering it to feminine gender) he apparently also meant that the Tahitian plants which he was discussing were to be equated with the Indian and Ceylon species. This interpreta- tion can perhaps therefore be viewed as a confirmation of the conclusions given above. It is interesting to note further that although much of the ‘Characteres Generum Plantarum’ was based on Pacific Islands plant collections, the “‘Appendix’’ with Athrodactylis does not specifically mention any Pacific materials. The reference to Rumphius suggests Forsters’ belief that the species of India and Ceylon extended into the East Indies. But there is no mention of Pandanus-tectorius (sic) of Parkinson (attributed by many authors to Solander, but never published by him; | Parkinson wrote it as a monomial hence it is illegitimate) which appeared in 1773 | (Journ. Voy. South Seas ... H.M.S. ‘Endeavour’ p. 46). There is no mention of any collection by either of the Forsters, although certainly they must have seen many Pandanus plants while they were in Tahiti. BIBLIOGRAPHY Forster, J. R. and G. 1776. Characteres Generum Plantarum, Londoni. Reede tot Draakestein. Henricum van. 1678-1703. Horti Indici Malabarici. Cytogenetics and Taxonomy of the Genus Globba L. (Zingiberaceae) in Malaya IV Distribution in Relation to Polyploidy by Lim Srew-Nco* Genetics Division, School of Biological Sciences University of Malaya**, Kuala Lumpur, Malaysia Abstract Of the twelve species, five subspecies and eight varieties of Globba in Malaya, some are exclusively allotetraploid (2n = 32, AABB), some exclusively hexaploid (2n = 48, AAABBB), one decaploid (2n = 80) and some with both allotetraploid as well as hexaploid chromosome traces. Globba species with only 2n = 32 occur in lowland forests and at moderate elevations, while those with only 2n = 48 generally in montane forests. Both are of rather restricted and localised distribution when compared to taxa with 2n = 32, 48. In the latter, the hexaploid infraspecific taxa are usually but not strictly montane in distribution. Where both the tetraploid and hexaploid taxa are montane in distribution, the hexaploids are of greater abundance. The reverse is true where tetraploid taxa occur in lowland forests and hexaploid in montane regions. Globba species in Malaya thrive in moist, partially shaded niches. An exception is G. marantina L. which colonises open habitats and has a wide distribution from India and Philippines to the Solomon Islands. The genus Globba L. (Zingiberaceae) is confined in distribution to the eastern Himalayas and southern China southwards to Malaysia and the Solomon Islands (Holttum 1950, Pendleton 1949). Of the 128 species recorded (in Index Kewensis, compiled up to 1966), twelve species, five subspecies and eight varieties occur in Malaya. These include eight new taxa which are described in Lim (1972 a). Of these Malayan taxa, three species, two subspecies and five varieties are exclusively allotetraploid (2n = 32, AABB), three species and a subspecies exclusively hexaploid (2n = 48, AAABBB), one species decaploid (2n = 80) and three species and a subspecies with both allotetraploid as well as hexaploid chromosomes races. (see Table. 1; Lim 1972 b). This paper presents the results of distributional studies of these Malayan taxa with special reference to their chromosome numbers. Materials and Method Sources of information for distribution mapping were the 363 herbarium specimens on loan from the Botanic Gardens, Singapore, and personal field collection data. Cases in which identification was doubtful and handwritten data dubious were rejected. In the mapping of each taxon, a distinction was made between herbarium and field collections. Herbarium collections are represented by solid circles, semi-circles and squares and field collections by outlined symbols. A single dot represents one to three collections from the same station, an enlarged symbol four or more. A large number of collections need not necessarily * Now Dr (Mrs) Chong Siew-Ngo nee Lim ** Current address: Department of Biology, Nanyang University, Singapore. 115 116 Gardens’ Bulletin, Singapore — XXVI (1972) represent increased density of occurrence of the taxon, as it might well represent over-collection. Absence of dots might merely mean under-collecting. With the available information, the maps (figs. 1, 2 and 3) represent what is known of Globba distribution in Malaya to date. No attempt was made to distinguish between the polyploids in the herbarium specimens. Taxa with 2n = 48 were distinguished from those with 2n = 32 in the mapping of field collections. Root tips as well as pollen mother cells were investigated for chromosome counts. Details are as described in Lim 1972 b. Information on the sources and chromosome numbers of fresh specimens are in Table 2 on p. 124 Herbarium specimens of these are deposited in the Herbarium, Botany Division, School of Biological Science, University of Malaya, Kuala Lumpur. Table 1. Intraspecific polyploidy in the Malayan species of Globba. Species ange neyo Species with 2n = 32, 48 (a) Species with 2n = 32 on ="S7 an = 9 1 G. fragilis Lim 1 G. pendula Roxb. G. pendula Roxb. 2 G. albiflora Ridl. G. pendula var. — G. albiflora var. elegans (Ridl.) aurea Holtt. Holtt. 3 G leucantha var. G. pendula spp. —- peninsularis Holtt. montana (Ridl.) 4 G. unifolia Ridl. Lim. 2 G. patens Miq. G. patens Mig. (b) Species with 2n = 48 G. patens var. -—— 1 G. curtisii Holtt. costulata 2 G. holttumii Lim Lim G. holttumii ssp. aurea Lim 3 G. cernua Bak. G. cernua Bak. G. cernua ssp. G. cernua =s crocea Lim crocea Lim (c) Species with 2n = 80 G. cernua ssp. = 1 G. marantina L. porphyria Lim 4 — G. variabilis Ridl. (d) Species with unknown G. variabilis se chromosome number ssp. pusilla 1 G. fasciata Ridl. Lim Distribution The distribution of Globba species with two anther appendages is recorded in figure 1 while that of Globba species with four anther appendages is recorded in figures 2 and 3. Globba species with only 2n = 32 or 2n = 48 show a rather restricted distribution compared to those with 2n = 32, 48. G. fragilis Lim (2n = 32) appears to be confined to Pulau Langkawi, Kedah (see fig. 3), G. albiflora Ridl. to Penang, G. albiflora var. aurea Holtt. to Gua Lambok and Sungei Betis in Kelantan, and G. unifolia Ridl. with its subspecific taxon to Trengganu and Kelantan. G. unifolia also occurs in Thailand. Trengganu might be its southern limit of distribution. G. fasciata Ridl. of unknown chromosome number, also has a very restricted distribution (see fig. 1). Cytogenetics and Taxonomy of the Genus Globba L. 117 G. curtisii Holtt. (2n = 48) has so far been found only in a restricted part of the Main Range, between Bukit Kutu, the Gap and Fraser’s Hill (see fig. 3). Further investigation might show that its distribution extends to other parts of the Main Range. G. holttumii Lim grows in large clumps in restricted localities in the Gombak Forest Reserve, Selangor (c.2500’), Genting Simpah, Selangor (2700’), Genting Highlands, Selangor (c.4000’) and Fraser’s Hill, Pahang (c.4000”). Both G. curtisii and G. holttumii are markedly montane in their distribution. G. holttumii ssp. aurea Lim, however, was collected from lowland Dryobalanops Forest, Lenggor Forest Reserve, Johore (c.300’). Four of the twelve species of Globba in Malaya show 2n = 32 and 48 and these are relatively widespread in distribution. G. pendula Roxb., “the commonest Globba in Malaya, occurring in all parts of the country.” (Holttum, 1950) was collected from all along the west coast of Malaya from Pulau Langkawi in the North to Singapore in the South (see fig. 1). Distribution seems densest in Perak, Selangor, Negri Sembilan and Malacca. On the eastern side of the Main Range, collections are from the interior of Pahang and from Kelantan. None of the 118 herbarium collections came from Trengganu and coastal Pahang. One single collection came from Pulau Tioman, off the Pahang coast. The species has a predominantly lowland distribution. Field specimens with 2n = 48 are from Fraser’s Hill, Pahang (c.4000’). Chromo- some races of 2n = 32 occur alongside those with 2n = 48 by the waterfall, near the old dairy farm on Fraser’s Hill. Field collections of specimens solely with 2n = 32 were made along the west coast from the foot of Gunong Jerai, Kedah to around Kuala Lumpur. The chromosome race with 2n = 48 appears to be confined to montane areas. Where G. pendula Roxb. occurs, plants are found in abundance. This is also true of the subspecific taxa: G. pendula var. elegans (Ridl.) Holtt. in the Fraser’s Hill region of Pahang and G. pendula ssp. montana (Ridl.) Lim in Penang. Outside Malaya, the species extends to India, Sumatra, and the Banka Islands. Common on the west coast from Penang to Malacca, in both lowland and montane forest is G. patens Miq. (2n = 32, 48) a species hitherto misidentified as G. aurantiaca Mig. see Lim (1972 a). Chromosome races with 2n = 32 seem confined to lowland forest up to c.2000’, while those with 2n = 48 are from Frasers Hill, Pahang (c.4000’), Bujang Melaka, Perak (c.2200’), Genting Highlands, Pahang (c.4000’) and Bukit Lagong Forest Reserve, Selangor (c.200-800’). Excepting the last, G. patens with 2n = 48 appears mainly montane in distribution. Single herbarium collections have been made from Johore and Kelantan. A field collection of a narrow-leaved and nearly glabrous form was made from Lenggor Forest Reserve, Johore (c.200’). This appears similar to the form collected from Ulu Kahang (Holttum, S.F.N., 10927), and has 2n = 32. At Fraser’s Hill, Pahang (c.4000’), Cameron Highlands, Pahang (c.4750’), and Sungei Lallang Forest Reserve, Selangor (c.250’), G. patens var. costulata Lim was found. It resembles the specimens from Bukit Kutu in Selangor by Ridley (mentioned in Holttum, 1950, p.37). This variety (2n = 32) apparently occurs in both lowland and montane forests. Overlapping in distribution with G. patens Miq. is G. cernua Bak, which also consists of chromosomal races with 2n = 32 and 2n = 48. Globba cernua Bak., however, is confined to montane forests between 2000’ and 5000’, within the region between Temangor, Perak and Gunong Angsi, Negri Sembilan. South 118 Gardens’ Bulletin, Singapore — X XVI (1972) of this, one single collection was made from Singapore. Both chromosomal races occur in abundance in Fraser’s Hill, Pahang and Maxwell’s Hill, Perak. More common are plants with 2n = 48 than those with 2n = 32. On Maxwell’s Hill, the two chromosomal races occur together in mixed stands. In Cameron High- lands, only plants with 2n -- 48 have so far been found. At Reid’s Third Quartz Ridge, 12th mi, Gombak Road, Selangor all plants collected had 2n = 32 chromosomes. Fruiting specimens in the herbarium collections from Larut Hills and Maxwell’s Hill in Perak, and Gunong Angsi in Negri Sembilan suggest that plants with 2n = 32 are also present. Two new subspecific taxa, G. cernua ssp. crocea Lim (2n = 32, 48) and G. cernua ssp. porphyria Lim (2n = 32), were collected from Jenka Forest Reserve, Pahang (lowland Dipterocarp forest, c.200’) and Bujang Melaka, Perak (c.1370") respectively. The two chromosomal races of G. cernua ssp. crocea were found side by side in Jenka Forest Reserve. Abundant occurrence along the jungle path from Kuala Tahan to Kuala Trengganu in the National Park in Pahang is reported by G. Smith in 1968 (personal communication). G. variabilis Ridl. (2n = 48), shows a rather widespread distribution from Gunong Korbu (Perak) on the western side of the Main Range and Kuala Klah (Kelantan) on the eastern side of the Main Range to Singapore (see fig. 3). It occurs mainly in lowland forest but also in montane areas. All living specimens collected from four main localities: Fraser's Hill, Pahang (c.4000’), Ulu Gombak Forest Reserve, Selangor (c.2500’), Sungei Lallang Forest Reserve, Selangor (c.250’) and Bukit Timah Nature Reserve, Singapore (c.500’) had somatic chromosome numbers of 48. The relatively high occurrence of fruiting materials in 9.8 per cent of the 121 herbarium sheets in the Singapore Botanic Garden collection seems to suggest the possibility of the existence of a chromosome race with 2n = 32. The possibility of these having 2n = 48, however, could not be completely ruled out. A new subspecies, G. variabilis ssp. pusilla Lim (2n = 32) was collected from Gunong Panti (1500’), Sungei Kayu and Sungei Sedili in Johore. So far, the distribution appears limited to the southern tip of the Malay Peninsula. An apparent exception to the general finding (that species with 2n = 32 show a rather restricted distribution compared to those with 2n = 32 and 48) is G. leucantha Miq. Brief cytological investigations of G. leucantha var. peninsularis Holtt. from two localities in southern Johore disclosed somatic numbers of 32. Globba leucantha shows a widespread but discontinuous distribution in the north and extreme south of Malaya (see fig. 1). The relative paucity of both herbarium and field collections may be related to the seasonal dormancy of the species. Globba leucantha var. peninsularis occurs in southern Johore and Singapore, reportedly “common in forest” (Holttum 1950). A few collections were from southern Trengganu, northern Pahang and Perak and two field collections were from Gunong Panti, Johore and Bukit Timah Nature Reserve, Singapore. G. leucantha var. bicolor Holtt. appears to be confined to the eastern part of the Main Range, in northern Pahang, southern Kelantan and Trengganu. G. leucantha var. violacea (Ridl.) Holtt. has been collected from Perak; and G. leucantha var. flavidula (Ridl.) Holtt. from Gunong Panti, Johore (100’). Although a direct cytological investigation of G. leucantha var. bicolor and var. violacea has not been conducted, the rather high incidence of fruiting materials in the two varieties seems to suggest that they have somatic numbers of 32. 0 f= ei ae ie —— cl ss eta iterate — ee eet ——— res ¢ i (ine ee ‘PERLIS (Rs. ra Miles 0 10 #20 30 40 50 eee ee ee ee ee 6° 00" SOUTH CHINA SEA 4° 00' | BON g PULAU e LANGKAWI THAILAND tien Ha fs) Land between 3,000ft. and 5,000 ft. eee Land between 1,000 ft and 3,000 ft. Globba albiflora Ridl. Globba albiflora var. aurea 60) Globba fasciata Ridl. Globba leucantha var. peninsularis Holtt. (fresh Globba leucantha var. violaceae (Ridl.) Holtt. Globba leucantha var. bicolor Holtt. Globba leucantha var. flavidula (Ridl.) Holtt. Globba pendula Roxb.(> 4 herb. coll.) Globba pendula Roxb. (1-3 herb, coll.) Globba pendula Roxb. ( 2n=32, fresh coll.) Globba pendula Roxb. (2n=48, fresh coll.) pendula var. elegans (Ridl.) Holtt. Globba pendula var. montana (Ridl.) Lim Globba unifolia Ridl(herb. coll.) eo Globba unifolia Ridl. (fresh coll.) " : > SINGAPORE Globba unifolia var. sessiliflorg Holtt. - Fig. 1 Distribution map of Globba species with two anther appendages in Malaya. @QOe2®22°* @ae Ko * Miles OR 1020) 300740 50 _ aS ee ee SOUTH CHINA SEA Land: above 5,000 ft. Land between 3,000 ft. and 5,000 ft. Land between 1,000 ft. and 3,000 ft. cernua Bak. s.s.(>4 herb.coll.) I@ ex ioe cernua Bak. s.s.(1-3 herb. coll.) Pee NE SE | @. - Aes e Fs ole) cernua Bak. ss. (2n=32, fresh coll.) i@ 1o cernua Bak. s.s. (2n=48, fresh coll.) . cernua . porphyria Lim (2n= 32) . cernua . erocea Lim (2n=32) . cernua . erocea Lim (2n=48) . patens Mig. (>4 herb. coll.) i JOHORE ig. (1-3 herb. coll.) a iq. (2n= 32, fresh coll.) iq. (2n=48, fresh coll.) ite). _ costulata Lim (2n=32, fresh coll.) ¢ SINGAPORE 2008 @ ¥H+O -°*ol7 Ill Fig. 2 Distribution map of Globba species with four anther appendages (G. cernua Bak. and G. patens Miq.) in Malaya. Ny | Po IPERLIS “Gp... we ‘i THAILAND Miles PULAU Rea 0 10 20 30 40 50 ea ; anne: LANGKAWI PENANG i ! \ ! ! \ ones M.S SOUTH CHINA i] Land above 5,000 ft. Land between 3,0001tO. and 5,000 ft. Land between 1,000 ft. 4 and 3,000ft. VY ¢ Curtisii Holtt.(herb. Coll.) We . Curtisii Holtt. (fresh colt.) Se ? fragilis Lim - A EP 7) ¢, Holttumii Lim 2S 8 JOHORE Holttumii ssp “aurea Lim Marantina L. variabilis Ridl. (herb. Coll.) Globba variabilis Ridl. (fresh colt.) Globba yariabilis ssp. pusilla Lim oo SINGAPORE Fig. 3. Distribution map of Globba species with four anther appendages (G. curtisii, Holtt., G. fragilis Lim, G. holttumii Lim. G. marantina — Cytogenetics and Taxonomy of the Genus Globba L. 123 A comparison of the distribution of Globba species with only 2n = 32 and those with only 2n = 48 shows that, with the exception of G. leucantha, both are rather restricted and localised in occurrence. Species with 2n = 32 have been found on lowland forests and at moderate elevations, while species with 2n = 48 usually occur in montane forests. The exception was G. holttumii ssp. aurea (2n = 48) found so far in lowland Dryobalanops forests in north- eastern Johore. G. marantina L. (2n = 80) was found in Ayer Itam, Penang and Jason Bay, Johore. Living materials for investigation were collected from the waterfall Botanic Garden, Penang. The paucity of collections may be attributable to its seasonal dormancy for about four months annually when it is nowhere to be seen. G. marantina is apparently localised in distribution in Malaya, but has a very widespread distribution in the Indo-Malaysian region — from India (G. strobilifera Zoll. & Mor. = G. marantina L., G. bulbifera Roxb. = G. marantina L., fide Index Kewensis, 1895, herbarium sheet examination of G. bulbifera Roxb. in comparison to living kpecimens of G. marantina L.) to Sumatra (G. bracteata Heyne = G. marantina L., fide Index Kewensis, 1895) to the Philippines, New Guinea, the Moluccas Islands and the Solomon Islands where G. marantina is the only species of the genus. Pendleton (1949) surmised that since the Pulau Islands, west of Solomons, were very similar to the Solomons, G. marantina might be there too. The ecological success of G. marantina may be attributed to its pro- pagation by rhizome and bulbils, both being very resistant to adverse conditions, and its ability to colonise efficiently open habitats where competition for survival would be less keen. The other Malayan Globba species thrive well only in moist, partially shaded niches. A study of intraspecific distribution in the four Globba species with 2n = 32, 48 shows that the subspecific taxa with 2n = 48 are generally but not strictly montane in distribution. Almost all the chromosomal races with 2n = 32 in G. pendula Roxb. s.s. are lowland in distribution, the one with 2n = 48 is solely montane. Those of G. patens Miq. s.s. show a similar distributional distinction, with the exception of the single collection of specimens with 2n = 48 from Bukit Lagong Forest Reserve, Selangor (c.200-800’). Both the chromosomal races of G. cernua Bak. s.s. are restricted to the highlands; but those of G. cernua ssp. crocea are found in lowland forests. In contrast, G. variabilis Ridl. s.s. (2n = 48) occurs both in the lowlands as well as in montane area, though predominantly in the latter. The subspecies pusilla has been collected from localities of moderate elevation. The ecological spread of chromosomal races with 2n = 32 of G. pendula Roxb. s.s. and G. patens Mig. s.s. was found to be more extensive than that of chromosomal races with 2n = 48. In G. cernua Bak. s.s., where the two chromosomal races occur together in montane areas, plants with 2n = 48 are more abundant than plants with 2n = 32. 124 Gardens’ Bulletin, Singapore — X XVI (1972) Acknowledgements This paper contains material extracted from a thesis accepted for the degree of Ph.D. in the University of Malaya. I wish to record my gratitute to Professor R. E. Holttum (Royal Botanic Gardens, Kew) and Dr. B. C. Stone (School of Biological Sciences, University of Malaya) for their kind assistance and advice. I wish to acknowledge the generous loan of herbarium specimens from the Botanic Gardens, Singapore, the Royal Botanic Gardens, Kew and the Central National Herbarium, Calcutta. I would also like to thank all those who have helped in the collection of live specimens and Miss Kuan Lai Wah for her efficient typing of this manuscript. Table 2. Source and Chromosome Numbers of Living Specimens of Malayan Globba Species. G. marantina L.: PENANG, Waterfall Gardens. lowland, LIM Siew-Ngo, KLU 4831, 2n = 80. G. cernua Bak. PAHANG, Western Hill, Fraser’s Hill, 4000’, LIM Siew-Ngo, KLU 4844, 2n = 32. ———., idem, KLU 4843, 2n = 48. —_—., idem, KLU 4832, 2n = 48. —, Bukit Peninjau, Fraser’s Hill, 4000’, LIM Siew-Ngo, KLU. $205. 20 =..42. ———., idem, M.£.D. POORE, KLU 4796, 2n = 48. —_—, Parit Falls, Cameron Highlands, 4700’, LIM Siew-Ngo, KLU 8233, 2n = 48. —., Robinson Falls, Cameron Highlands, 4400’, LIM Siew-Ngo, KLU 8235, 2n =-48. —, Gunong Jasar, Cameron Highlands, 4500’, LIM Siew-Ngo, KLU 8222, 2n = 48. , idem, *, 2a. 48: PERAK, Maxwell’s Hill, 3750’, LIM Siew-Ngo, KLU 4833, 2n = 32. —_—., idem, KLU 4834, 2n = 48. SELANGOR, Reid’s 3rd quartz ridge, Gombak, 12th mi., 1800’, A. LETHBRIDGE, KLU 8228, 2n = 32. G. cernua ssp. PAHANG, Jenka Forest Reserve, 200’, M.E.D. POORE, KLU 4791, crocea Lim an FZ. —_——., idem, I. CLEAR & C. C. HO, KLU 4817, 2n = 48. G. cernua ssp. PERAK, Bujang Melaka, 1370’, K. JONG, KLU 8240, 2n = 32. prophyria Lim G. unifolia var. KELANTAN, Sungei Lebir, 300’, B. C. STONE, KLU 7367, 2n = 32. sessiliflora Holtt. G. fragilis Lim KEDAH, Pulau Langkawi, lowland, K. C. CHEANG, KLU 4847, Ut = "S72. G. curtisii Holtt. PAHANG, Waterfall by old dairy farm, Fraser’s Hill, 4000’, K. JONG, KLU 4793, 2n = 48. Cytogenetics and Taxonomy of the Genus Globba L. 125 G. holttumii Lim G. holttumii ssp. aurea Lim G. X intermedia = G. patens xX cernua Lim G. patens Miq. G. patents var. costulata Lim G. variabilis Ridl. G. variabilis ssp. pusilla Lim , idem, KLU 4822, 2n = 48. SELANGOR, Gombak Forest Reserve, 2500’, LIM Siew-Ngo, “2 =<40, —_—-, Genting Simpah, 2700’, K. JONG, *, 2n = 48. ——., Genting Highlands, 4000’, LIM Siew-Ngo, *, 2n = 48. JOHORE;. Lenggor Forest Reserve, 300’, P. C. LEE, KLU 8206, 2n = 48. PAHANG, Parit Falls, Cameron Highlands, 4700’, LIM Siew-Ngo, KLU 4840, 2n = 48. ——., Robinson Falls, Cameron Highlands, 4400’, LIM Siew-Ngo, KLU 4841, 2n = 48. —-——, Gunong Jasar, Cameron Highlands, 4800’, LIM Siew-Ngo, KLU 4842, 2n = 48. ———., idem, KLU 4848, 2n = 48. SELANGOR, Gombak 22nd mi., 1500’, LIM Siew-Ngo, KLU BALL. 2 = 32. ———, Genting Simpah, 1800’, LIM Siew-Ngo, KLU 8219, 20y = 92; ———., Kanching Dryobalanops Forest Reserve, lowland, Honours students, KLU 4827, 2n = 32. — ———, Ulu Langat Forest Reserve, 700’, T. WHITMORE, *, 2 = 32. ——, Gombak Forest Reserve, (12th mi.), 1500’, A. LETH- BRIDGE, KLU 4849, 2n = 32. JOHORE,, Eengcor Forest: Reserve. 2007, P. C.. Lee, *, 2n = 32. PAHANG, by Golf Course, Fraser’s Hill, 4000’, K. C. CHEANG, KLU 6232, 2a = 46. —, Genting Highlands, 4000’, B. C. STONE, KLU 6588, 2n ‘= 48. PERAK, Bujang Melaka, 2200’, K. JONG, *, 2n = 48. SELANGOR, Bukit Lagong Forest Reserve, 200-800’, M. E. D. POORE, *; 2n'= 48. PAHANG, Waterfall by old dairy farm, Fraser’s Hill, 4000’, A. LETHBRIDGE, KLU, $209, 2n = 32. —-—---, Cameron Highlands, 4750’, W. L. CHEW, KLU 8231, Zhe. 32. SELANGOR, Sungei Lallang Forest Reserve, Kajang, 250’, J. DRANSFIELD, *, 2n = 32. SELANGOR, Gombak Forest Reserve (12th mi.), 1500’, LIM Siew- Ngo, KLU 8208, 2n = 48. - Sungei Lallang Forest Reserve, Kajang, 200’, J. DRANS- FIBLD, :*,.2n°= 48. PAHANG, Fraser’s Hill, 4000’, LIM Siew-Ngo, *, 2n = 48. SINGAPORE, Bukit Timah Nature Reserve, 500’, LIM Siew- Ngo, *, 2n = 48. JOHORE, Gunong Panti, 1500’, W. L. CHEW, KLU 8246, 2n = 32. 126 G. albiflora Ridl. Gardens’ Bulletin, Singapore — XXVI (1972) PENANG, Penang Hill, 1300’, LIM Siew-Ngo, KLU 8221, 2n = 372. G. pendula Roxb. SELANGOR, University Campus, Kuala Lumpur, lowland, LIM Siew-Ngo, KLU 4799, 2n = 32. ——-—-—-, Kanching Dryobalanops Forest Reserve, lowland, Honours students, KLU 4813, 2n = 32. -, Genting Simpah, 1800’, K. Jong, KLU 8213, 2n = 32. ———, Ayer Hitam Forest Reserve, 350, P. C. LEE, KLU 5201) 2 =" 32: PERAK, Ulu Kinta, off Ipoh, lowland, P. C. LEE, *, 2n = 32. PENANG, Waterfall Gardens, lowland, LIM Siew-Ngo, KLU S210, 20 =" 32. KEDAH, Foot of Gunong Jerai, lowland, LIM Siew-Ngo, KLU 8225, 2n = 32, , Jungle south of Gunong Jerai, lowland, LIM Siew- Ngo, KLU 8226, 2n = 32. PAHANG, Waterfall by old dairy farm, Fraser’s Hill, 4000’, LIM Siew-Ngo, KLU 8203, 2n = 48. —— —.,, idem KLU 4836, 2n = 48. G. pendula var. elegans (Ridl.) Holtt. PAHANG, idem, LIM Siew Ngo, KLU 4830, 2n = 32. G. pendula ssp. PENANG, Penang Hill, 1200’, LIM Siew-Ngo, KLU 8219, 2n = 32. montana (Ridl,) ———., Waterfall Gardens, lowland, LIM Siew-Ngo, KLU 8216, Lim Zu. = 32: ——., Ayer Itam Dam, 1090’, LIM Siew-Ngo, KLU 8229, 2n =="SZ, G. leucantha var. JOHORE, Gunong Panti, 1500’, J. SINCLAIR, KLU 8200, 2n = 32. peninsularis Holtt. SINGAPORE, Bukit Timah Nature Reserve, 500’, LIM Siew- Ngo, KLU 4839, 2n = 32. Bukit = Bill Gunong = Mountain Sungei = River Pulau = Island 5 = Sine specimen exsiccaitum LITERATURE CITED Holttum, R. E. (1950) The Zingiberaccae of the Malay Peninsula. Gard. Bull. Sing. XIII (1): 1-249. Lim Siew-Ngo (1972 a) Cytogenetics and Taxonomy of the genus Globba L. (Zingiberaceae) in Malaya I. Taxonomy. Notes Roy. Bot. Gard. Edinburgh 31(2) : 243-269. (1972 b) Cytogenetics and Taxonomy of the genus Globba L. (Zingiberaceae) in Malaya II. Cytogenetics. Notes Roy. Bot. Gard. Edinburgh 31(2) : 271-285. Index Kewensis — An Enumeration of the Genera and Species of Flowering Plants (1895) Vol. I. Oxford. ——— - (1901-1906 to 1966) Supplements I to XIII. Oxford. Pendleton, R. C. (1949) “The rain shadow effect on the plant formation of Guadalcanal.” Ecol. Monogr. 19: 75-93. Two new Theaceous plants from Malaysia and a proposal to reduce Tutcheria to a synonym of Pyrenaria by HSUAN KENG Department of Botany University of Singapore, Singapore 1. Introduction Late 1960, I examined several specimens, incl. Elmer 21628, 21832, and Wood SAN 16482 (Plate 1) from North Borneo which is now known as Sabah, a state of Malaysia. These were in the Herbarium of Singapore Botanic Gardens and identified as Camellia lanceolata (B1.) Seem. A study of the fruit (a 3-loculate capsule with thin woody wall, partly dehiscent) and the seed (exalbuminous, the embryo with large thin, contorted cotyledons) reminded me of those of Pyrenaria and Tutcheria. Later, through Professor C.G.G.J. van Steenis, I learned from Mr. J. R. Sealy, that the identification of these specimens as Camellia lanceolata was erroneous. Dr W. Meijer, the former Forest Botanist from Sandakan, Sabah, kindly sent all the theaceous specimens belonging to this group on loan to Singapore. But I failed to find any flowering material of this plant. He also informed me that the forest in Tawau area, where the collections were made, had since been felled. Early 1970, when checking some unidentified theaceous specimens, my attention was drawn to a specimen from the Malay Peninsula (Corner s.n., on Sept. 9, 1937, from Sungei Tahan, Pahang) (Plate 2). It had for many years been preliminarily identified as Euphorbiaceae, but Mr. H. K. Airy-Shaw of the Royal Botanic Gardens, Kew, wrote on an annotated sheet ‘‘cf. Theaceae’’, when he worked over the un-named Singapore sheets of Euphorbiaceae in February 1969 in preparation for the chapter in the new Tree Flora of Malaya. The fruit and seed structures of this plant reminded me again of those of Pyrenaria and Tutcheria. At my request, this specimen together with another old collection from the same area (Haniff & Nur 8067) and recognized by Ridley as a new species of Pyrenaria, was sent to Mr. Airy-Shaw for further confirmation. He agreed that these two specimens represent a hitherto undescribed species belonging to the Pyrenaria- Tutcheria complex. Originally, I planned to collect at least the flowering material of the Malayan plant. During a recent trip (June, 1971) to Pahang in Malaya with Dr. T. C. Whitmore, we reached the lower end of Sungei Tahan. Unfortunately, our efforts to find this plant failed. I decided then that it would be better to describe these two species of Pyrenaria (one from Sabah, one from Malaya) without further delay. A discussion of the reason why Tutcheria should be merged with Pyrenaria with a special note on their fruit, seed and seedling characters follows the description. 127 128 Gardens’ Bulletin, Singapore — X XVI (1972) Fig. | Pyrenaria tawauensis H. Keng, sp. nov. Habit sketch, fruit and seed (based on Wood SAN 16482, Supplemented by Elmer 21628) Two new Theaceous plants from Malaysia 129 2. Two new species of Pyrenaria from Malaysia 1. Pyrenaria tawauensis H. Keng sp. nov. (Plate 1, figure 1) Arbor ad 10 m alta. Ramuli puberuli. Folia disticha, subcoriacea. anguste elliptica vel anguste ellipto-lanceolata, 10-19 cm longa, 44.5 cm lata, apice acuta vel caudata, basi acuta, supra glabra, subtus verrucosa vel fere glauca, margine repanda, nervis lateralibus 8-12, bene intra marginem anatomosantibus; petiolo ca. 1 cm longo. Capsula ovoidea vel subglobosa, 3—3.5 cm longa, 2.5—3.5 cm in diametro; pericarpio tenuiter lignoso. Semina atropurpurea, 1.2—2.5 cm longa. A small or medium-sized tree, 10 m tall. Branches slender, puberulous. Leaves alternate, distichous, thin coriaceous, narrowly elliptic or narrowly elliptic- lanceolate, 10-19 cm long, 4—4.5 cm wide, acute or shortly to long-caudate at the apex and gradually or abruptly narrowed to the acute base, shining glabrous above, verrucous, puberulent or sub-glaucous beneath; lateral veins 8-12 pairs, oblique, curved and merged into the submarginal vein, slightly depressed above, elevated beneath; margin remotely serrulate or repandous; petiole about 1 cm long, slender, puberulent. Flowers not seen. Fruit broadly ovoid to depressed globose, 3-3.5 cm long, 2.5-3.5 cm across, verrucous, puberulous or glabrescent, usually 3-loculate. Pericarp thin woody (2-3 mm in thickness in mature specimen) eventually partly dehiscent along the sutures. Seeds usually 2 (rarely 3) in each locule, dorsally convex-rounded, the other surfaces variously angulate due to mutual compression, 1.2-2.5 cm long, 1.2—-1.4 cm broad, dark purple, shining. Specimens examined: Sabah (North Borneo), Malaysia: Tawau, alt. 100 m, G.H.S. Wood SAN 16482 (Type, Sing.) Oct. 28, 1955; A.D.E. Elmer 21628, 21832 (Sing.), Oct. 1922 to March 1923; C.E. Carr 26387 (Sing.) March 5, 1933. This species is characterized by the fruit with thin, woody, partly dehiscent pericarp. 2. Pyrenaria pahangensis H. Keng sp. nov. (plate 2, figure 2) Arbuscula c. 4 m alta. Ramuli glabrescenti. Folia pseudo-verticillata, membranacea, elliptica vel anguste oblongo-oblanceolata, 25-38 cm longa, 8-15 cm lata, apice acuta vel breviter caudata, basi cuneata, imo subcordata, supra glabra, subtus glabrata, margine serrulata vel subintegra, nervis lateralibus 7-8, bene intra marginem anastomosantibus; petiolo 0.25-0.5 cm longo. Capsula Ovoidea vel subglobosa, 3-4.5 cm longa, 2.5-4 cm in diametro; pericarpio cartilagioseo. Semina badia, 1.5-2 cm longa. A shrub or small tree, 3-4 m tall. Branches stout, glabrescent. Leaves pseudo- verticillate (i.e., 3-5 spirally congested to form a false whorl), membranaceous, elliptic to narrowly oblong-oblanceolate, 23-38 cm long, 8-15 cm wide, acute or shortly caudate at the apex and gradually narrowed to the cuneate and cubcordate base, shining, glabrous above, puberulent or glabrescent beneath; lateral veins 7-9 pairs, oblique to nearly perpendicular, curved and merged into the submarginal vein, rather faint above, distinct and slightly elevated beneath; margin remotely serrulate, nearly entire below the middle; petiole short, pulvinoid, 0.25—-0.5 cm long. Flowers not seen. Fruit broadly ovoid or subglobose, 3-4.5 cm long and 2.54 cm across, normally 3-loculate. Pericarp very thin, cartilaginous, (1-2 mm in thickness in the mature specimen), eventually partly dehiscent along the sutures. Seeds usually 2 in each locule, dorsally convex-rounded, the other surfaces variously angulate due to mutual compression, 1.5—-2 cm long, 1.2-1.8 cm broad, chestnut brown, shining. 130 Gardens’ Bulletin, Singapore — X XVI (1972) Specimens examined: Pahang, Malaysia: Sungei Tahan, E.J.H. Corner s.n. (Type, Sing.) Sept. 9, 1937 (Common shrub or treelet, in streams, off the main river); Teku, Gunong Tahan, Mohamad Haniff & Mohamad Nur S.F.N. 8067 (Sing.) June 21, 1922 (Small tree, 10-12 ft. high). This species is characterized by its sub-verticillate, short-petiolate leaves with a subcordate base and especially by its fruit with thin cartilaginous and partly dehiscent pericarp. 3. A proposal to reduce Tutcheria to a synonym of Pyrenaria with a special note on their fruit, seed and seedling characters The genus Pyrenaria was established by C. L. Blume (in Bijdr. 1119) in 1827, based on the Javanese species, P. serrata Bl. It is interesting to note from the original description, (““Pomum subglobossum, depressum, carnosum, guinqueloculare, loculis 2 pyrenis. Pyrenae l-spermae...’’) that Blume recognized the fruit as a pome, with 2 pyrenes in each of the 5 locules. From this character, the generic name Pyrenaria was derived. Most authors today, however, generally described the fruit as succulent, baccate or drupaceous. About twenty binomials have since been described, they were from E. India, Burma, Malay Peninsula, Borneo, Sumatra, Indo-China, Thailand and S.W. China (Yunnan). Some of them would probably be merely synonyms after critical revision; for example, out of 4 species described from Java only Blume’s original species stands (cf. Backer & Bakhuizen, Fl. Java, 1: 321, 1963). The genus Tutcheria, on the other hand, was established by S. T. Dunn (in Jour. Bot. 46: 324) in 1908, based on a Hongkong species, T. championi Nakai (formerly as T. spectabilis (Champ.) Dunn). Dunn clearly pointed out that the systematic position of Tutcheria is closely next to Pyrenaria which it resembles in flower and seed and from which it is distinguished by the dehiscence of its capsule and by its multiovulate locules. About ten speciecs of Tutcheria were described, mostly from Southern China, one from Formosa, and one from the Riukiu Islands. In Melchior’s monographic study of Theaceae in Engler’s Pflanzenfamilien (2nd ed. 21: 126, 1925), Pyrenaria and Tutcheria were treated separately as belonging to two different subtribes. Recently, in Sealy’s Rev. Gen. Camellia (p. 13, 1958) brief discussion on the classification of the subfamily Camellioideae (as Tribe Camellieae), Pyrenaria, Tutcheria, Camellia and Yunnania were treated together under a subtribe, Camelliinae. Since Yunnania is rather imperfectly known, the following brief discussion on flower, fruit, seed and seedling characters is concentrated on those of Camellia, Pyrenaria and Tutcheria. I. On flowers In general, the flowers of Pyrenaria and Tutcheria, like those of Camellia, are variable to a certain extent. Whereas the subdivision of Pyrenaria into two sections for instance was based on the number and degree of fusion of the styles (Melchior, 1.c. 138); the classification of Camellia into 12 sections (Sealy 1.c. p. 28) was primarily based on a pair of characters: styles united or free, and bracteoles and sepals clearly differentiated or not. A full range of variation in flower parts of Camellia was also given by Sealy (1.c. pp. 16-18). No attempt has been made to subdivide Tutcheria. Pyrenaria tawauensis sp. nov. Type! BASE ee ERE es Sa Plate 1. Type specimen of H. Keng Foreete 7 FLORA OF NORTH BORNEO DISTRIBUTED FROM THE HERBARIUM OF THE FOREST DEPARTMENT, SANDAKANS, SL BORNEO Betantcal nase ? & MALS Kame ae 3 2 Fe we # PRatra:t & a Lee & Ps Yat Date i Sy % { Crdlectog ate Ge + i : BARGG * e + * S 8 Herhar S SAN ‘ « a* 4 Pyrenaria tawauensis H. Keng nag 6 hte ek nw Ran am wandnad _. ap. 1 Fepamnarsis, fate sn: Aa seonk fumil5 Peont! >, es es Be Wien, Chute be Yuntes o (lew, Pim eed es Woy ate, Bir 9 et ow hang a Gerais fisis be. te. BTS es #4 Rertaer th. 4% Ne a ky 3 “sR tame se Aten © ate se abl — Raby yuna T BOC OPM ME No a0 _ Pyrenaria pahangensis H. sp. nov. Type! j j Mj yy Vij 7m» \ nt Yj WWW) ty 4 4% © Wy Wy) wi a j yy Ny ing b, Y y, 4 iy ny Pa, bey type j : y Yiu re Vj, 4449 WY yy Y, j yl Via j Yj, Y y % Ky, Hl y; Wty Wis i y Yj Ny ayy ahah RPO) ‘si j sosnunen, svonse AMOS I Plate 2. Type specimen of Pyrenaria pahangensis H. Keng 13] Two new Theaceous plants from Malaysia Fig 2. Pyrenaria pahangensis H. Keng sp. nov. Habit sketch, fruit and seed (based on Corner s.n. [Sept. 9, 1937], supplemented by Haniff & Nur SFN 8067). 132 Gardens’ Bulletin, Singapore — X XVI (1972) The number of locules in an ovary in all these three genera varies from 3-5. One particular point is the number of ovules per locule. Sealy mentioned (1.c. p. 18) that in Camellia, each locule contains 3-4 (very rarely 8) axile ovules attached near the apex. Dunn gave the number of ovules in each ovary in Tutcheria as ‘2-5’. Although Blume’s original description states that there are two ovules per locule in Pyrenaria, the occurrence of three or four ovules in each locule in this genus is in fact not uncommon. Therefore, there is no real difference between Pyrenaria and Tutcheria in the number of ovules per ovary-locule as Dunn once unduly stressed. 2.. On fruit The fruit in Camellia is a woody capsule, loculicidally dehiscent from the apex downwards (Sealy, l.c. p. 18). A typical form of the fruit of Pyrenaria (e.g. P. acuminata, plate 3, c) is baccate, the pericarp fleshy or soft woody, indehiscent, 3-5-loculate. On the other hand, that of Tutcheria, like Camellia, is a woody capsule, loculicidally dehiscent into 3-6 valves, the latter often deciduous and falling from the base. The fruit characters of Pyrenaria and Tutcheria will be further discussed below. 3. On seeds and seedlings The seeds of Camellia, Pyrenaria and Tutcheria are uniformly ex-albuminous. They are generally globose, hemispheric or plano-convex in Camellia, and much flattened, often irregularly angulate (due to mutual pressure) in Pyrenaria and Tutcheria. The most noticeable fact is that in Camellia, the embryo possesses a pair of thick, hemispheric cotyledons, while in Pyrenaria and Tutcheria, it possesses a pair of very thin and large, but contorted and twisted cotyledons. Following Martin’s phylogenetic classification of seeds (Amer. Midl. Nat. 36: 513-660, esp. 519-523, 1946), Camellia seed belongs to the ‘investing’ type, and Pyrenaria and Tutcheria seed, the ‘folded’ type, both types are classified in his ‘axile’ division, and both represent the zenith of seed evolution (see his Fig. 3 on p. 523). Morphological differences of these two types are obvious, nevertheless, their selective advantages are probably even more striking. In the ‘investing’ type, the two thick cotyledons generally remain in the seedcoat and embedded in the ground, serving as a food storage organ for the initial growth of a seedling. In the ‘folded’ type, in contrast, the two thin cotyledons, during the process of germination, gradually emerge from the seedcoat, rapidly unfold and spread out, thus performing the function of photosynthesis. Previously the present writer observed the germination of species of Camellia and Tutcheria (Keng in Univ. Calif. Publ. Bot. 33: 276, Fig. 3 A & B). Photographs of seedlings of two Malayan species of Pyrenaria are presented in Plate 3 (e to A) of this paper. As expected, the germination processes of Pyrenaria and Tutcheria so far as it concerns the few species observed, are identical. From the above discussion, it can be concluded that Pyrenaria and Tutcheria, as Dunn (l.c.) pointed out, are in fact, closely related. This can be further emphasized by their peculiar seed and seedling characters which are unique among the family Theaceae (cf. Keng, l.c. p. 280). It is also clear that their only substantial difference lies in their fruit character i.e. baccate, indehiscent, in the typical forms of Pyrenaria, and capsular in the typical form of Tutcheria. Is this fruit character so vital that demarcation of these two genera can be based on it alone? A negative answer could emerge from the following four examples. %, yn Uh, Why Wy, Plate 3. Two species of Pyrenaria P. acuminata Planch.: a-flowering branch; b-flower and flower-bud: c-fruits; d-seeds: e & f-seed germination; g-seedling. P. kunstleri King: h-seedling. Two new Theaceous plants from Malaysia 133 a. Lobelia-Pratia (Campanulaceae or Lobeliaceae). Traditionally, these two were treated as two separate genera merely on fruit characters alone: apically 2-valved capsule in Lobelia Linn. and baccate in Pratia Gaudich. Moeliono & Tuyn (Fl. Mal. I, 6(1): 122, 1960) recently fused them into one genus. b. Buddleja-Nicodemia (Loganiaceae). Likewise, the fruit of Buddleja Houst. ex Linn. is a septicidal, 2-valved capsule, and Nicodemia Tenore, a berry. The latter was reduced into a subgenus of the former by Leenhouts (Fl. Mal. I, 6(2):336-337, 1962). c. Euphorbia-Elaeophorbia (Euphorbiaceae). The fruit is drupaceous in Elaeophorbia Stapf., and capsular in Euphorbia Linn, Recently Webster (Jour. Arnold Arb. 48:397, 1967) reverts Elaeophorbia to Euphorbia, but makes a separate section of the genus for it. d. Lomatophyllum-Aloe (Liliaceae). Fruit of Lomatophyllum Willd. is a large fleshy berry, whereas that of Aloe Linn. is a dehiscent capsule. Based on ‘‘Phenetic diagrams’? from the use of a computer, Rowley (in Taxon 18:625—7, 1969) strongly queried the validity of keeping the status of Lomatophyllum as a genus merely on a single character basis. Undeniably, in a number of taxa (such as Leguminosae, Umbelliferae, Ulmaceae, Juglandaceae, to mention only a few), the fruit is a paramount diagnostic character. However, in the four examples cited above, and also in case of Pyrenaria and Tutcheria, the present writer tends to believe that the fruit character, in these instances, is probably merely of ecological or physiological, rather than of phylogenetic importance. To strengthen the argument that Pyrenaria and Tutcheria should be merged, the present writer would like to supply the description of two types of fruits from Pyrenaria species which appear to bridge the gap between baccate and capsulate fruits. The first type of fruit can be found in the two new species of Pyrenaria described in this paper, both from the Malaysian region. Their fruit walls are thin woody or crustaceous, partly dehiscent along one or few sutures in a rather irregular way when fully matured. The second can be found in a Chinese species, Pyrenaria camellioides Hu from Yunnan which is characterised by the fruit with spongy pericarp partly dehiscing septicidally from above, and partly dehiscing loculicidally from below. The apex of this fruit is slightly concave, with 5 bosses (which represent the style bases) around the depression, a situation extremely similar to the Thai species, P. garrettiana Craib. The fruit of the latter, however, is succulent and not dehiscent. Incidentally, Pyrenaria garrettiana Craib, together with two other species from Yunnan, has been segregated into a new genus, Sinopyrenaria Hu (in Act. Phytotax. Sin. 5:281, 1956) and Pyrenaria camellioides Hu, into another new genus, Glyptocarpa Hu (in Act. Phytotax. Sin. 10:25, 1965). Meanwhile, a third new genus, Parapyrenaria H. T. Chang (in Act. Phytotax. Sin 8:287, 1963), was proposed based on P. hainanensis H. T. Chang, from Hainan, China. All these three are probably superfluous, if we accept a reasonably broader generic concept ot Pyrenaria, comparable to that of Camellia as recently defined by Sealy (I. c. pp. 14-372). : For the reasons given above, the present writer therefore formally proposes to merge Tutcheria Dunn with Pyrenaria Blume, and to reduce the following known species of Tutcheria to Pyrenaria. 134 Gardens’ Bulletin, Singapore — X XVI (1972) Pyrenaria Blume, Bijdr. 1119, 1827; Mig. FI. Ind. Bat. 1:493, 1855, et Suppl. 1:484, 1861; Hook. f. Fl. Brit. Ind. 1:289, 1872; Melch. in Engl. & Prant. Nat. Pflanzenfam. ed. 2, 21:138, 1925; Back. & Bakh. f. Fl. Java 1: 321, 1963. Synonym: Tutcheria Dunn in Jour. Bot. 46: 324, 1908, 47: 197, 1909; Nakai in Jour. Jap. Bot. 16: 708, 1940; Li, Wood. Fl. Taiwan, 597, 1963; Hu & Chang in Chun, Chang & Chen, FI. Hainan. 1:495, 1964. Syn. nov. A list of new combinations follows. 1. Pyrenaria championi (Nakai) H. Keng comb. nov. Basionym: Tutcheria championi Nakai in Jour. Jap. Bot. 16:708, 1940. Synonyms: Camellia spectabilis Champion ex Benth. in Jour. Bot. 3:310, 1851 (non Berlese 1843). Tutcheria spectabilis (Champ.) Dunn in Jour. Bot. 46:324, 1908. Distribution: Hongkong. 2. Pyrenaria greeniae (Chun) H. Keng comb. nov. Basionym: Tutcheria greeniae Chun in Jour. Arnold Arb. 9:129, 1928. Distribution S. China (Kwangtung). 3. Pyrenaria hirta (Hand.-Mazz.) H. Keng comb. nov. Basionym: Gordonia hirta Hand.-Mazz. Anz. Akad. Wiss. Wien. 58:180, 1920. Synonyms: Tutcheria hirta (Hand.-Mazz.) Li in Jour. Arnold Arb. 26:64, 1945. Tutcheria villosa Wu in Engler, Jahrb. 71:192, 1940. Distribution: S.W. China (Hupeh, Kweichow, Kwangsi). 4. Pyrenaria microcarpa (Dunn) H. Keng comb. nov. Basionym: Tutcheria microcarpa Dunn in Jour. Bot. 47:197, 1909. Distribution: S. China. 5. Pyrenaria multisepala (Merr. & Chun) H. Keng comb. nov. Basionym: Tutcheria multisepala Merr. & Chun in Sunyetsenia 2:41, 1934. Distribution: S. China (Hainan). 6. Pyrenaria ovalifolia (Li) H. Keng comb. nov. Basionym: Tutcheria ovalifolia Li in Jour. Arnold Arb. 25:209, 1944. Distribution: §. China (Hainan). Two new Theaceous plants from Malaysia 135 7. Pyrenaria shinkoensis (Hayata) H. Keng comb. nov. Basionym: Thea shinkoensis Hayata in Jour. Coll. Sci. Tokyo 30 (1):45, 1911, Synonyms: Camellia shinkoensis (Hayata) Cohen-Stuart in Meded. Proefst. Thee 40:68, 1916. Tutcheria shinkoensis (Hayata) Nakai in Jour. Jap. Bot. 16:708, 1940. Distribution: China (Taiwan). 8. Pyrenaria symplocifolia (Merr. & Metcalf) H. Keng comb. nov. Basionym: Tutcheria symplocifolia Merr. & Metcalf in Lingnan Sci. Jour. L672 P93: Distribution: S. China (Kwangtung). 9. Pyrenaria virgata (Koidz.) H. Keng comb. nov. Basionym: Thea virgata Koidz. in Bot. Mag. Tokyo 32: 252, 1918. Synonyms: Camellia virgata (Koidz.) Makino & Nemoto, Fl. Jap. (ed. 2): 740, 1931. Tutcheria virgata (Koidz.) Nakai in Jour. Jap. Bot. 16:708. 1940. Distribution: The Riukiu Islands. The geographic range of Pyrenaria, after the merging of Tutcheria species into it, thus covers from E. India, Burma, Thailand, Indo-China to S. China, the Riukiu Islands, Taiwan and to Malesia (Malaya, Borneo, Sumatra, Java), and conforms closely to those of Camellia (Sealy, 1. c.), Coptosapelta (Steenis, in Amer. Jour. Bot. 56: 806, 1969) and a host of other genera. ACKNOWLEDGEMENTS I would like to express my sincere gratitude to. Mr. H. K. Airy-Shaw, Dr. R. C. Bakhuizen van den Brink, Jr., Dr. P. W. Leenhouts and Professor C. G. G. J. van Steenis for useful discussions, responsibility for the views expressed in this paper, however, is solely mine. I also like to thank the directors of the Singapore Botanic Gardens and the curators of the herbarium for the facilities provided, to Dr. C. X. Furtado for going through the Latin descriptions, to Dr. T. C. Whitmore for arranging a field trip to Pahang, Malaysia, to my wife, Mrs. Ro-siu Ling Keng for preparing two habit sketches, and to Mr. D. Teow for taking the photographs which are reproduced in the plates of this paper. a ' - h = nel : f ‘ vit. fT ee A We ean fqn v@ 44 7 4 evky: he. ate ,. oy ee >. a , ile Ligne at, wer . yalte Aomwlet) a i: ' hay M Mapegidiglise.2 lao nt Lae? i% 4 ‘ | intavt Gabi a ie wing jt yom * i a rua vs AT per! bol mnt ata Oo a f ) butt beki Beas! we (hin | sand > ato al a a | err yi 4, C11 Ww a ei mee hoi - ety SVIORN 1 easnqes os PROP: Atel ee & Ries te. » Ania 40) ‘at iz Osis | pain lsh i aavow om 7 oils win 5 tw eto ews “pili tnt iets a a i wr f r i we om 033 Mount Maguiling Bryoflora (Luzon) By P. T1xIER* Résumé L’auteur analyse ses récoltes personnelles faites au Mont Maquiling en 1965. Cette liste comprend 34 espéces de Mousses et 42 espéces et variétés d’Hépatiques. Il y a 16 Mousses nouvelles pour la station et une pour la Science. Les chiffres concernant les Hépatiques s’élévent 4 35 et 5 espéces et variétés nouvelles. Les espéces et variétés nouvelles sont: Endotrichella maquilinensis; Cololejeunea pseudoschmidtii, CC. haskarliana var. luzonensis et var. thermarum, C. selaginellicola, et C. armata. Summary The author analyses in this paper his collections from Mt. Maquiling of 1965. This list contains 34 species of Mosses and 42 species and varieties of the Liverworts. Of the mosses 16 are new for this hill station and one for science. Liverworts number 35 as novel records and new species and varieties 5. The species and varieties described here as new are:— Endotrichella maquilinensis; Cololejeunea pseudoschmidtii, C. haskarliana var. luzonensis and var. thermarum, C. selaginellicola, and C. armata. This paper contains the list of the author’s collections on Mt. Maquiling of 10th July 1965, consisting of numbers 1358 to 1457, a complete set of which is deposited at the “Laboratoire de Cryptogamie, Muséum National d’Histoire Naturelle” in Paris. The reader is reminded that Mt. Maquiling is a dead volcano, rising up to 1100 m, on the side of Laguna de Bay — the inland lake of central Luzon — at sixty kilometers from Manila. The annual rainfall at the College of Agriculture in Los Banos at the foot of the hills reaches 1930 mm annually for 179 rainy days (figure I). The botanical aspect of this hill station is renowned by the work of the great north American botanists early in the century: Merrill, Copeland and Elmer. I acknowledge with thanks The South East Asia Treaty Organisation which permitted me to stay in The Philippines, and Dr. D. Umali, Dean of The College of Agriculture, who kindly gave the material for my collecting trip. In addition, I express my gratitude to Professor J. V. Pancho of the Applied Botany Department. From his experience and knowledge of the mountain I profited much when he guided us in the forest of the volcano. To Dr. C. X. Furtado, Botanist in Singapore, I am grateful for his corrections of the Latin diagnosis in this paper. I MUSCI 1. Fissidens sylvaticus Griff. — Corticolous, no. 1435. Thailand, Cambodia, Vietnam, Sumatra, Java, Celebes, Borneo, Hongkong, Bataan, Luzon (M.Q.). 2. Leucoloma molle (C. Muell.) Mitt. — On twigs and trunks, nos. 1410, 1443, 1444, 1436. Widely distributed from Indo Malaya to Pacific Islands, Hawaii, China, Japan, Luzon (Maquiling and Mountain Province), Mindoro, Palawan, Mindanao. 3. Dicranodontium uncinatum (Harv.) Jaeg. — On twigs, no. 1403. India, Thailand, Japan, Moluccas, Luzon (Benguet and Mt. Banahao). * Laboratoire de Cryptogamie, Muséum National d’Histoire Naturelle (Paris), and, Faculté des Sciences, Phnom Penh (République Khmére) 137 138 Gardens’ Bulletin, Singapore — X XVI (1972) ‘5 Sa ' MANILA Laguna de Bay a Tagaytay amt Maquiling , \.Mt Banahao Taal Lucena »Batangas ern) MANILA and surroundings Rain/month mm Summer monsoon Autumn monsoon Temperature YZ Dry season OT EM AM J DOR 3 eo hep CLIMATOGRAM of LOS BANOS Fig. I. Southern part of Luzon. Climatology of Los Banos. Mount Maquiling Bryoflora 139 4. Leucobryum bowringii Mitt. — Corticolous, nos. 1420, 1436. Ceylon, Malaya, China, Japan, Formosa, Vietnam, Cambodia, Thailand, Bataan, Luzon, Sibuyan, Negros. 5. Calymperes serratum A. Braun — Corticolous, no. 1435. Malaya, Eastern China, Vietnam, Cambodia, Thailand, Fiji, Samoa, New Caledonia, Luzon (MQ). 6. Syrrhopodon tristichus Nees ex Schwaegr. — On twigs, nos. 1416, 1420, 1424, 1425. Ceylon, Sumatra, Java, Ambon, Borneo, Luzon, Negros. 7. Macromitrium cuspidatum Hampe — On trunks, nos. 1400 (c. fr.), 1410. Sumatra, Cambodia, Java, Borneo, Luzon, Mindoro, Palawan. 8. Macromitrium semipellucidum Doz. & Molk. — On branches, nos. 1432 (c. fr.), 1433. It is nearly impossible to separate M. semipellucidum from M. pungens. According to Bryologica javanica only one character is different i.e. the base of the leaf in M. semipellucidum is bordered with small cells. Luzon, Panay, Mindanao, Vietnam, Cambodia, Sumatra, Borneo, New Guinea. 9. Rhizogonium spiniforme (Hedw.) Bruch — On trunks in low positions, nos. 1436, 1437. Pantropical. 10. Hypnodendron vitiense Mitt. — Very common on trunks, in ridge forest, nos. 1430, 1443, 1446, 1447. Fiji, Solomon Islands, Luzon (MQ), Negros, Mindanao. 11. Mniodendron fusco-mucronatum (C. Muell.) Broth. — On rocks, circa 600 m; nos. 1416, 1455. Luzon, Negros, Biliran, Mindanao. 12. Endotrichella maquilinensis P. Tixier sp. nov. — On twigs, nos. 1400 (c. fr.), 1414, 1418 (c. fr.), 1424, 1425. 13. Endotrichella compressa (Mitt.) Broth. — On twigs, nos. 1400 (c. fr.), 1436. Borneo, Luzon. 14. Symphysodon subneckeroides Broth. — On trunk, nos. 1424, 1400. Sumatra, Java, Borneo, Panay, Luzon, Negros, Mindanao, New Caledonia. 15. Aerobryopsis longissima (Doz. & Molk.) Fleisch. var. prostratula (C. Muell.) Fleisch. — On twigs, nos. 1418, 1424, (c. fr.), 1399. Luzon. 16. Aerobryidium cf. filamentosum (Hook.) Fleisch. — On twigs, no. 1403. Near A. filamentosum by length and colour of leaves, differing only by shape of the cells. I also found this taxon in South Vietnam. Nilghiries, Sikkim, Ceylon, Yunnan, Szechwan, Sumatra, Vietnam, Laos. 17. Barbella cf clemensiae Broth. — On trunks, nos. 1401, 1437. Luzon, 140 Gardens’ Bulletin, Singapore — XXVI (1972) 18. Himantocladium cyclophyllum (C. Muell.) Fleisch. — On trunks, circa 700 m, no. 1455. Java, Celebes, Borneo, New Caledonia, Tahiti, Luzon (MQ), Mindoro, Negros, Biliran, Mindanao, Basilan. 19. Homaliodendron flabellatum (Sm.) Fleisch. — On trunk, no. 1457. India, Ceylon, Japan, Malaya to Hawaii Islands, Philippines, New Caledonia, Indochina. 20. Pinnatella cf. ambigua (Bosch. & Lac.) Fleisch. — no. 1435. This small plant, 1 cm high, may be allied to P. nana (Williams) Fleisch. Bhotan, Burma, Sumatra, Java, Thailand, Luzon, Palawan, Negros, Mindanao. 21. Eriopus microblastus Broth Very small specimen, epiphytic on Clastobryum cuculligerum. The drawing of Bartram is correct but it seems better to follow Brotherus and to separate E. microblastus from E. parviretus. The denticulation of the edge is, in the former species, denser, no. 1409. Luzon, Negros. 22. Chaetomitrium perarmatum Broth. — On twigs, nos. 1408, 1411, 1414, 1415. Mountain Province, Apayo district (Luzon). 23. Thuidium glaucinum (Mitt.) Bosch. & Lac. — On twigs, no. 1400. Himalaya, Southern India, Ceylon, Thailand, Cambodia, Vietnam, Japan, Luzon (MQ), Negros. 24. Clastobryum cuculligerum (Lac.) P. Tixier — Whitish mat on twigs, nos. 1409, 1412 (c. fr.), 1416 (c. fr.). Java, Thailand, Cambodia, Vietnam, Luzon (Benguet), New Guinea, New Caledonia. 25. Trismegistia calderensis (Sull.) Broth. — nos. 1441, 1444. Sumatra, Malaya, Luzon (MQ). 26. Acanthocladium piliferum (Broth.) P. Tixier comb. nov. Syn.: Acanthocladium longipilum Broth. Beih. Bot. Centralbl. 28, 2, (1909), 361. Raphidostichum eberhardtii (Thér. & Varde) Broth. Nat. Pfl. 2, 11, (1925), 434. Raphidostichum luzonense Broth. Nat. Pfl., 2, 11, (1925), 435. Raphidostichum piliferum (Broth.) Broth. Nat. Pfl., 2, 11, (1925), 434. On twigs and trunk, nos. 1426, 1445. Species of variable size, distributed from Thailand to Philippines. 27. Warburgiella cupressinoides C. Muell. — On branches, nos. 1413, 1421. Luzon, Panay, Mindanao, New Caledonia. 28. Acroporium diminutum (Brid.) Fleisch. — On twigs. nos. 1416, 1420. Sumatra, Java, Malaya, Luzon (MQ), Mindanao, Moluccas. Mount Maquiling Bryoflora 141 29. Acroporium hermaphroditum (C. Muell.) Fleisch. — On trunk, no. 1455. Thailand, Sumatra, Java, Celebes, Moluccas, New Guinea, New Caledonia, Luzon (MQ), Mindanao. 30. Acroporium secundum (Reinw. & Hornsch.) Fleisch. — On trunks and twigs, nos. 1402, 1425, 1426, 1440, 1446, 1454. Java, Sumatra, Borneo, Celebes, Ceram, Luzon (MQ); Negros, Mindanao. 31. Ectropothecium buitenzorgii (Bel.) Jaeg. — On twigs and trunks, nos. 1413, 1438. Widely distributed in South East Asia. 32. Ectropothecium falciforme (Doz. & Molk.) Jaeg. — On trunk and twigs, nos. 1413, 1426, 1442, 1444, 1448. Luzon, Panay, Negros, Sumatra, Java, Lombok, Borneo, Celebes. 33. Ectropothecium ichnotocladum (C. Muell.) Jaeg. — On trunk, no. 1398. Luzon (MQ), Bohol, Mindanao, Sumatra, Java, Borneo, Celebes, Ambon, Vietnam, New Caledonia. 34. Isopterygium albescens (Hook.) Jaeg. — Twigs, nos. 1415, 1419, 1436. India and Indochina, Malesia, Japan, Pacific Islands. Ii HEPATICAE 1. Metzgeria furcata (L.) Dum. — Epiphyllous, no. 1370. Cosmopolitan. 2. Chiloscyphus communis St. — On trunk in low position, no 1436. Assam, Vietnam, Cambodia, Sikkim, Japan, Java, Borneo, Queensland, Luzon (MQ). 3. Schistochila aligera Nees & Bl. — On trunk, nos. 1443, 1447. Java, Sumatra, Moluccas, New Hebrides, Nicobar, Borneo, Ambon, Philippines, Samoa, Vietnam, Thailand. 4. Lepidozia cf. subintegra Ludg. — no. 1439. South East Asia. 5. Herberta cf. minima Horik. — On trunk, very poor specimen, no. 1445. Formosa. 6. Trichocolea pluma Mont. — no. 1446. Cosmopolitan. 7. Radula anceps Lac. — Epiphyllous, no. 1378. Java, Luzon, New Caledonia. 8. Radula borneensis St. — Epiphyllous on Selaginella sp. no. 1359. Southern India, Borneo, Vietnam, Cambodia. 9. Archilejeunea mariana (G.) St. — Epiphyllous, no. 1445. Formosa, Botel Tobago, Thailand, Sumatra, Java, Sarawak, Philippines, Celebes, New Guinea, Mariannas, New Caledonia, Solomon Islands, Tahiti. 142 Gardens’ Bulletin, Singapore — X XVI (1972) 10. Ceratolejeunea maritima St. — On twigs, corticolous, no. 1409. Carribean region. 11. Cheilolejeunea (Xenolejeunea) ceylanica (G.) Schust. & Kachr. Epiphyllous, no. 1405. Tropical Asia, Ceylon, Pulau Penang, Ambon, New Guinea, Formosa, Vietnam, Cambodia. 12. Cololejeunea floccosa (Taeniolejeunea) (L. & L.) St. Epiphyllous, no. 1395. Japan, Ryukyu, Formosa, China, Philippines, Borneo, Java, Sumatra, Vietnam, Chittagong, Thailand, Cambodia, Laos, Tanzania. 13. Cololejeunea gynophthalma (Yaeniolejeunea) Bx. — Epiphyllous, no. 1396. Sumatra, Java, Vietnam, Cambodia, Malaya. 14. Cololejeunea maquilinensis (Taeniolejeunea) P. Tixier — Epiphyllous, no. 1394. Endemic. 15. Cololejeunea pseudoschmidtii (Globigerae) P. Tixier sp. nov. Epiphyllous, no. 1394 (TYPE). 16. Cololejeunea hispidissima (St.) P. Tixier comb. nov. (Syn.: Leptocolea hispidissima St. Spec. Hep., VI, 243, 1923). Epiphyllous, no. 1394. 17. Cololejeunea haskarliana (G.) Bx. var. luzonensis var. nov. Epiphyllous, no. 1375: 18. Cololejeunea haskarliana (G.) Bx. var. thermarum var. nov. Epiphyllous, no..1377- 19. Cololejeunea selaginellicola (Venustae) P. Tixier sp. nov. Very common, epiphyllous, no. 1383. 20. Cololejeunea armata (Venustae) P. Tixier sp. nov. — Epiphyllous no. 1383. 21. Cololejeunea flavicans (Lasiolejeunea, latebrosae) (St.) Mizutani. Epiphyllous, nos. 1360, 1362. Endemic of Luzon. 22. Cololejeunea aff. nymanii (Lasiolejeunea, latebrosae) (St.) Bx. — Epiphyllous, nos. 1364, 1397. Sumatra, Java, Borneo, New Guinea, Thailand. Vietnam, Cambodia. 23. Cololejeunea panchoana (Lasiolejeunea, latebrosae) P. Tixier — Epiphyllous, no. 1358. 24. Cololejeunea vulcania (Lasiolejeunea, latebrosea) P. Tixier — Epiphyllous, no. 1392. 25. Cololejeunea aff. yulensis (Lasiolejeunea, latebrosae) (St.) Bx. Epiphyllous, nos. 1361, 1363, 1368, 1384, 1388. Malaya, Sumatra, Java, Borneo, New Guinea, Cambodia, Vietnam. 26. Colura acutifolia $.J.A. — Epiphyllous, no. 1397. Borneo, Sumatra, Chittagong, Thailand, Vietnam. Mount Maquiling Bryoflora 143 27. Drepanolejeunea bakeri Herz. — Very common on the leaves. Endemic of Philippines, known also of Mt. Polis in Mountain Province. 28. Drepanolejeunea dactylophora (Nees) Spruce ex Herz. — Epiphyllous, no. 1383. Luzon, Mindanao, Malaya, Java, Ambon. 29. Lejeunea flava (Sw.) Nees. — Corticolous. Tropical Asia and Oceania. 30. Leptolejeunea foliicola St. — Epiphyllous no. 1360. Malaya, Sumatra, Java, Vietnam, Cambodia. 31. Leptolejeunea schiffneri Herz. — Epiphyllous, no. 1446. Andaman, Malaya, Sumatra, Java, Borneo, Celebes, Moluccas, Philippines. 32. Lopholejeunea applanata (R.B.N.) St. — Epiphyllous, no. 1446. Java, Ryukyu, China, Japan, Borneo, Sumatra. 33. Lopholejeunea subfusca (Nees) St. — Corticolous, no. 1401. Japan, Ryukyu, India, Ceylon, Malaya, Sumatra, Java, Borneo, Philippines, New Guinea, New Caledonia, Northern and Southern America, Africa. 34. Microlejeunea cucullata (Nees) St. — Epiphyllous, no. 1339. Tropical Asia and Oceania. 35. Ptychocoleus cumingianus (Mont.) Trev. — Corticolous, no. 1435. Andaman, Nicobar, Sumatra, Singapore, Malaya, Java, Borneo, Philippines, Indochina, Celebes, Ambon, New Guinea, Queensland, Marquesas, Samoa, Tahiti, Caroline Islands. 36. Pycnolejeunea bidentula St. — Epiphyllous, s.n. Vietnam, Formosa, Ryukyu, Java, Borneo, Philippines, New Guinea, Fiji. 37. Pycnolejeunea eximia S.J.A. & Tixier — Corticolous, no. 1432. Close to P. fitzgeraldii St. Vietnam, China. 38. Stenolejeunea apiculata (G.) Schuster — Epiphyllous, no. 1360, 1388. Formosa, Central Vietnam, Java, Cambodia, New Caledonia. 39. Frullania apiculata Verd. — Epiphyllous, no. 1435. Ceylon, Indo-Malaya to Philippines, Moluccas, New Guinea. 40. Frullania integristipula (Nees) Nees. — On trunk, 530 m. alt. no. 1433. Java, Malaya, Philippines. 41. Frullania squarrosa (R.B.N.) Dum. — Corticolous, no. 1435. Pantropical. 42. Frullania ternatensis G. — Corticolous, no. 1378, 1424. Malaya, Ceylon, Sumatra, Java, Vietnam, Borneo, Celebes, Ternate, Bataan, Philippines. 144 Gardens’ Bulletin, Singapore — XXVI (1972) lif DESCRIPTION OF NEW SPECIES Endotrichella maquilinensis sp. nov. (Fig. II). Plantae caespitosae, caules flavi, usque ad 4 cm alti, apice divisi. Folia caulina enervia, elongata, acuminata, 4-5 mm longa, 1.2 cm lata, in parte basilari leviter plicata, apice paucidentata, auricula relative parva. Cellulae apicales pariete incrassatae, 100 » longae, 10u latae; marginales 80 x 10 yu; centrales 90-100 vp X 10-15 p; alares scutulatae, quadrataeve, 20-30 » x 10-15 yu. Folia apicalia cum propagulis, pariete aurantiacea, 1-1.6 mm longa, 20 p» lata. Fructus proprius generis; seta 0.8 mm longa; theca ovalis, elongata, 2 mm longa, 0.8 mm Jata. Bracteae perichaetiales usque ad 2.5 mm. i TAN yt ENDOTRICHELLA 1400 Fig. II. Endotrichella maquilinensis sp. nov. 1: Habit; 2-3: Leaves; 4: Leaf with propagules; 5: Auricle; 6: Leaf cells; 7: Edge cells; 8: Denticulate apex; 9: Female inflorescence; 1/0: Capsule. Plant epiphytic up to 4 cm tall, branched at the top, light yellow. Stem leaves without veins, elongated, acuminate, with some folds in the basal part of the leaf. Apex with some teeth. Auricle relatively small. Cells in apex with thick walls, 100 » long, 10 » wide; cells of the edge 80 X 10 u; cells of the central part 90-100 » X 10-15 pw. Auricle cells lozenge-shaped to rectangular 20-30 yp long, 10-15 p» wide. Leaves of the top of the stem bearing propagules with orange coloured cell walls, multicellular, 1-1.6 mm long 20 wide. Fructification charac- teristic of the genus, seta 0.8 mm long. Ripe capsule horizontal, 2 mm long and 0.8 mm wide, oval, elongated. Perichaetial bracts up to 2.5 mm. Species very different from all those reported by Bartram in Mosses of Philippines and in his subsequent papers. Characterized by the elongated shape of leaves and the edge entire almost up to the apex. Resembles EF. dimorphophylla Tx. from South Vietnam. Mount Maquiling Bryoflora 145 THE COLOLEJEUNEA COMPLEX We now arrive at a group of 6 taxa. The first is Cololejeunea hispidissima (St.) Tx. conformable to the type of Stephani. Benedix, in 1953, rendered intricate the question by putting together in synonymy, under the name Cololejeunea haskarliana (G.) Bx., taxa which are evidently different. Mizutani, in 1961, reported that C. venusta Lac. was unlike C. haskarliana (G.) Bx. sensu lato because the Sande Lacoste’s species has papillous lobule cells. In Maquiling forest the author observed the sympatry of the taxa allied to Cololejeunea haskarliana in the locality and sometimes on the same leaf. In Tjibodas (Java) up to 4 different taxa on one leaf were collected. The author intends to return to this problem. The following three taxa are allied to Cololejeunea haskarliana: Cololejeunea pseudoschmidtii is related to the Section Globigerae of Benedix, Cololejeunea selaginellicola and Cololejeunea armata to Section Venustae of Mizutani. Cololejeunea pseudoschmidtii sp. nov. (Fig. II). Planta dioica, parva, pallide virens, sicca, ad substratum appressa. Caules usque 0.5 cm longi, 0.10 mm crassi, cum foliis 1.4 mm lati. Folia inter se 0.2 mm distantia. Lobus obovalis, 0.9 mm longus, 0.5 mm latus. Cellulae basalis intermedio leviter incrassatae 20-40 pm longae, 15-20 » latae haud papillosae; marginales quadratae hexagonales, 10-20 » longae, 10-15 wu latae, cum papillis humilibus. Lobulus elongatus, inflatus, 0.25 mm longus, 0.15 mm latus, papillosus ad carenam. ae Shi Fig. II. Cololejeunea pseudoschmidtii sp. nov. 1: Stem; 2: Leaf; 3: Basal part of the lobe; 4: Edge of the lobe; 5: Young lobule; 6-8: Mature lobule. 146 Gardens’ Bulletin, Singapore — X XV] (1972) Dens apicalis parvus, triangulari-unicellulatus. Dens medianus cum cellules in 3 series dispositis, una serie, duabus alteris unicellulatus. Papilla hyaline sphaerica, sub dente media 20 » in diametro. Stylus haud visus. Androecia lateralia, 0.3 mm alta, bracteis 2-jugis. Plant dioic, small, light green, applied to the substrate. Stem 0.5 cm long, 0.10 mm thick, with leaves 1.4 mm wide. Rhizoids hyaline. Leaves spaced on stem at 0.2 mm. Lobe oboval, 0.9 mm long, 0.5 mm wide. Basal cells with light intermediate thickenings 20-40 p long, 15-20 » wide, apapillose. Edge cells rectangular to hexagonal, 10-20 » long, 15-20 » wide with a low papilla thickened at the top similar to those of C. schmidtii, Lobule elongated, inflated on mature leaves, 0.25 mm long, 0.15 mm wide, sometimes papillose near the keel. Apical tooth weak, reduced to a triangular cell. Median tooth with 3 tiers of cells, one with 3 cells, the others with one cell only. Papilla hyaline, spheric, below the median tooth, 20 » in diameter. Male inflorescence lateral with 2 tiers of fertile bracts, 0.3 mm high. Cololejeunea hispidissima (St.) P. Tixier comb. nov. (Fig. IV). Small plant, light green, applied on substrate. Stem to 0.5 cm or more, 0.07 mm thick, with leaves 1.2 mm wide. Rhizoids hyaline. Leaves spaced at 0.25 mm. Lobe oval, trapeziform, 0.5 mm long, 0.4 mm wide. Basal cells making a short pseudo-vitta, cells 20-70 » long, 10-35 » wide with a slight papilla. Cells of the edge more or less isodiametric, with thin walls, 10-20 » x 10-16 m with cylinder-shaped papillae. Fig. IV. Cololejeunea hispidissima (St.) Tx. 1: Stem; 2: Leaf; 3: Basal part of the lobe; 4: Edge of the lobe; 5: Young lobule; 6 & 7: Mature lobule; 8: Propagule. Mount Maquiling Bryoflora 147 Lobule inflated, rounded, 0.25 mm long, 0.25 mm wide, keel without papillae. Apical tooth present on the young leaf, missing on the mature ones. Median tooth with 2 cells, young erect, repelled on the inferior edge. Hyaline papillae spheric, 10 » in diameter, under the first cell of median tooth. Fugaceous stylus 16 » high and 10 » wide. Propagules discoidal with 20 cells and 65 » in diameter. Fructification unknown. Cololejeunea haskarliana (G.) Bx. var. luzonensis var. nov. (Fig. V). Planta dioica, parva, pallida, ad substratum appressa. Caules 0.5 cm longi, 0.07 mm crassi, cum foliis 1.1 mm lati. Rhizoidea hyalina. Folio inter se 0.2 mm distantia. Lobus subtriangularis, 0.5 mm longus, 0.4 mm latus. Cellulae basales in pseudo-vitta 30-40 » longae, 30-15 p» latae, cum papilla crassa usque ad 25 p altae, marginales parvae, 10-15 » in diametro, papillosae. Lobulus ovalis 0.2 mm longus, 0.15 mm latus, carina papillosa. Dens apicalis unicellularis, dens medinus cellula arcuata, latere externo lobuli locata. Una cellula e dente medio usque ad marginem posticum. Papilla hyalina subsphaerica, 10 m in diametro. Stylus bicellularis, 40 » altus, 10 » latus. Propaguli disciformes, in statu 12 cellulares, 50 » in diam. Capitula feminea 0.4 mm alta. Bracteae florales cum lobo 0.4 mm longo, 0.15 mm lato, et lobulo 0.3 mm longo, 0.10 mm lato. Perianthia 0.3 mm alta, 0.2 mm lata. Dioic plant, small, light green, applied on substrate. Stem 0.5 cm _ long, 0.07 mm thick, with leaves of 1.1 mm wide. Rhizoids hyaline. Leaves spaced on stem at 0.2 mm. Lobe subtriangular, 0.5 mm long, 0.4 mm wide. Basal cells making a pseudo vitta, cells 30-40 p» long and 30-15 » wide with a strong papilla Fig. V. Cololejeunea haskarliana Bx var. luzonensis var. nov. I: Stem; 2: Leaf; 3: Basal part of the lobe; 4: Edge of the lobe; 5: Young lobule; 6-8: Lobule; 9: Propagule; 10: Perianth; 77: Lobule of the perianthal bract. 148 Gardens’ Bulletin, Singapore — XXVI (1972) up to 25 » tall. Cells of the edge, small, more or less isodiametric, 10-15 » in diameter, papillose on the edge of leaf. Lobule oval, 0.2 mm long, 0.15 mm wide papillose on the keel. Apical tooth unicellular, median tooth of one, arc shaped cell lying on the external face of lobule. One cell between the median tooth and the keel. Papillae spheric, hyaline, 10 » in diameter, stylus bicellular, 40 p» high, 10 » wide. Young (?) propagules, 50 » in diameter and with 12 cells. Female inflorescence 0.4 mm high. Perianthal bracts with a lobe 0.4 mm long, 0.15 mm wide and the lobule 0.3 mm long and 0.10 mm wide. Perianth shorter than perianthal bracts, piriform, 0.3 mm high, 0.2 mm wide. Cololejeunea haskarliana (G.) Bx. var. thermarum var. nov. (Fig. VI). Planta dioica, parva, pallide virens, ad substratum appressa. Caules usque 0.5 cm longi, 0.07 mm crassi, cum foliis 1 mm lati. Rhizoidea hyalina. Folia inter se 0.2 mm distantia. Lobus subtriangularis, 0.4 mm longus, 0.4 mm latus. Cellulae basales in pseudo-vitta, 30-60 m» longae, 15-30 yw latae cum papilla humili (10 p» alta): marginales parvae, 10-20 » X 10-15 yw metientes. Papillae per tutam marginem. Dens apicalis unicellularis, triangularis, pariete hyalinis. Dens medinus unicellularis, rotundatus cum dente medinus decussatus: 1 cellula unter dentem medianum et marginem posticum. Papilla hyalina subsphaerica, 25 » in diametro. Stylus unicellularis, 30 p» altus, 10 » latus. Capitula feminea 0.5 mm alta. Bracteae florales cum lobo 0.5 mm longo, 0.25 mm lato, et lobulo 0.3 mm longo, 0.15 mm lato. Perianthia piriformia, inflata, 4 plicata, 0.5 mm alta, 0.3 mm lata. Fig. VI. Cololejeunea haskarliana Bx var. thermarum vat. nov. 1: Stem; 2: Leaf; 3: Basal part of the lobe; 4: Edge of the lobe; 5: Young lobule; 6 & 7: Lobule; 8: Perianth; 9; Lobule of the perianthal bract. Mount Maquiling Bryoflora 149 Small dioic plant, light green, applied on substrate. Stem to 0.5 mm _ long, 0.07 mm thick, with leaves 1 mm wide. Rhizoids hyaline. Leaves spaced on stem at 0.2 mm. Lobe subtriangular, 0.4 mm long and 0.4 mm wide. Basal cells making a pseudo-vitta, cells 30-60 » long and 15-30 » wide with a low papilla 10 » high. Edge cells small, more or less isodiametric, 10-20 » long, 10-15 » wide. Edge papillose on almost all the periphery. Oval lobule, keel lightly papillose, 0.2 mm long, 0.15 mm wide. Apical tooth triangular with hyaline walls. Median tooth unicellular, crossed with apical tooth. One cell between the median tooth and the edge. Papillae hyaline, spheric, 25 » in diameter. Stylus unicellular, 30 » high, 10 » wide. Female inflorescence 0.5 mm long, perianthal bracts with a lobe 0.5 mm long, 0.25 mm wide and a lobule 0.3 mm long, 0.15 mm wide. Perianth piriform, inflated, with 4 folds, 0.5 mm high, 0.3 mm wide. Near to the preceding species, differing by the ornamentation of leaf cells, the disposition of the lobule teeth, the shape of the papillae and the perianthal bracts. Cclolejeunea selaginellicola sp. nov. (Fig. VII). Planta dioica, parva, pallide virens, ad substratum appressa. Caules usque ad 0.6 cm longi, 0.10 mm crassi, cum foliis 1 mm lati. Rhizoidea hyalina. Folia 0.2 mm inter se distantia. Lobus ovalis, 0.6 mm longus, 0.25 mm latus. Cellulae basales 30-60 » longae, 15-20 » latae, papillosae; marginales, pariete incrassatae rotundatae 10-15 mw in diametro, cum papillis altis conicis. Lobulus magnus 0.25 mm longus, 0.15 mm latus, papillosus usque ad + latitudinis. Margo superior lobuli cum uniseriatis cellulis hyalinais dens apicalis magnus, unicellularis, trian- gularis, parietibus hyalinis. Dens medianus unicellularis, sphaericus, 2 cellulis usque ad marginem posticum. Papilla hyalina parva sphaerica 7-10 p» in diametro. Stylus tricellularis, 40 » longus, 10 » latus. Capitula feminea 0.8 mm alta. Bracteae cum lobo 0.6 mm alto, 0.25 mm lato, et lobulo 0.45 mm longo, 0.08 mm lato, apice acuminato. Perianthia piriformia, 4 plicata, rostro notato, 0.8 mm_ alta, 0.5 mm lata. Dioic, small species, light green, applied on substrate. Stem to 0.5 cm long and more, 0.10 mm thick, with leaves 1 mm wide. Rhizoids hyaline. Leaves spaced on stem at 0.2 mm. Lobe oval, 0.6 mm long, 0.25 mm wide. Basal cells 30-60 » long, 15-20 » wide, papillose. Lobule large, 0.25 mm long, 0.15 mm wide with an angle between the keel and the edge of the leaf. Lobule papillose, up to 4 its width. Superior edge of lobule with a row of hyaline cells finishing with an apical, unicellular tooth, triangular, large, thin-walled. Median tooth reduced to a spheric cell; 2 cells between the median tooth and the edge. Papillae hyaline, small, spheric, 7-10 » in diameter. Stylus tricellular, 40 » long 10 p» wide. Female inflorescence 0.8 mm high. Perianthal bracts with a lobe 0.6 mm long, 0.25 mm wide and a lobule 0.45 mm long, 0.08 mm wide with an acute top. Perianth piriform with 4 folds and a strong beak, 0.8 mm high, 0.5 mm wide. Cololejeunea armata sp. nov. (Fig. VIII). Planta dioica, parva, pallide virens, ad substratum appressa. Caules 0.5 cm, 0.06 mm crassi, cum foliis 0.6 mm lati. Folia 0.15 mm inter se distantia. Lobus Ovalis, magis aut minus arcuatus, 0.4 mm longus, 0.25 mm latus. Cellulae basales, pariete pauciter incrassatae, 20-30 p» longae, 10-17 » latae, papillosae; marginales rotundatae 10 » in diametro. Lobulus rotundatus 0.15 mm longus, 0.10 mm latus, papillosus praeter ad basin. Margo superior lobuli cum cellulis hyalinis uniseriatim dispositis usque ad dentem apicalem ipsam hyalinam. Dens medianus, bicellularis 150 Gardens’ Bulletin, Singapore — XXVI (1972) Fig. VII. Cololejeunea selaginellicola sp. nov. 1: Stem; 2: Edge of the lobe; 3: Basal cells of the lobe; 4: Lobule; 5: Young lobule; 6: Perianth; 7: Lobule of perianthal bract. Mount Maquiling Bryoflora se LS, ; Oy EK oS zoey ZO | ex SSS = OD OYE SOE 1004 1b EER — FOE an S. ea all Fig. VIII. Cololejeunea armata sp. nov I: Stem; 2: Edge of the lobe; 3: Basal cells of the lo be; 4: Lobule; 5: Propagules; 6: Perianth; 7: Perianthal bract; 8: Lobule of the perianthal bract. $52 Gardens’ Bulletin, Singapore — XXVI (1972) decussatus cum dente apicali. Papilla hyalina, parva, sphaerica, 10 » in diametro. Stylus cylindratus, 30 p» altus, 10 » latus. Propaguli disciformes, in statu 12 cellulares, 40 » diam. Capitula feminea magis aut minus lateralis, 0.4 mm alta. Bracteae cum lobo, 0.4 mm longo, 0.18 mm lato, et lobulo 0.3 mm longo, 0.10 mm lato, margine crenulato, Perianthia piriformia, rotundata, 4 plicata, 0.4 mm alta, 0.25 mm lata, rostro notato. Dioic, small plant, light green, applied on substrate. Stem to 0.5 cm, 0.06 mm thick, width with leaves 0.6 mm. Rhizoids hyaline. Leaves spaced on stem at 0.15 mm. Lobe oval, more or less arc shaped, 0.4 mm long and 0.25 wide. Basal cells with thick walls, 20-30 » long and 10-17 p» wide; papillose edge with isodiametric cells about 10 » in diameter, with a papilla low and cylindric. Lobule rounded, 0.15 mm long, 0.10 mm wide, papillose except in the basal part. Superior edge with a row of hyaline cells finishing in apical tooth cell, hyaline, thickwalled. Median tooth of 2 cells, crossed with the apical one. The apical border of lobule from apical tooth to keel arc shaped. Papillae hyaline, small, spheric 10 p» in diameter. Stylus cylindric, 30 » high, 10 » wide. Propagules, 40 » in diameter and with 12 cells. Female inflorescence on the lateral side, 0.4 mm high. Perianthal bracts with a lobe 0.4 mm long, 0.18 mm wide and a lobule 0.3 mm long and 0.10 mm wide, the superior edge of the lobule crenulated. Piriform perianth, rounded with 4 folds 0.4 mm high, 0.25 mm wide with apparent beak. IV CONCLUSIONS This study devoted to few species shows the necessity of serious reflexion because of the complexity of Cololejeunea bryoflorula. 1. Interest of these collections The object of this work is to find out whether these gatherings bring an interesting contribution to the knowledge for Bryology for Mt. Maquiling. Accord- ing to Bartram (1939) sixty species of Mosses were found at the station. My list contains 34 species, 16 of which are new for Mt. Maquiling and one for science. This contribution shows an increase of known mosses by 20 per cent. There arose a few inquiries on the different gatherings of liverworts. The Herzog’s list concerning Baker’s collections had 28 species. My own has 42, 34 of which are new and 5 new for science. For liverworts a 50 per cent increase is found. The two figures based on a one-day collection show that further collecting trips to Mt. Maquiling would be worthwhile. 2. Biogeographical connections For mosses of my list, the community floristic coefficients between Mt. Maquiling and the different countries of South East Asia is established as follows:— 40 Java, Sumatra, Thailand 30-40 Vietnam, Borneo, Cambodia, Celebes 30-20 Ceylon, Japan, China 20-10 Himalaya, Malaya, Hawaii, New Caledonia. The main axis of affinities goes from Japan to Sunda Straits. Some relations with Hawaii and New Caledonia still remain puzzling. Mount Maquiling Bryoflora 153 LITERATURE HeErRzoG, T. (1931): — Hepaticae philippinenses a Cl. C. Baker lectae. Ann. Bryol., IX, 79-94. BARTRAM, E. B. (1939): — Mosses of the Philippines. Philip, J. Sc., 68, 1-424. TIxiER, P. (1966): — La végétation orophile de I’Asie du Sud Est. Le Mt. Maquiling et ses Epiphytes. — Science & Nature, no. 78, 3-11. (1969): — De Sematophyllaceis. I-Essai de revision des Clastobryaceae de Luzon, J. Hattori Bot. Lab., no. 32, 21-64. (1970): — Contribution a la connaissance du genre Cololejeunea en Asie due Sud Est. II/La section Radulae du sous genre Lasiolejeunea. Ann. Fac. Sci. Phnom Penh, no. 3, 174-190. A Pran f if | ee silat (aya ‘ ‘ ai ~ - a : i “4 > é i. » re ak Oe nid yt ae | . ae ‘4 na? > 9 é } _e% a A oe ve > * 455 aie G8, iii ea | >) 4h re ‘ J, %2 7teé nJ — he $6.55 ’ i ‘J | ' ’ a Tw ; (tu i My « rae t ip =i iste Si per cont ia ow a « glen r- pe re the er 7 ; ; we ik (i a rn \F sl nf : ‘ = ' i rer ae 7a a e 2. 3 , ryan + gia ‘- ‘ os i my y om 4 f reaaipe L New Cale Philippine Species of Nepenthes by SHIGEO KURATA* and Masami TOYOSHIMA#* This paper lists the results of two expeditions to the Philippines in search for Nepenthes. It records seven species and describes one of them and three hybrids as new taxa. All but one were collected during the three-week expedition made by both authors to the islands of Mindanao and Luzon in August 1965. N. burkei was collected by the senior author (S. Kurata) when he participated in the second expedition from December 1967 to February 1968 to Mindoro Island. Some Bornean species were also found but are excluded in this paper. The authors are grateful to Dr. C. X. Furtado for correcting the latin diagnosis. The new taxa are N. globamphora, N. mirabilis x alata, N. petiolata x alata and N. truncata x petiolata. 1. Nepenthes alata Blanco, Fl. Filip., ed. 1, p. 805 (1837). LUZON: Mountain Prov., Baguio, 1470 m. alt., Aug. 28, 1965, S. Kurata 1136-a 1136-b; en route to La Trinidad, 1500 m. alt., Aug. 28, 1965, S. Kurata 1135. MINDANO: Surigao del Sur, eastern slope of Mt. Legaspi, 270 m. alt., Aug. 19, 1965, S. Kurata 1101-a, 1101-b, 1104. Distribution. Philippines, Malaya, Sumatra, Borneo. Between the Luzon plants and those from Mindanao there are some differen- ces. The former are hairy on the pitchers and tendrils. The purple blotches or stripes are usually distributed on its pitcher wall. On the other hand, the pitcher of the latter is strikingly narrow but dilate in the lower part, and the hair is absent from its wall. The colour of the pitcher is mostly yellowish green and at times becoming brownish green near the mouth. 2. Nepenthes burkei Masters. Gard. Chron. Ser. 3, VI p. 492 (1889). MINDORO: Occidental Mindoro, Mt. Kidayap, summit, 1550 m. alt., Feb. 12, 1968, S. Kurata 2033, 2034, 2035. Distribution. Philippines (Mindoro and Panay). 3. Nepenthes (Montanae) globamphora Kurata et Toyoshima, sp. nov. (Fig. 1, pl. 1). — Nepenthes globamphora (nomen nudum), Kurata et Toyoshima, Journ. Insectivorous Plants Soc. 36: p. 15-22, 1966. Planta tenuis, breviter scandens, luteo-viridis. Caulis 30-80 cm altus, 4-5 mm crassus, cylindricus. Folia 8-12 cm longa, 1-1.8 cm lata, lanceolata, coriacea, sessilia, basi 4 amplexicaulia, utrinque glabra; nervi longitudinales utrinque 3; nervi transversales obscure, oblique ascendentes; cirrhus 15-20 cm longus, cylind- ricus, glaber, luteo-viridis saepe etiam lucido-purpureus. Ascidia 4-9 cm longa, 3-5 cm lata, subdimorphia, inferiora ellipsoidea vel globosa, superiora ampulliformia, omnia luteo-vel, rubro-viridia, diffuse purpureo-maculata, coriacea, glabra, intus omnino glandulosa, bialata, alis margine dentatis, dentibus 3-ciliatis; os orbiculare obliquum: peristomium 5-8 mm latum, sub-cylindricum, aequale, luteo-viride, margine exteriore revolutum, interiore denticulatum; operculum 2-3.5 cm latum, * Insectivorous Plants Society, Department of Biology, Nippon Dental College, Fujimi, Tokyo, Japan. 155 156 Gardens’ Bulletin, Singapore — XXVI (1972) Fig. 1. Nepenthes globamphora Kurata et Toyoshima: A. male plant with flowers and pitchers (x 0.3); B. upper portion of female plant with flowers (x 0.3); C. male flower (x 2); D. capsules (x 1). her on the lower stem (x 2.5). a pitc Nepenthes globamphora PLATE 1 al al ORNL RN LOR i tac intl Mab tabs Philippine Species of Nepenthes 157 cordato-orbicularum; calar ca. 1 mm longum, filiforme. Inflorescentia 10-15 cm longa, racemosa, pedunculus 6-9 cm longus, ferruginio-tomentosus, pedicelli 3-4 mm longi, uniflori; sepala 4, ovata, 2-2.5 mm longa, 1-1.2 mm lata, extus minute tomentosa; columna staminae sepalis brevior, glaber, antherae 8, uniseriatae; cap- sula 17-20 mm longa, 4 mm lata, fusiformis, brunneo-nitida, glabra, valvae lanceo- latae. MINDANAO: Surigao del Sur, eastern slope of Mt. Legaspi, 270 m alt., Aug. 22, 1965, S. Kurata and M. Toyoshima 1128 holotype in Herb. NDC (Nippon Dental College) 1129, 1130, 1132. This new species is easily distinguishable from all others by the peculiar pitcher which has an unusual roundness (hence the epithet “‘globamphora’’), and by the trifid ciliae fringing the pair of wide wings — a very unique occurrence in the genus. The lid has no appendix on its lower surface where nectar glands are sparingly distributed. The glands are very small and deep. The remarkable feature of this species is that the inner surface of the pitcher is wholly glandular. When dry the specimen is red-brown, and the lower surface of leaves and flowers are more reddish. The lid has a ceramic luster on its lower surface. This plant is comparati- vely common on the eastern slope of Mt. Legaspi, but its distribution seems limited to a narrow area. 4. Nepenthes merrilliana Macf., Contrib. Bot. Lab. Univ. Pennsylv., III, mecu7..t. (1911). MINDANAO: Surigao del Sur, Camp David (a lumberer’s camp, 10 miles upper stream of the mouth of Carrascal river). 150 m alt., Aug. 20, 1965, S. Kurata 1117, 1118, 1119, 1120-a, 1120-b, 1124, 1125, 1126; eastern slope of Mt. Legaspi, mo m alt.. Aug. 19, 1965, S. Kurata 1116, 1121, 1122, 1123, 1127. Distribution: Philippines (Mindanao), Sulawesi. 5. Nepenthes mirabilis Druce, Rept. Exch. Cl. Br. Isl., 1916, p. 637 (1917). MINDANAO: Surigao del Sur, on a river side near the Carrascal Bay, 20 m alt., Aug. 19, 1965, S. Kurata 1107. Distribution: Southern China, Indochina, Malaya, Philippines (Mindanao), Borneo, Java, Sumatra, Sulawesi, New Guinea, Parau, Australia (York Pen.). 6. Nepenthes mirabilis x alata, hybr. nov. Hybrida naturalis inter N. mirabilis et N. alata. Folia lanceolata, herbacea, glabra, petiolata. Ascidia 10-15 cm longa, 3-4 cm lata, monomorphia, cylindrica, bialata, alis fimbriatis, luteo-viridia, herbacea, glabra; os orbiculare obliquum: peristomium 2-3 mm latum, subcylindricum, aequale; operculum 3-5 cm iatum, orbicularum, facie inferiore prope basin appendiculatum; calcar ca. 1 cm longum, bifidum. Inflorescentia ignota. MINDANAO: Surigao del Sur, on a river side near the Carrascal Bay, 20 m alt., Aug. 9, 1965, S. Kurata 1111-a (type in Herb. NDC), /1/1-b. This specimen has thin, herbaceous, petiolate leaves and cylindrical pitchers with orbicular lids. The pitcher is intermediate between its parental species; the strong folds on the boundary between the waxy zone and the glandular zone are derived from N. mirabilis whilst the appendix on the lower surface of the lid agrees with N. alata. 158 Gardens’ Bulletin, Singapore — XXVI (1972) 7. Nepenthes petiolata Danser, Bull. Jard. Bot. Buitzg. ser. III, Vol. IX, p. 353 (1928). MINDANAO: Surigao del Sur, eastern slope of Mt. Legaspi, 270 m alt., Aug. 19, 1965, S. Kurata 1108. Distribution. Philippines (Mindanao). 8. Nepenthes petiolata x alata, hybr. nov. Hybrida naturalis inter N. petiolata et N. alata. Folia elliptico-lanceolata, coriacea, glabra, petiolata. Ascidia 10-15 cm longa, 3-4 cm lata, monomorphia, cylindrica, bialata alis fimbriatis, luteo-viridia, diffuse purpureo-maculata, coriacea, minute tomentosa; os orbiculare obliquum; peristomium ca. 2 mm latum, sub- cylindricum, aequale; operculum 3 cm latum, ovatum, facie inferiore prope basin appendiculatum; calcar ca. 1 cm longum, filiforme. Inflorescentia ignota. MINDANAO: Surigao del Sur, eastern slope of Mt. Legaspi, 270 m alt., Aug. 19, 1965, S. Kurata 1113-a (type in Herb. NDC), J1/3-b. This specimen has elliptical leaves with petioles and cylindrical pitchers with ovate lids. The colour of the pitcher is yellow-green with purple blotches and/or stripes and is densely covered with short hairs which seem to be the same of that of N. petiolata. On the other hand, the small glands and the appendix on the lower surface of the lid seem to be derived from N. alata. A lustrous belt on the boundary between the waxy zone and the glandular zone on the inner surface of the pitcher is also from N. alata. Both species N. petiolata and N. alata are found growing in the same area. 9. Nepenthes truncata Macf., Contrib. Bot. Lab. Univ. Pennsylv., II, p. 209, t. IT (1911). MINDANAO: Surigao del Sur, Camp David, 150-300 m alt., Aug. 20, 1965, S. Kurata 1105, 1106, 1115. Distribution. Philippines (Mindanao). 10. Nepenthes truncata x petiolata, hybr. nov. Hybrida naturalis inter N. truncata et N. petiolata. Folia oblonga, apice truncata, coriacea, supra glabra, subtus villosa, petiolata. Ascidia 10-15 cm longa, 3-4 cm lata, monomorphia, inferne ventricosa, superne cylindrica, bialata, alis fimbriatis, luteo-viridia, diffuse purpureo-maculata, coriacea, tomentosa; os orbi- culare obliquum; peristomium 5-8 mm latum, subcylindricum, margine exteriore expansum, margine interiore denticulatum; operculum 4-5 cm latum, cordato- ovatum, facie inferiore prope basin appendiculatum; calcar ca. 1 cm longum, bifidum. Inflorescentia ignota. MINDANAO: Surigao del Sur, eastern slope of Mt. Legaspi, 270 m. alt., Aug. 22, 1965, S. Kurata 1109-a (type in Herb. NDC), 1109-5. This specimen has wide coriaceous leaves with petioles, and the lamina is cordate at the apex. The petiole and lamina are densely covered with brownish coarse hairs beneath. The pitcher has a wide and expanded peristome. The lid has on its lower surface an appendix and two kinds of nectar glands: one small and deep, surrounded with a wide rim, the other large and shallow, surrounded with a narrow rim. The above characters show that this specimen is a natural hybrid between N. truncata and N. petiolata. At the place where this plant was collected, both parental species and the other i.e. N. alata were growing together. Studies in the basidium Spore-spacing and the Boletus spore by FE. J. H. CORNER, Botany School, University of Cambridge A geometrical analysis of the 4-spored homobasidium is made on the assumption that spore-spacing is determined by sterigmatic spacing on the basidium-apex. Given, then, the values of spore-width and the interval between adjacent spores of the tetrad, other details of the basidium-unit can be worked out except the lengths of the spores, sterigmata and basidium. The relations between the parts are summarised as equations; their values in ratios of spore- width are tabulated in an appendix. Observations on Mycena illuminans with subglobose spores are given as a check on the theory. The smooth Boletus-spore is shown to be the unornamented endospore of a subglobose spore with endosporic ornamentation that has suffered compression and reduced spacing through the adaxial displacement of the sterigmata. The spores of Strobilomyces and Phyllo- boletellus appear as the more primitive. The use of this theory is shown by analysis of the cruciform spores of Marasmius nigricans. During the tumultuous years 1942-1945, when Dr Kwan Koriba maintained the scientific status of the Singapore Botanic Gardens, I was permitted to work with him in the upstairs laboratory of the office. We studied trees, their seasons and their reproduction, and I was allowed a more personal investigation, which had long intrigued me, into the construction of the basidiomycete hymenium. Some of this work has been published (Corner 1947, 1948). It has shown that the lengths and widths of basidia, spores and cystidia are bound by specific constants which have generic trends. The dimensions of these parts are usually given inde- pendently in taxonomic descriptions, but they are related in consequence of the characteristic of the hypha that produces them. The relations can be displayed as sporographs, basidiographs and so on, as explained in those papers, but their interconnection lies in the dependence of spore-width on that of the basidium. This is the matter that I now take up, and it becomes the problem of spore-spacing on the basidium. With many detailed observations for such genera as Amanita, Lepiota, Hygro- phorus, Marasmius, Russula, Entoloma, Agaricus and Coprinus, I had intended an extensive survey but the long hours necessary for such a work have never recurred. I take it up for the special case of Boletus because in the course of my account of the rich Boletus-flora of Malaysia (Corner 1972) the problem has loomed continually and is fundamental to the classification of the species by means of their distinctive spores. My thesis is the geometrical analysis of the position of the spores on the basidium. It is not an explanation of their arrangement by physical means but a guide to understanding how the spore grows to its full size and shape and, indeed, how it may develop its superficial structure. I have set forth this analysis in the simplest terms, step by step, so that it may be followed by mycologists who, like myself, may have lost touch with advanced mathematics. 159 160 Gardens’ Bulletin, Singapore — X XVI (1972) Most basidiomycetes have four spores on the basidium. There have been several accounts of the development of individual spores (Corner 1948, 1968; Malencon 1958; Perreau-Bertrand 1967). I propose to consider the tetrad as a unit. The four spores are regularly spaced. Their arrangement can be affected by adjacent basidia. The spacing is affected by the contour of the hymenium. Spore-shape results from the way in which spore-volume is contained in this spacing. A prime factor is the width of the basidium. Yet the tetrad of one basidium does not contact that of another; the spores of a tetrad are not in contact; they are separated from the basidium by sterigmata; maturing basidia are separated from each other by immature. To secure this spacing there must be action at a distance. The force appears to be electrostatic repulsion at free surfaces; the whole mechanism collapses when wetted. To prove this, I had intended to grow basidia in an induced electric field, but this has not materialised. I hope, however, that this article will Open a new line of enquiry from which some real understanding of the precision of the hymenium will come and better appreciation of the basidiocarp which houses it. The toadstool is essentially an umbrella opened geotropically. Method The sporing basidium is a delicate turgid structure that readily shrinks in contact with a fixative. As my object was to make fine measurements of spore-size and the intervals between spores on the basidium, it was necessary to examine undistorted living basidia under high enough magnification to make reasonably large camera lucida drawings. I first used dry mounts of thin strips of hymenium for surface-view and moderately thick sections for side-view, and observed them with an oil-immersion lens. Condensation on the coverslip, however, in the high humidity of Singapore brought failure and I had to resort to aqueous mounts. Thus I learnt that sooner or later, after ten minutes with some species or an hour or two with others, the apex of the wetted basidium collapsed and the sterigmata inclined, even jumped, inwards; to be effective the basidium had to be superficially dry. By dint of perseverance I managed to obtain for subsequent analysis many accurate camera lucida drawings of a great variety of agarics, boleti, polypores, stereums and clavarias. The present article merely summarises these results because names are not available for most of the fungi, and the labour is great; about a thousand measurements are needed for accuracy in any one species. The two aspects of the basidium, end-view and side-view, need to be combined. From this a geometry of the basidium emerges to relate the shape of the basidium- apex via the sterigmata with the spore-spacing and, thus, with spore-shape. As the spores are borne at the same level above the basidium (though there are exceptions), one can obtain in end-view an optical section of the tetrad. Focussing down, one can see the basidium-outline in its optical section at maximum width and, on then focussing slightly upwards, even the tips of the sterigmata. However, it is easier to see these tips on discharged basidia before they have collapsed. The four spores, as seen in end-view, occupy the corners of a square and one can imagine them to be confined by two spore-circles, an outer circumscribing circle S’ (Figure 3) and an inner inscribing circle $’”’, to which may be added a third spore- circle S” passing through the centres of the spores. Then there is the real basidium- circle as the outline of the basidium, which I call w, and for the sterigmata one Studies in the basidium 161 can imagine a sterigmatic circle M passing through the tips of the sterigmata. The circles are concentred on the long axis of the basidium. S’ exceeds the basidium circle which can often be seen to lie within S$”; the M circle lies slightly outside or abaxial from $”’. These real and imaginary circles, derived from end-view of the basidium-unit, can be projected on to the basidium apex, as in Figure 3. Since a basidium cannot be examined both in end-view and side-view, an average must be taken from a set of measures for each. In end-view basidia are often seen to be compressed and ellipsoid in optical section. Such can be avoided but they cannot be recognised in side-view, for which the average is necessary. The figures in the following pages are based on such averages from sets of twenty to one hundred drawings. Following the system adopted in my previous papers, I continue with this notation: — nis the spore-number per basidium. D is spore-length. d is spore-width; d rad, when measured in side-view, and d tan, when measured tangentially or at right angles to d rad. is the ratio D/d. is the length of the basidium. “Se w is the maximum width of the basidium. M is the diameter of the sterigmatic circle. m is the diameter of the sterigmatic patch circle. S’ is the diameter of the outer spore-circle. S” is the diameter of the middle spore-circle. S’” is the diameter of the inner spore-circle. (S’), (S”), (S”’) and (m) refer to the inner spore or endospore when needed to be distinguished. o is the ratio of spore-interval to d. 8 is the spore-angle. 5 is the angle subtended by the half-spore on the spore-circle. «: is the angle subtended by the sterigmatic patch at the centre of the curved apex of the basidium. Evidence of spore-spacing The following considerations supply the evidence that the spores of a tetrad are regularly spaced. Elongate spores. A tetrad of elongate spores in end-view appears as four circles equidistant round the basidium-apex (Figure 1; Buller 1924, fig. 84-86, 91; Corner 1964 p. 234). The long axes of the spores are parallel with that of the basidium; the spores face this axis and are not inclined. This is the main evidence for supposing the imaginary circles S’ and S”’ and the lateral spacing factor ‘ o’. 162 Gardens’ Bulletin, Singapore — XXVI (1972) 20 Oa ZS, OG Ff GAL 500 Figure 1. Spore-tetrads in end-view, x 800 (a. x 500); a, Oudemansiella; b, f, Amanita; c, Pluteus; d, Panaeolus; e, Psathyra; g, Boletus ravenelii, with two normal basidia and iwo pairs of approximated basidia having their eight spores set in an ellipse. Interfering tetrads. Usually a basidium develops its tetrad freely at some distance from other sporing basidia. New basidia are intercalated and take the place of those that have discharged their spores and become ineffective. Occasion- ally two basidia mature so close together that their tetrads are distorted and, as seen in end-view, their eight spores are combined into one imaginary ellipse (Figure 1). This re-arrangement shows that sporing basidia act as spacing units and influence their neighbours. Where two S’ circles approach or would intersect the adjacent spores of each tetrad are repulsed outwards. The unit has a boundary at S’. The adaxial patch. In many verrucose or echinulate spores, for which Russula and Thelephora are good examples (Malencon 1958; Corner 1968), the warts or spines develop all over the spore except in a small area on the adaxial side of the spore just above the apiculus. I call this smooth area the adaxial patch. It is the part of the spore impinging on the inner spore-circle S”’ and it shows that this imaginary circle is a boundary defining a central ‘dead space’ over the basidium- apex where spore-growth is forbidden. It is the sharpest boundary in the tetrad. Tangentially compressed spores. The tetrads of Panaeolus (Figure 1) and some species of Coprinus (Buller 1922, fig. 104; 1931, fig. 29-32), show radial or adaxial compression of the spores; d rad is less than d tan. In contrast with globose spores that are strictly confined by the one spacing given by S’ and S”’, they bulge laterally; a lateral spacing has to be considered as well as a radial. More spectacular are the trigonous and cruciform spores in various agarics (Figure 12). Evidently there may be variation in the lateral spacing and in the intensity of inhibition from S’ (as shown by abaxially echinulate spores), while that of S”’ is rigid. Here, however, the exception comes with the Boletus-spore. Studies in the basidium 163 Reversed spores. Tetrads of homobasidiomycetes do not bear reversed spores; that is, spores do not develop with the apiculus turned abaxially. This may happen with diads in normally tetrasporous species (Corner 1950, fig. 271). In such cases the two spores are evidently so far apart that the sterigmatic spacing by the sterigmatic disc is ineffective; nevertheless, S’ and S”’ operate from the basidium- apex to maintain the spore-shape, even if o is also ineffective. 6-8-spored basidia, When n is 3, the spores in end-view fit the angles of an equilateral triangle; when n is 4, they fit a square; when n is 5, they fit a regular pentagon. But, when n is 6 or 8, they do not fit a regular hexagon or octagon; they are disposed in an ellipse, as with interfering tetrads. This ellipse, however, seems not to be plane but to have decurved ends. Four spores occupy the central part of the ellipse and one or two spores at each end lie at a slightly lower level through the obliqueness of their sterigmata. The same effect may be seen in the tetrads of Russula and Lactarius. | have not succeeded in examining such hexads and octads in any detail. The obvious material is to be found in Cantharellus, Craterellus and Pterygellus (Corner 1966), but the basidia of these fungi are too long for suitable study in end-view. The regular disposition of the spores proves, nevertheless, that spacing factors operate through the ring of 6-8 sterigmata placed peripherally at the basidium-apex with its superimposed dead space. Towards the geometry of the basidium Certain features of the basidium-unit need more explanation. Positioning of the spores. They are positioned by the sterigmata. These out- growths, at first normal to the curved apex of the basidium, become parallel with the basidium-axis and develop at their attenuate tips the sterigmatic discs. From these the spores, inhibited by S$”, develop abaxially to the limit of S’ and then elongate within these limits, parallel to the basidium-axis, if they are to become larger. The transverse sterigmatic discs appear to be the initial spacers and the apiculus of the mature spore is the adaxial feeler which indicates, but seldom coincides with, S’”. Careful measurement of the spore- and sterigmatic circles have shown me that, as a general rule, M= S” — 0.7d; for a single spore, that is, the tip of the sterigma is situated external to S”’ by a distance of 0.15d. If spore-spacing is determined by sterigmatic spacing, the cause of this cannot be discovered by simple observation because it must lie within the basidium. It must be connected with the hyaline cap which forms at the apex of the basidium shortly before the sterigmata emerge (Corner 1948). The substance of this cap forms, evidently, the walls of the sterigmata and those of the spore-rudiments; presumably it becomes electrically charged at the surface. It must be partitioned into four blocks which underlie the sterigmatic patches, or bases of the sterigmata (Corner 1948). I have pursued the geometrical consequences of the projection of this patch on the plane of the spore-circles and its hypothetical function in spore- Spacing (p. 169). Spore-angle. The direction in which the spore grows from the transverse sterigmatic disc was studied by Buller. He concluded that this angle was at 45° to the long axis of the basidium. I have measured the angle on camera lucida drawings and, though it is impossible to be precise with such minute structures, I agree with Buller’s conclusion. It is supported by the following argument. 164 Gardens’ Bulletin, Singapore — XXVI (1972) With reference to Figure 3, ca MTC is the spore-angle 0. If this is 45°, then MO=7C tin Gr= og sin 45° = 0.35d ~ or, for the whole basidium-unit, M = S” —2MC = S” — 0.70d This is the value of M found by direct measurement of S”’, S’ and M. The longitudinal plane of S”’ lies adaxially to that of M by this small fraction 0.15d. The ratio d/w. The positioning of the sterigmata and that of the spores about the longitudinal axis of the basidium so that the basidium circle (in end-view of the tetrad) lies within S” imply that there is a close relation between spore- width d and basidium-width w. These are two easy measures and their relation is an important feature of the basidium-unit. For the smooth ellipsoid spores of many homobasidiomycetes from Amanita to Clavaria, I find that in general d = 0.6 to 0.65w. Large globose spores may increase the ratio to 0.8w and narrow spores may decrease it to 0.5w. These differences may occur within a genus; the latitude in positioning of S’ may be the explanation, but specific differences in spore-width generally imply differences in the valve of w. Thus, a basidium 8 wide will bear spores 5u wide; one 10» wide will bear spores 6.34 wide. Despite records to the contrary, I have not seen a tetrasporous homobasidiomycete in which -d equals or exceeds w. The exception among the disporous is Clavulina with d = 1.18w (Corner 1948). No species has basidia of constant width. There is always some variation. Hence I have used as values of w and d either the mean of their variation or, when there have been sufficient measurements, their averages. Lateral spacing in the tetrad. As a measure of this feature, I take the ratio ~ of the spore-interval to spore-width, as seen on the spore-circle. In Figure 3 KL WAKE oS DS 2CK d Measurements of the tetrads of various basidiomycetes with isodiametric spores (d rad equals d tan) show that in general o = 0.5. The spacing can also be expressed by the angle 6 (Figure 3), as explained on p. 169. The basidium-apex. The projection of the spore-circles and other items of the basidium-unit on to the basidium-apex in median longitudinal section implies knowledge of the curvature of the apex, but this is a difficult matter and I can give only an approximation. The clavate basidium, fully grown but without sterigmata, has a hemispherical apex. When the hyaline cap forms, the apex protrudes slightly and in the long basidia of agarics with polymorphic basidia (such as Coprinus) this projection is lengthened and becomes almost cylindric with hemispheric apex. The question is whether it is ellipsoid or paraboloid. To test this, I measured the diameter of the basidia with hyaline caps, yet without sterigmata, at certain levels. These were the level of maximum width w and those at 4, 4 and ? of the vertical distance between the apex: of the basidium and the plane of w. I compared these results with those that can be calculated for similar chords of a sphere (or ellipse) — and a parabola, as shown in Table 1. Studies in the basidium 165 Table 1. Diameter of the basidium-apex, as ratios of w, at successive levels from the apex. Levels w 0.25w 0.5w 0.75w ellipse 1 0.661 0.866 0.968 observed 1 0.687 0.854 0.954 parabola l 0.438 0.750 0.938 The apex is clearly ellipsoid but, as rounding off of a clavate apex with slight protrusion, it is probably ellipsoid with a small dome of spherical curvature. This is the dome between the sterigmatic bases that struts them and collapses when wetted. Globose spores. Longitudinal alignment cannot be discerned directly from such spores but, from their positioning on the sterigmata, they clearly obey the dead space defined by S$”. Presumably they have lateral spacing and external limits, defined at least by the total effect of the basidium-unit. Spore-length. A globose spore fills the space between the limits of S’ and S”’. If more protoplasm enters the spore, it elongates within these limits. For a given spore-volume, the closer these limits the longer the spore will be. But spores vary in the way that they grow into and fill this space. Some are globose from the first and then lengthen; others expand gradually as they lengthen and become pip-shaped, obovoid or clavate. None of these factors, however, has prime signi- ficance in spore-spacing, though the limits of S’ and S”’ explain why ellipsoid spores vary much in length though little in width, and conform to a sporograph- locus. Sterigmatic length. Since the sterigmata become parallel, their length has no immediate effect on spore-spacing. They may lengthen abnormally, however, and remove one or two spores of the tetrad from the spacing effect of the whole, and this may be usual in Tremellaceae, but it is exceptional in homobasidiomycetes and leads to the abortion of the spore. Many exceptional states occur in Gastero- mycetes with their variously disorganised basidium-mechanism. I have been unable to detect any relation between the length of the sterigmata and other features of the basidium-unit. The length seems to depend on the hydrostatic pressure in the basidium and to be related with spore-number. Thus in species with n =3 or 2 or 1, as variations on n= 4, the sterigmata are progressively longer. Sterigmatic length can, therefore, be neglected in basidium- geometry. Basidium-length. With any one kind of basidium as defined by its equation (Corner 1947) w depends on /, but 7 does not otherwise determine spore-spacing. In all species / varies, but the resulting differences in w are so slight that the shorter basidia bear spores of practically the same shape and size as the longer. Thus, in tetramorphic species of Coprinus, there are not four sizes and shapes of spores. Nevertheless, as always, there are exceptions. At least two species of Hygrophorus have normally basidia of two sizes, very different in length, and they bear spores of very different size (Corner 1936). For the purpose of spore- spacing, basidium-length can also be ignored. 166 Gardens’ Bulletin, Singapore — XXVI (1972) Figure 2. Diagrams of basidium-units, treated as cones, divergent in the convex hymenium (upper left), convergent in the concave hymenium (upper right), and divergent or paraboloid in the plane hymenium (lower figures). The basidium-unit, The whole basidium with its spores can be regarded as an inverted cone expanding from the base (inverted apex) to the value of S’. Within the cone the unit develops; outside the cone it restrains other basidia. The effect is well seen in clavarioid basidia. Their sterigmata often diverge slightly and their long spores may also diverge and appear in optical section on the spore- circle as short ellipses. I suppose that this divergence is connected with the convex curvature of the hymenium, developed on the outside of the cylindric or clavate fruit-body, and that this curvature makes the basidia slightly divergent (Figure 2). In the agaric hymenium the basidium-unit becomes paraboloid with sterigmata and spores parallel to the long axis of the basidium. But the poroid hymenium is concave, lining tubes; the basidia are convergent and their cone- effects interfere; the outer spore-circles of adjacent units conflict and the whole unit is distally compressed. This is the problem in Boletus, many species of which Studies in the basidium 167 have long and narrow spores with convex adaxial face, known as the boletoid spore. Superficially this spore may resemble a Clavaria-spore; actually it is developed in a more complicated manner because the sterigmatic circle is displaced inwards. The hydnoid hymenium resembles the clavarioid. Its basidia are often small and their divergence may explain the prevalence of globose or subglobose spores. The stereoid hymenium with plane extension resembles that of the agaric but the basidium-units are not strictly aligned because of the thickening hymenium; new basidia may project beyond the influence of the mature and have slightly divergent sterigmata. Asymmetric basidia. Very occasionally a basidium that has grown aslant develops a typical spore-circle asymmetrically on the exterior side of the apex. Whether or not this is the effect of light, gravity or some other external stimulus on the hyaline cap, the configuration proves that the sterigmatic arrangement is a structural unit. Measurements. I have relied on my own measurements from living material. The basidium-unit is an inflated, semi-rigid structure prone to collapse, and it is not certain that measurements from dried material or that fixed in alcohol- formalin agree exactly with those from the living. It is tempting to use the published data in descriptions, but I have avoided these not only because they often refer to preserved specimens but because it is clear that immature basidia are often measured and such will give misleading results. The geometry of the basidium The geometry, as I have analysed it, is set out in Figure 3. The upper part shows the spore-tetrad, as seen in optical section from the end-view of the basidium, together with the spore-circles S’, S$”, S’”, the basidium-circle w, the sterigmatic circle M, the square joining the spore-centres C, and the tangents from the long axis of the basidium A to the spores. Then there are shown four small circles m on the spore-radii; they are the orthogonal projections of the sterigmatic patches on to this transverse plane. The lower part of the figure shows the longitudinal geometry of the basidium-apex which is drawn as a hemisphere on the base ww of maximum width of the basidium. The right half of the basidium-apex shows the sterigmatic patch (arc Hw) and the angle Haw which it subtends; I call this angle « . The various circles of the upper figure are projected on to the base-line ww. The lower right-hand spore represents a globose spore in side-view as it would fit on to the sterigma which, for clarity, has been omitted. If the upper figure is rotated through 90°, it represents the spore-circles above the basidium. Many relations of the parts can be derived from this figure. I give merely ’ the more useful that can be reduced to the minimum data d and c , when n = 4. It must be noted that the symbols S’, 8’, S’’, M, w, d and m refer to diameters and not radii. 168 Gardens’ Bulletin, Singapore — X XVI (1972) = —-— fe =e se ww ee Se SF es ew ewe ee SP se se as 8 = . § “sf _< i ! 1 a 1 1 i I ! | | I ' ’ be ; : S' S.» feos Mefab Ai os oS" Mai \Gr S$} Figure 3. The geometry of the spore-circles in relation to that of the basidium-apex in median longitudinal section; constructed for ¢ = 0.5, w = 1.65 d. For explana- tion, see text. Studies in the basidium 169 err re Te GE GE) sees ttc ies boca iv wba reassess orca accsenecsacvdanscosesevess (1) because CC = CK + KL + LLC, and by definition KL = do, beetacate d d ae ee ge ot to) en eR SoS os ca oa'y ac nc aan vals gne wndoohewelin apie buyuiewcana ait (2) because S” = JAC and, as S” (diameter) is hypotenuse to the rt. < triangle with sides CC, then (2AC)2 = 2(CC)2 AC = Jae = J id “(1 + oy as from (1) Ns ES eI MR EI a aie cakes iw tah Wigs On ncenensuansstenassavsedencesevassenedsege (3) NY ccs cee icon adh sansa dace Mags vos esedl tiandhusddecdessiorionaees (4) Now it is convenient to turn these relations into functions of § . In Figure 2, P is the point where the tangent from A meets the spore. PC d 2 d 1 . sin § = —— = — x — = SN earas cum CECT ORE E TEE CEP CETT ETT TT SeT rere (5) eae Seer emcee fC +c + o) or cosecS = 2 (1 + o) d = SS POSED Sacra treanenndemiinasnnnt Oe cise Ssdvahbiuivedeos satus (6) sin 6 d d(1 + sin 8) S’ = 8S” + d = — + d = ————— =_ (4 Cosec§) ciccecccccccccccsecceeccecteceeeceecers (7) sin § sin 6 o XS =d{ 20 +0) +1] d d (1 — sin 8) S’”’ = §S” — d = — — d = — —— = d (cosec§ — 1) ooeceiccceccccccccceccceccceeeeeeees (8) sin § sin 6 or S”=d[ 2(1+o0)—-1] ane PO ek a aN Seah in las sawn awianghcdnnae sumqagle ass enepeee s (9) or M=d{[ /2 (1 +o) -0.7] cosec § = oe et eae ee ee eee. 0 a, (10) Sterigmatic patch hypothesis. I assume that the spacing of the sterigmatic patches provides that of the spores by transmission through the sterigmata. The symmetrical partition of the hyaline cap in the basidium-apex into four areas, set as far as apart as possible on the level of w, may be the physical basis. The geometrical consequence must be traced by the projection of the sterigmatic patch on to the transverse plane of the spore-circles. In Figure 3 any circle with centre on AC’C and inscribing AP will provide the spore-spacing, but the sterigmatic patch implies certain conditions. Its projection must lie within the basidium- circle; it must place the sterigmatic tips on the sterigmatic circle; and it must be related with w (that is the perpendicular ww in Figure 3). For convenience I use m’ for the diameter of the projected circle of the sterigmatic patch and denote the circle accordingly. The most likely position for the centre of m is C’ where AC cuts the S”’ circle (in the upper part of Figure 3). This m circle contacts the basidium-circle, as can be seen from the following argument with reference to Figure 4. 170 Gardens’ Bulletin, Singapore — XXVI (1972) A C B c oe WO) 5" Figure 4. Construction to prove that the m circle with centre C on the S” circle contacts the basidium circle w. For a circle with centre on AC and with AP as tangent, then C’ P’ = C’ B, as radii of this circle. C’P’ = AC’ sin 6, or in this case S’’ sin § d (1 — sin 6) sin § d (1 — sin 6) from (8) 2 2 sin § C’B = AB — AC’, or in this case = 4 (W = ore) d (1 — sin 4) Now S’”” = from (8) sin 6 m = S’’ sin 6 ex hypothesi w= 5S’ + m= 8S” + S’ smo = SS’ (1 + sino) Therefore, d (1 — sind§) (1+ sin§6) d (1 —sin 68) oO): ae oo Saeed Vee eS 2 sin§ 2 sind 2 sin§ C’B — = 1 — sin2§ — 1 + sin§ = sin6 — sin2§ d d (1 —sin8) : 2. Therefore C’B = C’P’ Studies in the basidium 171 The following equations may therefore be added ee NE PN le 5a Vol sib Si din'n aveca vce GAibsee die ol nen dk Cees Dek cies Sou naseareanas (11) iS ize sah ret i ae SCN ee SINR ir sa ca dda ced aernscannevr eves “Weastuondoecdioesteamteths (12) tt This curious relation between w and the spore-circles follows from that between w and d as determined by o and 6. Thus w= 8” + m and S” = S” —d m = S’” sinS =(S” — d) x — = AY ic” d a d2 Therefore w = (S” — d) + ————— Mt (Sys dts’ di Oe (8”.4.¢d) 6" —d) S 4” S 4 AY ” AS S74 = from (3) Si ce ih eek eS Ss) Pn | ea (13) d (1 — sin 8) d (1 — sin2 §) because S”’ (1 + sin §) = ———————- x (1 + sin §) = ———————_ sin 6 sin § d cos2 § = = d cots§ coss sin 6 The longitudinal relations may now be considered. Let the sterigmatic patch circle m be projected on to the basidium-apex, as in the lower part of Figure 3. It occupies arc Hw and subtends 2 HAw or « Bisect « and the bisecting radius AF will meet the arc Hw where it is intersected by the projection of the sterigmatic circle M. That is, if the centre of the longitudinal sterigmatic arc on the basidium is projected parallel with the long axis of the basidium up to the level at which the spores are developed, it becomes the sterigmatic tip, whence the spores grow abaxially with the spacing initiated on the basidium-apex. This is one of the most remarkable agreements between the geometry of the basidium and the observation of the spore-circles, the sterig- matic circle and the spore-angle. At first there seems to be no reason for the position of the sterigmatic tips, other than structural convenience; now it is seen to be an integral part of the basidium-unit. The tips are eccentric in the sterigmatic patch circle. It would be interesting to compare this construction with the basidium of Gasteromycetes with sessile or subsessile spores. By means of the angle « , other relationships in the basidium can be found which have value in checking reconstruction from specific data. 172 Gardens’ Bulletin, Singapore — XXVI (1972) ford MEW COG vga ksinasnoesvancce'esy ous Geallatineaasmasencie ceuee ene atabadenninte caees yee (14) 1 — sin § OS oa ssn nnasiee an nen un ote hina mmc oaebanesns ale ae eee (15) 1 + sing Am 2Am 2Am because cos“« = ——— = = —_______- from (12) AH w S’”’ (1 + sin 8) A ig m Ss” S”’sng% Soer— ea.) Am = — —- — = — — — = from (11) z 2 2 2 Z Therefore S’”” (1 — sin 6) 2 1 — sin § COS 0 = —AA > X« = 2 S’” (1+ iné) 1+ sin6g SY a m Alternatively, Os OC Se mn isis ige sn cucigeapas canes seine s sean alvs eile cana (16) Ss” +m 1 — cos «© sin § = Fronts (1S) oa ccsscsnexcesenucanshumesenadeemes ss hen tes aaenpege eg aden (17) 1+ cos« S” —m ST a riiiccdvuscevcns usanachen eon 0 sy aoanne aapheduusan sw ickiens sco entinngean sued: emus aeaaennnnnnnnn (18) d MC S”—M 2 S’—m because sin 9 = —— = —————- x —- = Bi, 2 d d 5S” —M d = ——____- froma (16) auncbincsviincc ee tRD Mewes coneweue cee (19) sin 6 1 + cos« M = w / ———-——_, from (14) erie ALAR An oso ecoe nce (20) 2 1 =w /————— from, (15) 1 + sin$ d (1 — sin 6) mn macnn ee eR ee (21) sin § do d (i — sin 8) because S’”’ = S” — d = —— — d = —___—_—_- sin 6 sin 6 y SU M =d cot§ cos§_ ,/————— Sram: 420) ated, 4) 3)..2.. fer (raar | (lazy) (62)... (62) 7" GE | G1) B. nanus 5.38 10.96 | 5.58 | 8.35 | 10.98 | 6.81 | 8.32 | for d’ 4.03 10.96 | 6.93 8.10 | 9.48 | ford ratio to d’ 1 0.441 1.04 | 1.55 | 2.04] 1.26 | 1.54 ratio to d ne ne ae 0.924 72 | BOT 142.73 (number) on .. | (250) | (180) | (46) | (46) | (42) | (92) B. phaeocephalus .. pout Mes 11.65 | 7.10 | 8.68 | 12.20 | 8.37 | 9.87 ratio tod “4 ay 1 0.810 Poo | 1.98 | 20 | 184) 217 (number) + .. | (102) | (160) | (S50) | (50) | (82) | (66) B. umbilicatus et , at) ad 13.06 | 7.69 | 8.86 | 12.33 | 9.12 | 10.85 ratio to d F out ay 0.720 1.43, | 1,65. |..2:30:1.. 1,70.) 2.02 (number) +e wo |? (oo) |. GO) (9) (9) (6) | (38) 180 Gardens’ Bulletin, Singapore — X XVI (1972) Table 4 — continued. Spore-measurement in Boletus. Observed values Theoretical values* fx! ei? (py) d o Sie M w Ww B. ravenelli - to Ae Gaze 10.22 | 6.01 | 6.81 | 9.96 8.49 ratio to d df OPK, & 0.720 1.43 | 3.62 | . 2.37 2.02 (number) + .. | (132) | (92) | (40) | (40) | (22) | (69) B. aureomycelinus all ae 10.89 | 6.00 | 7.90 | 10.47 8.69 ratio to d cf. ae all 0.575 1.24 | 1.62 | 2.14 1.78 (number) se .. | (126) | (56) | (28) | (28) | (36) | (64) P. cingulatus he sw 1 Sz 11.98 | 6.80 | 8.96 | 11.63 9.95 ratio to d a AS ae 0.615 L3i:t i e8 Zoe 1.91 (number) - .. | (188) | (80) | (40) | (40) | (7)! (73) Ss’ AN < * calculated from M=S’” — and w= Sed iS? d’ outer spore, d inner spore. (number) refers to the number of observations. Various errors in observation and construction can be imagined to explain the differences in positioning of the sterigmatic patch circles, but without under- standing of the physical forces involved I do not think that they help the solution. I mention one possibility because it is a step in the adaxial displacement of the sterigmata. In Figure 5 I have drawn w’ w’ as the false base of the basidium-apex; it is the base that makes the basidium-apex hemispheric instead of ellipsoid and, as will be seen, it has the effect of shifting the sterigmata upwards and adaxially to compress the spore-circles. The sterigmatic patch, nevertheless, seems to fit not the arc of / HA’ w’ but the upper half of this arc. It appears that the compres- sion of the cone-effect of the basidium-unit in the concave hymenium acts on the sterigmatic patches before emergence of the sterigmata. This is a part of the more general problem of the hymenium-level where some force causes basidia to stop elongating and to mature; thus cystidia are basidia that have exceeded this level, often before it has become established in the developing hymenium, and remain sterile. The hymenium-level is another apparently electrostatic effect of the fruit- body. B. phaeocephalus (Figure 5). At first sight this appears to be a normal recon- struction but the S” circle lies within the w circle, a spacing circle on S”’ with radius S”’ w greatly exceeds the sterigmatic patch, and the observed spacing, whereas a spacing circle on M with radius Mw almost fits the tangent to the spore; and the sterigmatic arc is much shorter than Hw. I detected no exospore but comparison with Figure 6 shows that the spore of B. phaeocephalus is an endospore developed from an adaxially displaced sterigmatic patch. This construc- tion probably represents the normal state for the smooth boletoid spore. 18] Boletus phaeocephalus, with w’w’ as the false base of the basidium-apex; x 5000. Studies in the basidium Figure 5. 182 Gardens’ Bulletin, Singapore — X XVI (1972) B. albo-ater (Figure 6). This reconstruction shows the excessive o’ spacing which leads to the disappearance of the exospore and its substitution by the endospore with (m) spacing. The centre of the (m) circle is not on S’”” but midway between S’”’ and M (on the inner limit of the endospore). Construction of « “ on the false base gives the sterigmatic arc with the radius of the half angle contacting the M ordinate. B. nanus (Figure 7). In this reconstruction I have drawn a normal spore on the right with its evanescent exospore and durable endospore. The centre of the (m) circle almost coincides with M; that of the m circle is slightly abaxial from S$”’, but for both the radius is now to S” which coincides with w; a spacing circle with centre on S”’ and radius S”’ w is clearly of no consequence. On the left I have drawn an imaginary ellipsoid spore (with endospore and _ exospore) and an imaginary globose spore; the volume of the latter (6.8 x 7.ly) and that of the ellipsoid endospore (8.0 x 4.6u) are equal to the volume of the normal endospore. They are positioned according to the spacing factor o’ for the normal exospore and its theoretical value of w (as from the table in Appendix 2). These spores are eccentric because S” is 14.47» for o’ and 12.59 for o. The recon- struction shows how a globose spore of normal spacing relations with the basidium, such as might have occurred in the ancestor of Boletus, may have been transformed into the smooth boietoid spore by compression of the spore-circles commensurate wiih the concave hymenium. B. retisporus (Figure 8). In this reconstruction (S”’), for the inner spore, almost coincides with M; they are 0.3 apart. It appears that sterigmatic spacing is more compressed than spore-spacing; thus M is displaced adaxially towards S”’ and the wider gap which results between exospore and endospore may allow the development of the reticulate ornamentation all over the spore without a smooth adaxial patch. In this case the (m) circle contacts the w’ ordinate of the false base. It is likely that B. longicollis agrees with B. retisporus though the measures of S’” and M (in Table 4) do not show it. The basidia of B. longicollis collapse very easily in aqueous mounts and my measures of the spore-circles are probably too small. (B. longicollis has also been described as B. altissimus Mass. and B. singa- porensis Pat. et Baker). Strobilomyces velutipes (Figure 9). The reconstruction is complicated by the necessity to distinguish the spore-circles S$’, S” and S”’ for the exospore from (S’), (S”) and (S”’) for the endospore. Two reconstructions are shown. That on the right fits the smooth boletoid spore and that on the left is the true spore. The spore on the right has exospore and endospore concentric, but this would allow ornamentation to be developed all over the spore, as with B. retisporus. The normal spore has a smooth adaxial patch, where the membrane of the exospore can be seen to touch the endospore, and the warts are graded in height from the shortest on the adaxial face to the tallest on the abaxial. The endospore has, therefore, to be constructed eccentrically to the exospore. The m circle for the normal spore gives too large a spacing but the (m) circle with centre on S”’ and radius S”’ w gives the correct spacing for the endospore. This arrangement means compression of S’ to (S’) without compression of M, and S”’ remains unaffected. The spore is produced on a wide basidium with much less compression than with the smooth boletoid spore and is, accordingly, broadly ellipsoid and even subglobose. The whole basidium-unit is the least modified and, presumably, the most primitive among boleti. 183 mS" MwS (S)S' Figure 6. Boletus albo-ater, with exospore and endospore; x 5000. A Studies in the basidium Gardens’ Bulletin, Singapore — X XVI (1972) 184 ns Ce — d in on the left, imaginary ellipsoid he normal endospore but altere 38 Cc fo) fae} LY a) “hole a oe ~ » 5) 3) fe —_— OY om) Se 8 — Y on om -m- ore a Sta F- 2 gos eos 282 a nn oO 22O £838 Sos bee SU 3 weg Qo So 4 Figure 7. 185 Studies in the basidium (5) S' mim SSM w S" A Figure 8. Boletus (Heimiella) retisporus; x 4500. Gardens’ Bulletin, Singapore -— X XVI (1972) 186 ----------------- = ee ee ee core Figure 9. Strobilomyces velutipes with a normal spore on the left and an abnormal spore, constructed as a smooth spore, on the right; x 4800. ™ oo Y —_ — Ww a C S io S Y) t+ ad » 3 sz wn = 1) Vv — at oe s o = ~ = = a o = ao) = = 3 — wn ~ { 8 S nn S ~~ 3 £ 4 EF 8 — = Y — wi, aS nbs a - a S a 3 Oo — ay v Tp) 5 = = a) lp) iL — no Pe UE eto ole lee a sa Oe a *. a oo = = == ww ew ww se Ree ese Pe ee ww we ee Pe ere ee ee ee elU S = aS S v = ™ = ie) aS aS) = ~ A Gardens’ Bulletin, Singapore — X XVI (1972) 188 Boletus emodensis; x 5000. Figure 11. Studies in the basidium 189 Having no detailed analysis of the curvature of the basidium-apex in this species, the reconstruction has been made on a hemispheric apex. It is probably a reconstruction on the false base with the effect that the radius which bisects « meets the sterigmatic arc where intersected by the M ordinate. Porphyrellus sect. Graciles (Figure 10). Having no fresh material to study in this interesting case allied with Strobilomyces, I used alcohol-formalin material of an undescribed Malayan species. This did not give reliable measures for the spore-circles. I reconstructed the basidium-unit from measures of d and w and the ratio w = 1.655d for d’ tan, which gives o’ = 0.5. The measures were as follows: — exospore 17.36 X 8.41,» (radial), X 9.16 (tangential); endospore 17.36 X 7.26, with truncate conical warts 0.7— 1.24 high (mean 0.954) on the sides and abaxial face, 0.1-0.3 on the adaxial side; (average of 30 spores); basidium 32 X 15.16u, with the sterigmatic circle 11.84 wide (average of 50 basidia and 40 sterigmatic circles). The reconstruction on a hemispheric apex for the basidium is similar to that of Strobilomyces but the spore is compressed and elongate through the adaxial shift of M towards S’”. The point is shown by the value of S’ which is 27» in the reconstruction though, theoretically, it should be 28.6u. The adaxial shift of M is the principal change leading to the smooth boletoid spore. Thus, this section of Porphyrellus appears to be half-way between the state of the basidium-unit in Strobilomyces and that in Boletus with smooth spore. B. emodensis (Figure 11). The high compression of these spores is shown by the upward and adaxial displacement of the sterigmata on the wide basidia. Though I have constructed the basidium-apex as a hemisphere, the false base would need to be lifted even higher, as shown by the dotted line, to fit the sterigmatic patches; the dotted line gives w= 10.lu, which is the usual width of basidia in smooth spored boleti. Then, as further evidence of compression in this figure, the real value of w slightly exceeds that of S”; the m circle has centre M and radius Mw; the (m) circle has centre S”’ and radius S’” M. B. longicollis belongs in this same alliance of Boletellus with striate-costate spores. So, apparently, does B. obscurecoccineus Hoehn. with faintly striate spores equivalent to the endospore of B. emodensis. It would seem that in this alliance the series can be traced from the broadly ellipsoid spore to the smooth boletoid. Unfortunately other species in Malaya have n = 2 or 3, and comparable results could not be obtained. Phylloporus This genus is close to Boletus in its wide sense and differs in the lamellate hymenium. Among fifteen Malayan species for which I have accurate measures (Corner 1970), thirteen have d = 0.42w to 0.56w, but in P. hborneensis d = 0.58w and in P. coccineus d = 0.71w. Evidently there is the tendency to the compression of the spore-circles in this genus without the development of the fully tubular hymenium. I worked out the details for P. cingulatus (Table 4); they showed the boletoid features of close agreement between the values of S” and w and of the location of the M ordinate through the m circle on the false base. Gardens’ Bulletin, Singapore — X XVI (1972) 190 — ee Se Se Oe Ee = Te Figure 12. Marasmius nigripes; x 5000- Studies in the basidium 191 If, as I consider, the poroid Boletus has been derived from a lamellate agaric of Paxillus-Phylloporus affinity, then it seems that the characteristic of the Boletus-spore was developed in this ancestor, but both Paxillus and Phylloporus have smooth spores. This conflicts with the idea, put forward in previous pages, that the primitive Boletus had an ornamented endospore. Therefore the monotypic South American Phylloboletellus Singer stands out because it has the phylloporoid hymenium and the striate-costate spore of Boletellus (Singer 1964). According to Singer’s measurements, in Phylloboletellus chloephorus d = 0.78w and this implies both o’ and o spacing. Certainly it suggests a primitive boletoid fungus, though apparently without adaxial patch. This is supplied by Gloeocantharellus with ornamented spores and apparently no exospore. For three of its five species d= 0.59 to 0.65w (Corner 1969). There appears to be a single spacing factor with external ornamentation, as in Russula and without any of the elaboration of the boletoid spore. Marasmius nigripes Schw. The remarkable spores of this fungus illustrate the use of geometrical analysis (Figure 12). In adaxial and abaxial view the spores are cruciform; in side-view they are trigonous and in end-view crescentic. They develop from the spore- rudiment into a shortly clavate form (6 X 3), typical of many species of Marasmius; this form is shown in dotted outline in the figure. Then shoulders develop to extend laterally and abaxially while the apex lengthens into the third arm and the spore-body swells to full size. There are no radial arms against the boundaries of S’ and S”’, Figure 12 is based on averages from ten basidia in side-view and ten in end-view; the basidium apex is drawn as semicircular. On this reconstruction I then drew (i) the radial tangents to the spore-body and spore-arm; (ii) the sterigmatic patch circle (centre S”’, radius S’” w); (iii) a spore-body circle b inscribed on S” between S’”” and S’, and found to my surprise that it fitted the tangent to the spore-arm; (iv) a spore-arm circle v (centre S”, radius S” to the apex of the arm), and found to my surprise that it contacted adjacent spore-bodies and almost reached the long axis of the basidium; (v) an outer spore-circle a (centre A, circumscribing the spore-arms); (vi) an outermost circle u circumscribing the v circles, as the outermost limit of the basidium-unit. This figure leads to the following conclusions: — (i) the m circle may be the prime spacing circle for the spores; that it does not contact the tangent in the drawing may be due to the difficulty in locating the exact tangent in this region where the curvature of the spore is changing. (ii) the m circle gives © = 0.42 or w = 1.5ld, which corresponds with w = 1.52d for average measures of spores (9-12 x 8-llu) and basidia (25-32 xX 8-10,). (iii) the 5 circle, contacting the spore-arm tangent, gives « = 0.44 or w == 1.55d. (iv) the actual spore-body (with radius measured along the dotted line joining adjacent spore-centres in Figure 12) gives c = 0.61 or w = 1.84d. (v) the spore starts with theoretical spacing of o for the mature basidium; it then reduces this to o’, grows radially to the allowance of oc’, but this extension occurs only about the mid-length of the spore; the spore-body is itself compressed tangentially to a value between o’ and co. 192 Gardens’ Bulletin, Singapore — X XVI (1972) (vi) in final form, as shown by the v circles, the spore comes to fulfil by some physical means the whole spore-sector. The compression of the spore-body, mentioned in (v), may bear on the nature of the long, narrowly clavate spores of many species of Marasmius, as another instance of spore-compression, similar to that of Boletus. I have seen a fine mucilage sheath round many Marasniius spores. On germination of the spore of M. nigripes, it is only the spore-body that swells. References Buller, A.H.R. (1922) Researches on fungi. Vol. Il. Longmans, Green and Co.; London. Buller, A.H.R. (1924) Researches on fungi. Vol. II. Buller, A.H.R. (1931) Researches on fungi. Vol. IV. Corner, E.J.H. (1936) Hygrophorus with dimorphous spores. Trans. Brit. mycol. Soc. 20, 157-184. Corner, E.J.H. (1947) Variation in the size and shape of spores, basidia and cystidia in Basidiomycetes, New. Phytol. 46, 195-228. Corner, E.J.H. (1948) Studies in the basidium I. The ampoule effect. New Phytol. 47, 22-51. Corner, E.J.H. (1950) Monograph of Clavaria and allied genera. Ann. Bot. Mem. 1. Corner, E.J.H. (1954) Further descriptions of luminous agarics. Trans. Brit. mycol. Soc. 37, 256-271. Corner, E.J.H. (1964) The life of plants. Weidenfeld and Nicolson; London. Corner, E.J.H. (1966) A monograph of cantharelloid fungi. Ann. Bot. Mem. 2. Corner, E.J.H. (1968) A monograph of Thelephora (Basidiomycetes). Nova Hedwigia Beih. 27, 1-110. Corner, E.J.H. (1969) Gloeocantharellus. Nova Hedwigia 18, 793-805. Corner, E.J.H. (1970) Phylloporus and Paxillus in Malaya and Borneo. Nova Hedwigia (in press). Corner, E.J.H. (1972) Boletus in Malaysia. Government Printer. Singapore. Malencon, G. (1958) Le dévelopment des spores chez les Phylactéries. Bull. Soc. mycol, Fr. 74, 423-435. Perreau-Bertrand, J (1964) Recherches sur la différentiation et la structure de la paroi sporale chez les Homobasidiomycétes a spores ornées, Annls Sci. nat. Bot. ser. 12, 8, 639-745. Singer, R. (1964) Phylloboletellus. Nova Hedwigia 7, 129. Studies in the basidium 193 Appendix 1 Summary of the relations between the spores and the basidium (n = 4) d = spore-width; m = diameter of the sterigmatic patch circle; w = basidium-width; M = diameter of the sterigmatic circle; S’, S”, S”’” = diameters of the outer, middle and inner spore-circles respectively. eo = the sterigmatic patch angle; § = half the spore-subtending angle; @ = the spore angle; @ =the spore-spacing factor. Text reference a 0 Sr a a a ae (1) fain’ d S*=d/2(1 Se ame aN Ra ss fw badip dnd w ves db uvnsde sus auc huetbeacs¥amder GnSevkde (2), (6) sin § d (1+sin 6) f= 5" 4+d=d f2ai +o)t+d= ae rise =a tiv + GOSEO) = 8 SCORE hoc 5 ccc escsiewcne (7) sin d(1 — sin 6) 8” =S" —~d=d /2(1+ 0) —d = ————-= d (cosec § — 1) er eee Ce. Sore eee (8) sin § Ss +e 2) and J —S”= 2d a Bed Lhd SV Ae —=1+sin§ = 1 — sing a eae —- =w Ss” Fe So wcos « M= = =e | Se a ART eee anrrren. Aa a remnren (14) 2 1 + oor 7 M = dcotscosd eS acs sea Mt AOR aBES cn alth veecapacertncadons ee. ie (22) 1 + sin § Me OF ot (oes § 07) Sf J 21 SY OOTY oooh eens seceshsnagah en denecens (9) A i cosec 6 =—— -l= aE en Ane 5 eee BOR ii hioe ek Aton sinc dratindawahedaths cxwsisesa (10) df 2 2. 1 + cos« I RE IE) AEE RE RO. ARE. A ORE See eee, PE (15) ) Fae oe — cos « ; ee eI ON Gal Savin pais as eied bah vi ekabeavadehccvcsedet ninteVeeseadicceungescoemerencscses (11) Nie he ee i eae TT ee ee, eee ee eee (12) (13) Ss d 1 S’—M 1 — cose SAT cecceeccc ese ec ete netec ene ea ene nateneneneeeens (17) 5S” (2( +0) S” sn@ 1+cos« i, ag 1a m 1 — sin 5 A fa a ia cdl sts stdensstcwsssstescnrascacececcanecsteccsseccesescaceante (15) , tm. J2+3n8 5 ad sin @ = se = cosec § — cot §cos § hcp ood cE ek feuedi onpine (18), (23) d S” sin § 1 + sin§ 5S’ —M Gardens’ Bulletin, Singapore — X XVI (1972) 194 IE SP VT o8b BP ofS I 09S 67 o19 OV C9 OT v9 6E $9 VI oL9 LS 089 8V OL OS oCL iL oGL LC oLL L 08 20 bl oLE 61 o8€ EV 6€ CE olP Of ovv LT SP 18 9D WET OLD ST o8P i£E o6P € ofS CS oS 8 oS i€E oLS LV 09 6 “CT OC 8T £7 IT LC CS oCE VC olD CV oP IT OV OT .6¥ 87 oS VI 9S LP 09 IS G9 /OT cL /£ 008 006 SZ PT6LV'S PO8PL'V P£900'V PICSC'E Prrlr'c P6PIE'C PICST'C PO886'T P6618 T P9619'T P6rLr' I P6£67C'T P6LOT'T P6TT6'0 P690L'0 PTETB'0 PO86L'0 PEV9L'O PILIL‘O Pp9P9'0 POL79'O PTL09°0 POrss 0 P6LSS'0 P987S'0 P6r6r'0 P6SSv'0 P60IV'0 P69SE'0 P6767'0 PISO'S P9S9'P P9S9°S P9s9°9 POEE'V POS6'E POS6'P POS6'S PEO9'e PCV E PCVT'V PCVE'S PTL8°7C PSes’c PSes'e PSes'y P9CI'C P87é8'T P878°C P878'E P9L6'T PL89'T PL89°C PL89°€ Pres T Psrs'T PSPS" PSPS'€ PILOT Pror' I Pror'c Pore POTS T PC9T'T PC9C'C PC9T'E pogve I PICT'T PICT? PICT’ Peoc'T P0860 PO86 T P086°C PCHO'T P8E38'0 P8E8'T P8E8°C P8L3°0 PL69°0 PL69'T PL69°C P9OTL'O | PSsss'd Poss PSSS’c Pelps'oO | Privo Priv Priv'c W NY NY S O Ul SUONeLIeA YIM pUN-UINIpIseg oy} JO syred oY} Ul sUOTeLIeA Z XIGNdddV oy ae OS Oem PUBLICATIONS OF THE BOTANIC GARDENS SINGAPORE 1. The Agricultural Bulletin of the Malay Peninsula (Series I). ‘Only Nos. 3, 5, 7, 8 and 9 available, at 20 cents each. 2. The Agricultural Bulletin of the Straits and F.M.S. (Series I). Vols. 1-10, 1901-1911, monthly issues. Many parts available. Price: $5 per -volume, 50 cents per part. 3. The Gardens’ Bulletin, Straits Settlements (Series III). Vols. 1-11, 1912-1947. on Vol. 1 (1-5) January-May 1912 issued under title of Agriculbaeatil Bull oe Ain: of the Straits & F.MLS. Prices on application. 4. The Gardens’ Bulletin, Singapore (Series IV). Vols. 12-25, 1949-1971. Vol. 26 Pt. II in print. Price: Vol. 13 Pt. 1 (New impression) $12 gh copy, $20 per “eS Vols. 12 & 14 $13 per vol. Vol. 15 $20 per vol. Vols. 16-25 $25 per vol. Individual parts vary in price. Frequency: Parts are published as materials become available. Bee 5. Materials for a Flora of the Malay Peninsula, Monocotyledons. Parts 1, 2 and 3 remain available. Price: $10.per set, $5 per part. 6. Annual Reports. Many from 1909 onward remain available: Prices vary. 7. (a) Malayan Orchid Hybrids by M. R. Henderson and G. H. Addison. $15. z (b) Malayan Orchid Hybrids, Supplement I by G. i. Addison. $21. 8. A Revised Flora of Malaya. *“ (a) Vol. 1, Orchids, by R. E. Holttum. $30 (3rd ed. 1972 ‘ileus (b) Vol. 2, Ferns, by R. E. Holttum. $20 (2nd ed., 1968). (c) Vol. 3, Grasses, by {H. B. Gilliland. $30 (1971). 9. Boletus in Malaysia by E. J. H. Corner. $50 (1972). Items 1-6 obtainable from the Director, Botanic Gardens, Cluny Road, § 90. : Items 7-9 obtainable from the Government Printer, Government Printing C O Upper Serangoon Road, acai chs a: Prices quoted are in Singapore Dollars eee Overseas postage is extra — gS oa ; “ae ~ EOE GEOL YEE ERIE EE” iE ASP a is NO REF Rr RECEIVED i ins Fy, 3 THE BEC 9.9 1975 a ; ; o “LIBRARY a GARDENS’ BULLETI SINGAPORE . Vol. XXVI, Part I 15th September, 1973 CONTENTS ; PAGE 4 Dctrmons Ee: EEeuCRCY and Habitat of Tree Species in the Rain Forest a Be of Ulu Kelantan - - - . - 195-210 _ Horrrum R. E.: A New Bamboo from Mt. Kinabalu - - - - 211 Marie A. Martin: Notes on the Vegetation of _the Cardamon Mountains, a _ Cambodia - : : - - 5 - 213-222 _ Hsvan Ken & Heastert E. AS The x ylopia malayana fruit: significance of its dehiscence - 223-225 ¢ _Sitior0 Kurata: Nepenthes from Borneo, Singapore and Sumatra - . 227-232 a HsuAN KENG: Annotated list of seed plants of Singapore (I) - - - 233-237 * ‘Draxsriet De? Korthalsia hispida Becc. in Malaya - - = - 239-244 ay R so A. N. and Lee War CuIN: Pollen Vogue and Gomalnation in Some ee Orchid Hybrids - - - - : 245-257 “Nayar M. P.: A Review of the Genus Creaghiella Stapf. (Melastomataceae) - 259-261 ) CK H TIONG-KHENG: New Plant Disease Records for Sarawak for 1970 and 1971 263-268 BA KOCHUMMEN K, M. & Wuitmore T. C.: Notes on the Systematy of Malayan oP . be Phanerogams XVIII-XXxII - - 269-287 ANNE : JoHNSON: A survey of the occurence of epiterranean soil algae in Singa- f E: pore Island - _- - - - - - - . 289-302 LiL J.: Pollination in Dioecious Figs: Pollination of Ficus fitulosa by i i \y Ceratosolon: hewitti - - - - - - - : 303-311 To be purchased at the Botanic Gardens, Singapore Price: S $18 Published by Authority é ‘ : : Printed by the Singapore National Printers (Pte) Ltd Sr 1973 ae een Frequency and Habitat of Tree Species in the Rain Forest of Ulu Kelantan by T. C. WHITMORE Forest Research Institute, Kepong* Abstract In 1967 a survey was made as part of the Forest Resource Reconnaissance Survey of Ulu Kelantan district. 26, 628 trees were enumerated > 4ft girth on sample strips 1 chain wide, in primary rain forest mostly below 2800ft elevation. For this analysis the survey area of 1672 acres has been divided into 4 classes on geology and altitude, comprising sedimentary and granitic rocks and below or above 750ft. The occurrence of different tree species, genera and families in these 4 classes is listed in a series of Tables. No such extensive analysis has previously been made. There are many groups which are markedly more common in one or other of the 4 habitat classes and several groups have a distri- bution in Ulu Kelantan which counters subjective preconceptions. It is suggested that at any altitude species’ distribution is likely to be more closely correlated with relief and soil than with geology. INTRODUCTION Our knowledge of the habitat preferences of Rain forest trees in southeast Asia is very largely based on the subjective impressions of forest botanists not backed up by measurements of tree frequency in particular habitats or precise records of the nature of the habitats themselves. An enumeration of all trees over and including 4 ft girth on a small sample of the Lowland Rain forest of Ulu Kelantan district, northeast Malaya has been the stimulus to evaluate the relative frequency of the various tree families, genera and species occuring there, and to analyse their distribution between the two major kinds of rock in the district, and at low or higher elevations. This is of course, background information of fundamental importance when attempting to summarise our knowledge of particular species for such publications as the new Tree Flora of Malaya (Malayan Forest Records 26, Whitmore 1972, 1973 a). Species with horticultural and some with known pharmacological potential enumerated in this survey have already been discussed in Whitmore (1971). It is well known in Malaya, as a matter of observation, that different species have different altitude preferences, and there is a growing awareness that the rock type on which a forest occurs, to some extent at least, appears to determine its species composition, in a manner tentatively analysed below where we attempt to investigate a little further these suspected discriminants to species distribution. THE FOREST SURVEY The forest survey was conducted between June and November 1967 by Forest Resource Reconnaissance Survey teams under the able direction of Mr. N. Cerra, working from Kepong as an American Peace Corps volunteer, as part of an operation which has now covered the lowland forest of the whole country. * Present address: Mead Hall, Saffron Walden. Essex CB10 IUX, England. 195 196 Gardens’ Bulletin — XXVI (1973) The forest was divided into a series of ‘phototypes’ from the study of aerial photographs. Teams then went into Ulu Kelantan and systematically sampled these types throughout the district, at a low intensity. The sample lines were 1 chain wide and up to several miles long, but divided into 10 chain lengths, so individual plots of 10 sq. ch. (1 acre) are the basic unit. The lines were cut across the ‘grain’ of the country so as to sample ridge tops, hillsides and valleys; altitude was recorded; the precise position of each line was recorded on a set of 1 inch maps. Access was mainly by river, and from any one camp a series of lines was surveyed; the inset of Fig 1 shows the location of each cluster of survey lines. The total sample area was 1,672 acres; 26,628 trees were enumerated and named according to the preferred vernacular list in Kochummen & Wyatt-Smith (1964). The upper limit of most lines was below 2,800 ft, a few went up as high as 3,800 ft. Thus the whole survey is restricted to the Lowland Tropical Rain forest formation (Symington 1943, Burnham & Whitmore 1969). Because this was a reconnaissance survey no attempt was made to take a rigorously, objectively selected sample. It was adequate to cover different photo- types throughout the whole district. It follows that it is unwise to multiply up from the results of this sample to find tree numbers over the whole of Ulu Kelantan. The results are, however, likely to be indicative for this part of Malaya. No pretence is made that the present analysis indicates frequency and habitat everywhere in Malaya; indeed, as pointed out below, there are, in Ulu Kelantan, several marked departures from what are suspected to be the preferences of species and groups elsewhere in Malaya. The forest data were divided into stands for each of which a table was prepared stating, inter alia, the number of stems of each vernacular-named group occurring and the total basal area of all the trees enumerated. The stand tables have been the basis for the present analysis. Each stand is either one whole survey line or part of one. Each one contains homogeneous forest of one of the forest types recognised by the Forest Resource Reconnaissance Survey. For the present analysis the stands were divided into two groups, those below 750 ft elevation and those above, to separate lowland dipterocarp forest from hill and upper dipterocarp forest in the sense of Symington (1943).* Regrettably, nearly all the stands transgress the 750 ft contour, and have had to be included on the side where they mostly lie. This can only blur the altitudinal preferences of species and groups and weaken the clarity of the analyses presented below. ULu KELANTAN Ulu Kelantan district was in 1967 mostly still under primary rain forest. The major areas of settled agriculture are shown in the inset to Fig 1. Logging had then not long begun, though by early 1972 much of the district had been let out on logging concessions. The total area of this administrative district is 4,817 sq. mi. and about 80 per cent was then still forested. Within the forested part of Ulu Kelantan there is little flat land except in the Lebir/Aring basin in the east; mostly the rivers run through narrow, steep-sided valleys. The main mountains lie to the west and east (Fig 1) and this is also, roughly, the distribu- tion of granite, of Triassic age, which forms the Main and east coast ranges * Symington drew the boundary at 1,000 ft. I have divided at 750 ft on the advice of Mr. P. F. Burgess who considers the latter a more realistic elevation especially, as here, in hilly country. Tree Species in the Rain Forest of Ulu Kelantan 197 5000 ft 1200 ft 600 ft Fig. 1. Ulu Kelantan District, location and relief. Taman Negara also shown, S. and E. of Ulu Kelantan and partly over-lapping. Inset, location of sample sites, rock types (granite dark, sedimentary rocks pale) and approximate main area of cultivation (hatched). Geology from Alexander 1963. Cultivation limits from Wyatt-Smith (1964). 198 Gardens’ Bulletin — XXVI (1973) of the Peninsula respectively. The centre of the district is mainly of Permian and Triassic sedimentary rocks, which are described further below. The geology is not known in detail, but the granite/sedimentary rock boundary is fairly accurately known; only eight of the 222 survey stands lay so close to the boundary that their geology is doubtful. The stands were classified as lying on either granite or sedimentary rock and the analysis of species frequency was made for four groups, low granite (below 750 ft), high granite (over 750 ft), low sedimentary and high sedimentary; in these four classes 113.8, 448.8, 716.4, 393.0 acres had been sampled respectively; it can be seen that low granite was distinctly under- sampled. SPECIES AND FAMILY FREQUENCY So few have been the extensive, quantitative analyses of composition and habitat in Malayan Lowland Rain forest that, when quoting the relative frequency of species, one is thrown back on observations scattered through the pioneer work of Foxworthy (1927). Table 1 shows the 21 species in order of decreasing frequency which have 10 or more stems per 100 acres in this survey. The first column gives the average density, the next four columns give, for the ten commonest species, the density in the four habitats. A major weakness in this table is that many species were not individually recognised in the forest. Dipterocarpaceae dominates the table, partly because it is a very common family but partly also because every species has a vernacular name. Burseraceae and Sapotaceae are both very common in Malayan Lowland Rain forest but their species were not distinguished from each other. Nevertheless, the top third of the table is probably roughly correct, only lower down would one expect other species, especially of Burseraceae and Sapotaceae. Note that the order of the first six species does not vary between the habitats, though the order does vary lower down. The important exception to this generalisation is Shorea curtisii* (seraya) which is almost confined to hill ridges where it sometimes forms almost pure stands; this species is so common in hill forest that it still easily comes top of the list, despite its low frequency in lowland forest. The next two species (S. parvifolia, S. leprosula) are both light red merantis which are fast growers and light demanders and come up gregariously in clearings. They are amongst the commonest trees everywhere in Malaya. In Table 2 the 42 families encountered in the survey are arranged in order of decreasing frequency. Again, the first column gives the average density (in stems per hundred acres rounded off) and the next four columns relate to the four separate habitats. One sees at once why these forests are known as dipterocarp rain forests, the family Dipterocarpaceae is four times as abundant, amongst the trees > 4 ft girth, than Leguminosae, which comes second, and this is true of all four habitats. As in the species table the order of frequency of the families differs between habitats low down the table, though the first four families are in the same order in every one. Of the two biggest tree families in Malaya Euphorbiaceae comes surprisingly high in the list compared to other surveys which have been made (Whitmore, 1973 b); Rubiaceae, which is actually slightly larger than Euphorbiaceae in Malaya in number of species, comes much lower, this is because very few of them reach 4 ft girth. Leguminosae always come high in a survey of this nature, but * Species names throughout follows the new Tree Flora of Malaya (Whitmore 1972 and 1973a) or, for families not yet revised, Kochummen & Wyatt-Smith (1964). Tree Species in the Rain Forest of Ulu Kelantan 199 Table 1. The commonest of the species identified in the forest, those with on average more than ten stems per hundred acres granite sedimentary ee” Now" high” low high 1. Shorea curtisii(Dipterocarp.) .. xr 125 21 126 20 345 2. Shorea parviflora (Dipterocarp.) . . m 108 91 93 127 95 3. Shorea leprosula (Dipterocarp.) . . a} 80 a7 88 i 82 4. Koompassia malaccensis (Legum.) he 51 36 39 66 45 5. Shorea dvatis (Dipterocarp.) * e 40 26 33 51 34 6. Dipterocarpus cornutus (Dipterocarp.) 1 35 40 13 56 24 7. Elateriospermum tapos (Euphorbi.) =e 33 16 45 20 49 8. Dryobalanops oblongifolia (Dipterocarp.) .-. 31 17 3 66 6 (Shorea multiflora (Dipterocarp.) Ae 23 16 11 30 25 a ie palembanica (Legum.) oN ye 23 18 26 23 22 11. Shorea platyclados (Dipterocarp.) A 21 — — — — ( Koompassia excelsa (Legum.) .. yg 18 —_ _ — _- , | Cynometra malaccensis (Legum.) a 18 — — — — 14. Dipterocarpus crinitus (Dipterocarp.) $i i5 — — = — 15. Shorea bracteolata (Dipterocarp.) o 14 — — — — 16. Shorea guiso (Dipterocarp.) at iu 13 — — — — 17. Parashorea lucida (Dipterocarp.) cs 12 — — — —- 18. Pometia pinnata (Sapind.) . on 11 — -— “= os Mesua ferrea (Guttif.) ter ~¢: 10 “— — — — 19.< Endospermum malaccense (Euphorbi.) ne 10 — = — — | Ochanostachys amentacea (Olac.) dé 10 a a a5 = Family name abbreviated in brackets; stems per hundred acres rounded off to nearest whole number. Myrtaceae and Lauraceae are usually lower and Burseraceae higher. The complete absence of Flacourtiaceae, with several big trees with well known vernacular names, is curious. Alangiaceae, Proteaceae, Symplocaceae and ‘Theaceae would also all have been expected in Ulu Kelantan. It is a well known weakness of Forest Dept. survey teams in Malaya to assign an unknown tree to either medang or kelat, and it is likely that trees of these missing families, and one must assume Flacourtiaceae too, have been mistakenly included in Lauraceae or Myrtaceae and thus contribute to the unexpectedly high figures for these two tamilies. 200 Gardens’ Bulletin — X XVI (1973) Basal Area and Tree Numbers per Acre Sedimentary rock has a higher basal area of trees > 4 ft girth per acre than granite, due to the much lower basal area per acre on granite over 750 ft than elsewhere (Table 3). In Ulu Kelantan (as shown below, Table 9) the important hill forest timber species Shorea curtisii is relatively rare on granite; doubtless this contributes to the low basal area. It can be seen from Table 3 that stems per acre are more numerous above 750 ft than below. These figures reflect a bias in the sample. In the hills big trees tend to be commonest on ridges and sparse on hill sides and in valleys and it can clearly be seen on Table 4 that survey lines above 750 ft have tended to follow ridges rather than to cut across the lie of the country, which becomes increasingly difficult with increasing elevation. The slightly larger number of stems per acre on low sedimentary rocks than on low granite, and other slight differences in Table 3 may not be statistically significant. An analysis of variance would be required to demonstrate that they are not due merely to chance variation in the data, and this has not been attempted because of the difficulties involved in extracting the figures from the survey summaries. HABITAT PREFERENCES OF SPECIES AND GROUPS Limitations to the Analysis The main analysis with which we are here concerned is that which attempts to elucidate how far individual species are confined to particular habitats, in this case defined on altitude and underlying rock. Two main weaknesses of the analysis have already been pointed out; firstly, that the forest data have been arranged in stands many of which cut across the boundary at 750 ft between lowland and hill forest; secondly, that the forest staff do not recognise many individual species outside Dipterocarpaceae but do recognise genera (e.g. petai: Parkia; sepetir: Sindora, etc.) generic groups (e.g. mempening: Lithocarpus, Quercus) or whole families (e.g. nyatoh: Sapotaceae). The result of this second weakness is, as of the first, an obfuscation of revealed habitat preferences. Individual species (or indeed groups within them) must be expected to show the greatest specificity for habitat, and to lump species together can only lead to blurring. A third and serious source of weakness is that the rocks have only been divided into the two great categories granite and sedimentary. There are consider- able variations in the granites of Malaya, often over very short distances, in texture and composition. No attempt has ever been made to map these. The sedimentary rocks are even more varied and include, in Ulu Kelantan, the pre- Carboniferous Teku schists in the north, which are the only large area of schist in the country, plus large areas of predominantly argillaceous Permian sediments and of predominantly arenaceous Triassic sediments. The various sedimentary rocks are usually interbedded and have in places been metamorphosed, especially along the boundaries with the granite. In places they are variously calcareous. No record was made of the rock type of the survey lines, indeed it is unlikely to have been homogeneous along their length, and the published geological records are not sufficiently detailed for high precision. It is for these reasons that the analysis is restricted to granite versus sedimentary rocks, as the boundary between these two major types is believed to be well and accurately known. Tree Species in the Rain Forest of Ulu Kelantan Table 2. Frequency of Families Leguminosae Myrtaceae Lauraceae Anacardiaceae Burseraceae Euphorbiaceae Olacaceae Fagaceae Sterculiaceae Guttiferae Moraceae Sapotaceae Sapindaceae Se A Se a ee Lee ce I ie eee cee ee i en Annonaceae ys Dilleniaceae " UMyristicaceae 18. Polygalaceae 19. Tiliaceae 20. Apocynaceae 21. Lecythidaceae 22. Bombacaceae 23. Thymelaeaceae 24. Rutaceae 25. Celastraceae 26. Combretaceae 27. Ebenaceae 28. Rhizophoraceae 29. Hypericaceae 30. Meliaceae 31. Loganiaceae 32. Rubiaceae 33. Simaroubaceae 34. Juglandaceae 35. Malvaceae 36. Ulmaceae 37. Melastomataceae 38. Oxalidaceae 39. Coniferae 40. Magnoliaceae Theaceae sat { Linaceae Dipterocarpaceae . . average 634 161 122 76 73 72 35 41 39 37.8 wid 33 af 24 20 14 14 122 10 8.0 1.0 Ts 5.0 4.1 3.1 3.0 2.8 2.0 1.9 1.7 1.6 1.5 1.3 0.6 0.5 0.2 0.2 0.9 .04 .02 .0006 .0006 201 granite sedimentary low high low high 439 602 587 815 139 153 182 140 73 122 124 i33 57 94 4 68 25 69 64 109 48 90 67 66 50 67 39 71 47 35 46 36 40 60 30 31 41 69 37 37 21 33 38 47 34 43 29 30 27 40 33 22 24 24 a | 20 16 29 14 21 11 14 iF 8.9 23 16 14 9 ‘| 8 16 12 13 8 12 10 6.1 9.8 Ts 7.9 11 6.3 8.8 5.9 12 i 53 3.1 1.8 6.0 6.2 2.8 3.3 1.8 6.8 is 0.9 1.3 eres 4.6 0.9 3 4.2 1.8 1.8 2.9 2.0 4.3 1.8 1.3 2.5 2.3 — 20 1.9 13 3.6 2.9 1.6 = 1.8 0.2 1.8 0.8 1.8 2.6 0.6 1.8 —- 11 1.4 1.8 -— a2 -— — 4.9 0.5 0.1 -— — 0.5 0.1 — —- 0.4 0.1 — 0.9 0.2 1.3 0.8 = pe _ 1.8 _- —- 3.9 — = -—— 0.1 — —— — 0.1 — In Tables 2 and 5-14 the figures given are for stems per 100 acres. 202 -Gardens’ Bulletin — XXVI (1973) Table 3. Basal area and tree numbers per acre granite sedimentary low high low high Basal area (sq ft) i <2 A Fe iP 47.9 52.9 56.7 average each rock .. af ae - 49.5 54.3 overall average 2M bd B - Se 52.7 Stems per acre. . a se v So bee 17.3 14.9 17.3 average each rock .. at 4 ie 14.6 15.8 overall average 7 bu ae bi 15.9 Table 4. Representation (as percentage of total acreage) of different topographic sites granite sedimentary ao ST SO ge LS Fe — Both rocks we, | 0- 500- 1,500- 2,500- 0- 500- 1,500- 2,500- Coral’ “aititude ihye Meah son 1300. 7.500 4.00) MO” sah Sar i. they &%. nie. & 8 eet | Wee ee gine J 67.\.. hillside fcc. top4PZedeoBOee. Te avi TneomtpZQa 1ftisn ollidestiiateeeeee 22) nadkdgecepyidel8ad wideaigine ask. 23 24. te Fe ee area sampled (acres)... dx. -—i.. 41.8-u2518+ 91Sruns48dcanc a@ienaeeda ie 0 based on an unpublished analysis by P.F. Burgess of 1087.1 out of the total 1,672 acres. A fourth potential source of weakness is that valleys, hillsides and ridge crests might have been sampled in different proportions in the four classes, and if species have topographic preferences a bias would emerge in the results. Examin- ation of an unpublished analysis by Mr P. F. Burgess of 1087.1 acres (out of the total 1.672 acres), see Table 4, shows that, fortunately, the proportion of the topographic classes does not vary largely either with rock type or altitude for the two altitude groups 0-750 ft and over 750 ft considered here. But, because there are differences of the order of 10 percent, comments on the present analysis are restricted to major differences in species’ density. It has been noted already that the area of low granite sampled was disproportionately low, and this is borne in mind in the comments which follow. All these limitations must be borne in mind in the analysis which follows, which is presented as a series of tables showing stems per 100 acres. Individual species are shown as far as they were recognised (the Dipterocarpaceae dominat- ing the lists), then genera (either straight from the survey data or by adding together species’ frequencies), then families. It is interesting to see how far genera and families do show habitat preferences, and whether these support the pre- conceptions forest botanists have about such things, though in general these big groups cannot be expected to show much uniformity in habitat. Tree Species in the Rain Forest of Ulu Kelantan 203 It must also be remembered that only trees 5 4 ft. girth were sampled. The effect of this may be illustrated by considerating two genera of the Sapindaceae. Xerospermum (gigi buntal) is actually far commoner in the forest than Nephelium lappaceum (rambutan hutan), but quite the reverse is shown in the result of this survey, because its trees rarely attain 4 ft girth and so only a few exceptionally large individuals have been counted, whereas rambutan hutan grows to a larger tree and so a larger fraction of the total population of the species has been included. In general little is known about species or group preferences for sedimentary or igneous rocks in Malaya as this is the first analysis made of a large body of data. Comments on the tables are therefore restricted to particular points which confirm or conflict with our existing subjective impressions. In fact, rock type itself probably exerts a less direct controlling influence on the distribution and frequency of plant species than the nature of the relief and of the soil (which is partly dependent on relief) which themselves are not directly related to rock type as expressed in this analysis. In the hot, wet climate of Malaya all soils are very strongly leached and consequently poor in plant nutrients; physical soil properties are probably usually more important to plant life than chemical ones, notably the depth to which the rock has weathered and the particle sizes formed by weathering. Granites tend to weather very deeply and to form well-structured soils, such as are seen along road cuttings in the Main Range, though quartz-rich areas may yield very shallow and coarse soils. Sedimentary rocks tend to weather less deeply, especially where metamorphosed; shale-derived soils are much more finely textured than sandstone-derived soils. Further, the relief of granite country tends to be less rugged than of sedimentary where precipitately sided knife-edge crests often replace broad, rounded ridges. The task of collecting detailed data on topography and soils, such as would be needed for a fully analysis, would be formidable in rain forest country, and has never been attempted over more than a few acres. On small areas the problem of correlating plant distribution with habitat is con- founded, in these excessively species-rich forests, by the probability that there is not enough space in a small area of any particular habitat for all the species to grow which are ecologically suited. The Analysis The first important conclusion results from a glance at Tables 5-12. It can be seen at once that nearly all species, except some of the rarer ones, occur in all four habitats, but that the frequency of occurrence varies in nearly all cases. This shows how misleading qualitative impressions of species’ distribution can be, and how important it is that ecological analyses of the rain forest are not based solely on presence or absence. Groups showing a marked preference for granite. (i) Below 750 ft, Table 5: Hibiscus floccosus (kangsar) and Pterygota horsfieldii (kasah) are valley species not known previously to prefer granite. The genus Hopea is usually considered commonest at high elevations; the preference here for lowland forest is unexpected. 204 Gardens’ Bulletin — X XVI (1973) Table 5. Groups showing a marked preference for granite and below 750 ft elevation granite sedimentary @) low high low high a) species Dipterocarp. Shorea assamica .. uh JO9 if 0.2 3.3 = S. kunstleri i] 0; , fiom 8.7 re» = Euphorbi. Endospermum malaccense Pe im eee 12 8.7 7.6 Maly. Hibiscus floccosus . . af s: 4.9 0.5 0.1 -- Mor. Artocarpus rigidus. . fd og 43 8.0 7.1 5.6 A. scortechinii (elasticus) dag ue 7.9 5.6 2.4 1.5 Sterculi. Pterygota horsfieldii fi. s 5.3 2.9 0.4 0.3 (b) genera Dipterocarp. Hopea . . it a - 1G 3A 27 22 18 Legum. Parkia . . au aft pea sblg 11 7.3 6.1 (c) families Myristicaceae B, 4 a iO 223 16.1 14.4 9.4 All the species of the forest survey are included in the tables 5-12 except for a few obviously mistaken identifications. (ii) Above 750 ft, Table 6: Shorea platyclados (meranti bukit) is the highest growing Malayan Shorea and a very important timber tree of upper dipterocarp forest from 2,500—4,000 it; it is locally gregarious; its almost complete absence from sedimentary rocks in Ulu Kelantan is certainly not true of Malaya generally. Inspection of Table 4 shows that none of the subsample Burgess has analysed is on sedimentary rocks over 2,500 ft, and the result for meranti bukit is therefore almost certainly an artefact due to sampling bias. Table 6. Groups showing a marked preference for granite and above 750 ft elevation granite sedimentary low high low high (a) species Dipterocarp. Shorea platyclados a3 . eh 73 0.4 4.3 Rubi. Anthocephalus chinensis v% ae 1.8 pay 0.3 1.8 (b) genera Anacardi. Campnosperma .. me . 2.6 6.9 2.0 1.3 Jugland. Engelhardtia te! * 5 err ee But — — Sterculi. Heritiera : Mey i 25 11 10 (c) families Annonaceae B “ 7 i (GS 29 14 21 Anthocephalus chinensis (cadamba) (kelempayan) is a species of valleys and alluvial soils in Borneo; we have noticed it common on hill slopes, e.g. at Genting Sempah, and this preference for hill forest is borne out by this survey. Engelhardtia (dungun paya) is well known as a genus of hill and mountain forest in Malaya. Tree Species in the Rain Forest of Ulu Kelantan 205 (iii) Just granite, Table 7: None of the preferences revealed here was previously suspected. Table 7. Groups showing a marked preference just for granite granite sedimentary (a low high low high a) species Bombac. Bombax valetonii . a rae Pe 6.9 0.4 0.8 Dipterocarp. Dipterocarpus appendiculatus . . be 4.4 0.2 — — Sterculi. Pterocymbium javanicum TF 8.8 7.3 2.0 1.0 (6) genera — nil (c) families Bombacaceae ra ist a a 12 12 5.3 a Fagaceae . . is ve his ne 40 60 30 31 Meliaceae ret sik ee fe 3.6 2.9 1.6 — Sterculiaceae _ pe < es 41 69 37 37 Groups showing a marked preference for sedimentary rocks. (i) Below 750 ft, Table 8: Dryobalanops oblongifolia (keledan) is a riverine species. The other Dryobala- nops in Malaya D. aromatica (kapur) is known strongly to prefer sedimentary rocks; both are found mainly east of the Main Range. Mesua ferrea (penaga) is a species of poor soils and dry sites, this is consonant with its marked preference for sedimentary rocks. Table 8. Groups showing a marked preference for sedimentaries and below 750 ft elevation granite sedimentary low high low high (a) species Dipterocarp. Dryobalanops oblongifolia. . M 17 Chas 66 5.6 Shorea bracteolata . . rs a 19 2.9 21 13 S. multiflora - sre 2 16 11 30 25 S. parvifolia m. he be 91 93 127 95 S. singkawang he om me — — 3.9 0.8 Guttif. Mesua ferrea me MS “ 4.4 3.3 14 12 Magnoli. Aromadendron elegans ‘ia a — — 3.9 -— (6) genera Combret. Terminalia be. f. si = — 3.9 _— Dipterocarp. Dipterocarpus oe ane he 68 48 94 69 (c) families — nil (ii) Above 750 ft, Table 9: Shorea curtisii (seraya) is the most important timber tree of hill dipterocarp forest; its distribution seems to be correlated with sites liable to periodic drought, which tend to be mainly shallow, sedimentary-derived soils and hill ridges, as is amply confirmed by the present survey (P. F. Burgess, unpublished observations). Swintonia, Vatica and Calophyllum are well known to be commoner in the hills. Their preference for sedimentary rocks was not known, though all are known as genera of ‘poor’ soils or sites. 206 Gardens’ Bulletin— X XVI (1973) The preference of the genus Shorea and of the family Dipterocarpaceae for this habitat completely obscures the markedly differing preferences of the many individual species enumerated and reflects the abundance of S. curtisii. Table 9. Groups showing a marked preference for sedimentaries and above 750 ft elevation granite sedimentary @ low high low high a) species Dipterocarp. Shorea curtisii se! + 33 21 126 20 345 S. guiso 2% ” fy 7.9 11 11 16 (5) genera Anacardi. Rengas* IP se y 11 21 22 35 Swintonia y be Me. 9.7 27 28 63 Dipterocarp. Shorea - 8 = ; pee 468 372 653 Vatica .. oh aS ie 1.8 6.0 7.7 14 Guttif. Calophyllum a ie, Td 17 29 23 34 {c) families Anacardiaceae val ™ - 53 25 69 64 109 Dipterocarpaceae af: re me <5 Oe 602 587 815 Coniferae ne ae ne se i fee — — 1.8 * Gluta, Melanochyla, Melannorhoea, Semecarpus (iii) Just sedimentary rocks, Table 10: Koompassia malaccensis (kempas) is one of the commonest trees everywhere in Malaya. Its preference for sedimentary rocks was not suspected and may not be typical of the whole country. Myrtaceae is a family which, throughout Malesia, becomes prominent in ‘poor’ sites, its preference in Ulu Kelantan for hills and for sedimentaries is not unexpected. Table 10. Groups showing a marked preference just for sedimentaries granite sedimentary low high low high {a) species Dipterocarp. Dipterocarpus crinitus ra . 4.4 5.8 20 21 Shorea dasyphylla . . aso an 1.8 1.6 2.1 2.3 S. faguetiana ; aks as 0.9 0.2 1.5 poe S. macroptera a , + 0.9 Yi 3.3 S. ovalis ce ae le 26 33 51 34 S. ovata wd fs sa Meee — 14 1.8 S. sumatrana , a a — 0.8 0.3 Legum. Koompassia malaccensis - ne 36 39 66 45 Mor. Antiaris toxicaria m a — 0.1 0.8 Artocarpus integer . . ” << — 0.4 0.3 Olac. Ctenolophon parvifolius oe bas 1.8 2.0 5.7 4.3 (b) genera Celastr. Lophopetalum 331 £ 0.9 1.3 ee 4.6 Legum. Koompassia + aye =e 54 56 84 63 Polygal. Xanthophyllum ot ™ 7.0 7.6 16 12 Rhizophor. Pellacalyx 1.8 0.9 2S 2.0 (c) families Myrtaceae Lo is ae if 73 122 124 133 Tree Species in the Rain Forest of Ulu Kelantan 207 Groups showing a marked preference for forest below 750 ft, Table 11: Dipterocarpus oblongifolius (keruing neram) is a riverbank species. Pometia pinnata var pinnata (kasai) is, in Malaya, a valley species. Barringtonia (putat), as a genus, is commonest in valleys and along rivers. Euodia (pepauh) is nowhere common in Malaya. It has been considered a genus preferring the hills, an impression contradicted by this survey. Pentace (melunak) likewise has been considered mainly a hill genus. Table 11. Groups showing a marked preference for elevation below 750 ft irrespective of rock granite sedimentary low high low high (a) species Dipterocarp. Dipterocaropus oblongifolius . . se 09 — 0.4 -— Dipt. cornutus + kunstleri .. gn 48 eA 60 29 Shorea balanocarpoides wy ‘es 7.9 if 5.2 3.3 Olac. Ochanostachys amentacea bes a 20 4.4 15 5.6 Sapind. Pometia pinnata v. pinnata... fo 18 9.1 i 9.7 (6) genera Lecythid. —_ Barringtonia ¥ ae me 11 6.3 $s 5.9 Logani. Fagraea af Ai es 1.8 0.2 1.8 0.8 Oxalid. Sarcotheca OR supe .2sleupsant 8 3 pend Su pbbr Dt | hee kie supers nilo? ‘rstinof every, aneaigg irotillo? iand’ se 635 Hee rOA POIhYf eatoe xiviag ailuoinus ee Hiellgzes} able wisdimctmpynol om: 2 imrotitee mezicA a aba iat WSTSI 2 Mpa aclgpiqe jwise) zustoy dizad inset ‘sabe SHOES softy 8 otsaie i er: ‘ egnol mun T-2 sorisini = sBUDBV 9Emu hy aveisy fisogs sgn! mre @ sibomoint” SEitfitie: :oardalg “oninr . {ois Unfeitic otuain onioo eudisigiem 2QnbD tant O1-@ stars! ’: aide 7 | eee ae a ” . - ‘ = ay > +. “ oa n 2. sdivabol :aiieitia ginivss ,seanol mer OFA) syolagotaipmtatitse cor — re weseala dil osgnol fm 2 seTocine " + + » % poe & ." M oh Fr a < ac i ad guar wee a 8 2 x te er sey v » * : " ie a, _~ “ Samrong é he hae oe + Ps . K% @ © x < ‘ . v ee ODDAR MEAU CHEY Sones 9 » i 3 - 2 “**, @ Banteay ss ae + ee s A as Tory Chma | Y; 3 a 2 i - »* ro at? Ge « a ah “ - «a? we a* Seas Tohen Mean che weve Saez + « ” =e . a ae o A a is Phnom - PREAH VIHEAR Lom phat = % < Kulen + ae ogee e a «4° * Ayan As RATTANAKIRG 4 a“ Ye Beng.’ * Thine « + ~ Py @ SIEM REAP Viealea s ei ase sone baal Ps ¢ Es »* Me s me 2 Fadeeci ag = 6 tt << Ls @ oe . a e st Ue + * BATDAHMBARNG ae A « 7 « ail =r me ¢ ac ‘ Me + e Pailin f ; MONDOLMIAI ; ¢ : -j x “ oF + z x Saat 4taaa ie 6 WRATIE "s > a wean ee PURSAT ae fen menerom = Tt, ry x i° * Peam Leach oh at oh 7 % Prus x e as ag * “h +KG, CHHNANG id es ‘ - snvoel = x ¥ > e Ven eng i , ® Psi 4 Ws ® “a4 Bee rik SOOO + + ~ 4 é “ % + ¢ : ' ¢ iat meriany a . as 2 es & i e rs aa* 2 % Ba a ry? i % S aed * < C NT . es D hhemarak - ; 4 \ Vhowrrin villee Thma *, PuNoH Pb : 1000 Kr Pang * ke. SPU 4 i : ; ® & - 6 () 7 ae sO kleirem te KOH KONG + © Sapehle ° ae : Pasa @svay rithe +++ Limite des Khets (Province) la = A a +# * ni) oot ane 36 Sew sat is" Yo _wuthnet 6 tonlie 2 tl joolb « ze aodeteion vii sree | ove Yiniehg ofatiot oftiiving wikaueld ; pn Syproneoy oalt_ gait: zhese sidibe olf lo saysced (HONS ysl) esgelliv yelsM tne BHA woiisils® 10 ie Mlle Cena” by beliate Me ile j re PKs. Ju my j tae ¥ th q OM volar ‘a no bia werner tnt tooa A; ence el velbbR ab calle pps gnidém rol feeratind orl hedmnils aid A dime (B2de% sit | Je, WAR sodhsrta) ‘jak mV ohiqilis opal a) ‘ a chisen bees ligills at’ (CSB> esti) ignsdt> ie ake 0 mou ) bobs live le A at : am rte, ately producing im nat a “ ea a » o* Cn philnaandl i vv iene 0, kee SUED Sse ORR Ate id) wise sigaut nobret) ijn iit ae ih ersG wevask jie Australia: sts rile. i HH ris om pis ri a‘ , ee is Cabesiedrwit ox onosane Presi me ~s.* = +f at) dlace cor we __ Form ls nga ea dd +3 vives so Fis al Ms anal! y, producing. male cones, vn ial a nE®, i. ie Sy. i | ~ SAvoret or otter eabies of as nes, such a inst, 7%. corthare Mercier’ {3 oe": a-neeilled, (Davedicd, fram & on wre acre! tale ComveN, amp % ee ; iat | Korthalsia hispida Becc. in Malaya by J. DRANSFIELD Herbarium Bogoriense, Bogor Indonesia On 7th January 1973, while in Singapore, I accompanied Dr. E. A. Heaslett on a one day field trip to Gunung Panti in Johore, Malay Peninsula. Despite bad weather, we were able to observe about 55 taxa of Palmae in the forest of Panti Fast. This total indicates the richness of the palm flora: amongst those species observed was one novelty, a Korthalsia vegetatively distinct from all other Malayan species of Korthalsia. Within the genus Korthalsia in Malaya (Furtado 1951) there are two types of ochrea (extension of the leaf sheath beyond the insertion of the petiole); in one type such as that found in K. scaphigera Griff., K. echinometra Becc. and K. scortechinii Becc. (see fig 1 a, b) the ochrea is swollen and hollow and forms a spiny chamber around the leaf sheath of the leaf above, and is favoured by ants as a nesting place. The relationship of the ants with the rattan has been described many times, and it seems likely that particular ants are specific to different Korthal- sia species. In the other type as found in all other Malayan species of Korthalsia such as K. flagellaris Miq and K. rigida Bl. (see fig 1c, d), the ochrea is not swollen and forms a tightly sheathing tube around the leaf sheath of the leaf above: this type of ochrea may remain entire or becomes split or tattered, and may be armed or unarmed. The Korthalsia species new to the Malay Peninsula from Panti East Forest Reserve, however, has a third type of ochrea, a type found in four species (see fig 1 e): K. robusta Bl. of Sumatra. K. macrocarpa Becc. of Borneo, K. squarrosa Becc. of the Philippines and K. hispida Becc. of Sumatra and Borneo. In these four species, the ochrea is shaped something “like an elongated ass’s ear” (Beccari 1918): the ochrea is not sheathing, being open opposite the petiole, the two edges inrolling to produce an open-ended spiny tube, pale straw in colour, diverging from the axis at an angle of about 20 degrees. In K. squarrosa the ochreas are slightly different in that they do not diverge far from the stem, but the ochreas visible in Beccari’s plate (Beccari loc. cit.) are just below an inflorescence and such ochreas are often slightly abnormal. Within this tube-like ochrea in the Sumatran and Bornean species, ants make their nests. When the rattan is touched, the ants within rustle in unison in each ochrea by banging their heads against the dry ochrea, each ochrea slightly out of phase with the next. I have only heard this rustling noise in K. robusta, K. hispida and K. macrocarpa; I have not heard it in K. scaphigera or K. echinometra and it is quite likely that the ant species in these last two are different. 239 240 Gardens’ Bulletin — XXVI (1973) The Gunung Panti plant is indistinguishable from Korthalsia hispida, a mis- understood species from West Sumatra, Aceh, and Borneo, first collected by Beccari (P. S. 673 — Beccari 1884) at Ayer Mancur, West Sumatra, and later included by Beccari in Korthalsia robusta Bl. non Mart. (Beccari 1918). In July 1972 in Propinsi Jambi, Sumatra, (Dransfield in press) I found specimens of a Korthalsia, vegetatively fitting the description and plate of K. hispida but with inflorescences totally different from those of the widespread Sumatran taxon, K. robusta, the vegetative parts of which were unknown to Beccari. The Jambi specimens are identical to Meijer 2411, Nunukan, Borneo (Meijer 1956). There are hence two species in Sumatra, K. robusta and K. hispida, and Beccari appears to have confused the two in his monograph (1918) — the fertile specimens referred by Beccari to K. robusta belong to K. robusta; the vegetative specimens referred to K. robusta belong to another species originally described as K. hispida. In fact vegetatively K. hispida is more distinct than K. robusta and K. macrocarpa and it is quite possible that these last two represent one widespread variable species common to Sumatra and Borneo. Because of the confusion in this group of Korthalsia it is considered useful to give a complete description of Korthalsia hispida. Although the relationship of Korthalsia macrocarpa and Korthalsia robusta is not yet clear, I am presenting this note in order to draw attention to this easily recognized group of Korthalsia species. Korthalsia hispida Becc. Beccari, in Malesia //, (1884) 71., Beccari, Ann. Roy. Bot. Gard. Calc. XII, 3 (1918) 148. Plate 99. Meijer, W. Penggemar Alam 36 (1957) 60, Fig 3. Slender, clustering, thicket-forming rattan, branching at ground-level. Stems to 1.0 cm in diam. without sheaths, with sheaths 1.5—1.8 cm. Internodes 20-25 cm long. Sheaths bright shiny green when fresh, covered with sparse, scattered, shiny black spines to 2.0 cm long, easily breaking just above the base and minute c. 0.5 mm long, easily detached spiculae. When young, sheath covered with chocolate coloured scurfy indumentum. Ochrea 25-30 cm long, like a very narrow elongate horn, diverging at an angle of c. 20 degrees from the axis, shiny brown within, straw-coloured without, thorny-spiculate and indumentose as the leaf sheath, truncate and shallowly bilobed at the apex, papery in texture, open opposite the petiole, rolled, often filled with vicious biting ants. Leaf to 1.8 m long: cirrus to 90 cm, petiole to 10 cm. Petiole and rachis yellowish-green, semi-circular in cross section, to 5 mm wide, armed with scattered, reflexed black spines below, above, spines in groups 3-5 forming claws, separated by c. 3 cm, distance of separation decreasing above to c. 5 mm at tip of cirrus, Leaflets 7 on each side, alternate below, subopposite above, separated by 7-10 cm, with ansae 6 mm long, the lower two leaflets narrowly cuneate-rhomboid, to 3 cm broad, upper leaflets broader, 15-20 cm long by 7-9 cm broad, irregularly plicate, longly acuminate praemorse at upper margins, bright green above, grey-white indumentose below. Leaves sub- tending the terminal inflorescence much smaller. Inflorescence system lax, terminal, single stems dying after flowering. the upper 3-8 nodes producing partial inflorescences. Partial inflorescences to 30 cm long, emerging through splits in the upper leaf sheaths; prophylls and other bracts tubular below, variously split and tattered above, 5-10 cm long, straw-coloured, papery, densely covered in chocolate-coloured indumentum and small spiculae. Bracts on axis subtending branches, each terminating in rhachillae, 2-6 in number. 241 Korthalsia hispida in Malaya 10cm OCHREA TYPES IN KORTHALSIA Fig. 1 on- ida, e) piny, d) K. sp. (? juvenile of K. rigi shaped ochrea: b) K. echinometra. N.B. hole made by ants. N , Central Sumatra) — ochrea unarmed. Horn- =| cs & rs} a) Pp aa) on & S oo = Q i fe) on ° 2) [aa] aoa sig _& oS 'g | 3 OQ ”~ oO S.8 5M SE _F = & =<: © sagh o> mn) vt ( @ Tae hows 3 Miu 5 N Braces VANE Saas a) q aeBGs SL25 oY Ss) ogs SaQ =2_s 53. nASHM 242 Gardens’ Bulletin — X XVI (1973) Rhachillae pendulous, to 15 cm long, 1.2 cm in diam, clothed in a tight spiral of bracts, adnate laterally, each bract subtending a condensed branching system with one hermaphrodite flower. Bracts to 1 cm wide and 5 mm high, pale brown with deeper brown fugaceous scurfy indumentum, irregularly split and lobed at the apex. Flowers pale brown in colour, exserted between the tight bracts. Sepals 3, free, imbricate, ovate, shiny, straw-coloured, to 3 mm long. Corolla with a tube c. 3 mm long and 3 free imbricate rounded triangular lobes above, 4 mm long and 3.5 mm wide, during growth of the ovary after fertilization, the corolla tube splitting and being carried out of the bract axils on the tip of the ovary, at the base of the style. Anthers 6, 2 x 1 mm, borne on a short staminal tube, this shortly epipetalous. Pollen pale yellow. Ovary at anthesis, to 2 mm high, with style to 4 mm, chestnut brown, scaly. Style grooved longitudinally and tipped with 3 minute approximate stigmatic lobes. Fruit when not damaged crowned with remains of style, surrounded by corolla and stamens, at least 1.2 cm long, and 1 cm wide. Mature fruit unknown. Specimens examined Type not seen, though Plate thereof in Beccari (1918) examined. SUMATRA Aceh: Nainggolan s.n. April 1931, Bukit Plawi (BO). Jambi: Dransfield 2620. 22.7.72. Kampung Penetai, Kabupaten Kerinci, streamside, Hill Dipterocarp Forest, alt 200m (BO, L, BH). BORNEO East Kalimantan: Meijer 2411. 5.12.53. Nunukan, near S. Binusan. dry lowland Dipterocarp Forest (BO). MALAY PENINSULA Johore: Dransfield 3037. 7.1.73. Gunung Panti East. Lowland, reshwater swamp forest. (SING, BO, L, BH). Korthalsia hispida differs from the other species having long horn-like ochreas, in the hispid appearance of the sheaths, ochreas and inflorescence bracts, produced by the dense minute spiculae, and in the lax, not approximate, slender infiorescence branches. K. macrocarpa, K. robusta, and K. squarrosa all have squat inflorescences made up of robust, fat, approximate rhachillae, unlike any other Korthalsia species (see Beccari 1918). The presence of this group of long-ochrea Korthalsia species in Malaya is not surprising; the absence of collections of it, is, especially as it grows in a very famous, much-visited botanical locality. Acknowledgements I should like to thank Dr. E. A. Heaslett for introducing me to Gunung Panti East, and Ahmad Shukur and Samsuri, plant collectors, Botanic Gardens, Singapore for their help in collecting palms on Gunung Panti: thanks are also due to Sdr. Damhuri, draughtsman, Herbarium Bogoriense, for preparing the figures. Korthalsia hispida in Malaya BAKO CARELESS 0) aay) ae ee Ae ~ ad) in »S Pesan pics eR SN on vig sae SOE SAK ALA cy RANE Deas Wek Ph aw Ves, “sr¢ A Py > Ae ella eat Fig. 2. KORTHALSIA HISPIDA BECC. 243 Illustration of a node and most of one internode and the subtended partial inflore- scence, taken from near the apex of a flowering stem; in all 3, nodes were producing partial inflorescences. N.B. the hispidity on all sheaths, ochreas, and bracts, Dransfield 2620, Jambi, Sumatra. Drawing by Damhuri. 244 Gardens’ Bulletin — X XVI (1973) References Beccari, O. (1886): Piante Ospitatrici, Malesia 2, 62-78. Beccari, O. (1918): Asiatic Palms — Lepidocaryceae. Part 3. The Species of the genera Ceratolobus, Calospatha, Plectocomia, Plectocomiopsis, Myrialepis, Zalacca, Pigafetta, Korthalsia, Metroxylon, Eugeis- sona, Ann. R. bot. Gard. Calcutta 12. Dransfield, J. (in press): Notes on the Palm Flora of central Sumatra. Reinwardtia & (4). Furtado, C.X. (1953): Palmae Malesicae 11. Korthalsia. Gdns’ Bull., Singapore 13 300-324. Meijer, W. (1957): Rotans van Nunukan. Penggemar Alam 36 51-64. Pollen Viability and Germination in Some Orchid Hybrids by A. N. RAo and LEE War CHIN* Department of Botany University of Singapore Introduction Singapore has become an important orchid breeding center producing many well-known hybrids of international acclaim. Such successful results are achieved by the continued efforts of scientists at Botanic Gardens and local horticulturists over a period of three decades. In Singapore the first hybrid that appeared was Vanda Miss Joaquim in 1893 (Holttum, 1953). Nevertheless, our knowledge about the cytogenetics and pollen physiology of local orchid hybrids is very limited. The importance of such studies need not be over-emphasized since pollen viability and germination form the essential requirements to produce good and the required hybrids. Not every cross pollination will bear fruit and it is the common experience of every orchid breeder that more than 60 per cent of the pollinations made would end up with no fruits or fruits without seeds. One of the main reasons for such failures could be the pollen sterility and some of these problems are under study (Rao and Goh, 1970). Very few studies are made on orchid pollen germination. Molisch determined the optimal sucrose concentration for 6 European species (Withner, 1959). Miwa (1937) germinated the pollen of certain orchid species and their hybrids to test the longevity and viability of stored orchid pollen. Curtis and Duncan (1947) studied 11 species in 7 genera. The present paper outlines the morphology of pollinia, pollen structure, viability and germination of some well-known local hydrids. Materials and Methods Pollinia of the following hybrids were collected from Mandai Gardens now known as Singapore Orchids Pte. Ltd.: (i) Arachnis Maggie Oei var. Red Ribbon, (ii) Arachnis Maggie Oei var. Yellow Ribbon, (iii) Aranda Hilda Galistan, (iv) A. Lucy Laycock (long spray), (v) A. Lucy Laycock (short spray), (vi) A. Wendy Scott var. Greenfield, (vii) Aranthera Lilliput, (viii) Vanda Poepoe and (xi) Vanda Josephine (from the garden at Botany Department, University of Singapore). Fresh pollinia from open flowers were collected between 2-3 p.m. and used within 1 hr. after collection, using half a pollinium in each case. The following culture media were used: (a) distilled water as the control medium (5) stigmatic extract — the appropriate number of fresh stigmas were ground in 10 mls. of distilled water and the clear filtrate of the mixture was then used for germination. The number of stigmas used in such preparation varied from * Currently at Botanic Gardens, Singapore. 245 Pollen Viability and Germination in some Orchid Hybrids 247 Figures 19-25: Pollen germination in Vanda Josephine. Fig. 19: in dist. water, note ihe short tubes. Fig. 20: in stigmatic medium at 2 stigmas/10 mi of dist. water; all 4 grains in the tetrad germinated. Fig. 21: in 4% sucrose a & b different sized pollen tubes with tapering ends. Fig. 22: in salt-boron medium, a— pollen tubes typical match-stick shaped, some with vacuoles; b — short tube, with broad ends. Fig. 23: in GA (10-7 mg/L) medium short pollen tubes with exudates (arrow). Fig. 24: in borax medium, with all the grains in tetrad germinated. Fig. 25: in kinetin (10-6mg/L) medium with pollen exudate (arrow), See opposite page Figures 1-18. Figs. 1-9: Pollinia of different orchid hybrids. Figs. | & 2: Arachnis Maggie Oei var Red Ribbon and var. Yellow Ribbon respectively. Figs. 3-6: Aranda Hilda Galistan; Lucy Laycock (long spray); Lucy Laycock (short spray); Wendy Scott var. Greenfield respectively. Fig. 7: Aranthera Lilliput. Fig. 8: Vanda Poepoe. Fig. 9: Vanda Josephine. a-column, b-} stigma, c-pollinia. Figs. 10-18: Tetrads showing microspore arrangement and supernumerary spores. Fig. 10: Arachnis Maggie Oei var Red Ribbon, Fig. 11: A. Maggie Oei var. Yellow Ribbon. Fig. 12: Aranda Hilda Galistan. Fig. 13: A. Lucy Laycock (long spray), Fig. 14: A Lucy Laycock (short spray). Fig. 15: A. Wendy Scott var. Greenfield. Fig 16: Aranthera Lilliput, Fig. 17: Wanda Poepoe. Fig. 18: Vanda Josephine. 248 Gardens’ Bulletin— XXVI (1973) 3, 1, 2, 3 stigmas/10 mls. of distilled water. In some cases all the four concentrations were used, in others only 1 or 2. The total length of the stigmas ranged from 3-8 mm in different hybrids. Their sizes and weights are: Length of Wt. of Plants stigma I stigma (in mm) (in mg) Arachnis Maggie Oei var. Red Ribbon 8 198 A. Maggie Oei var. Yellow Ribbon 8 180 Aranda Hilda Galistan 5 101 A. Lucy Laycock (long spray) 5 96 A. Lucy Laycock (short spray) 5 109 A. Wendy Scott var. Greenfield 6 87 Aranthera Lilliput 3 16 Vanda Poepoe 8 84 V. Josephine 8 85 (c) Sucrose medium with varying concentration of 1-10%. (d) Borax medium — Sodium tetraborate was used to prepare the media with concentration ranging from 10° to 10 mg./L. (e) Salt boron medium (modified Knop’s medium). Formula: H;BO,; — 100, Ca(NO;)2.H,O0 — 300, MgSO,. 7H,O — 200, KNO; — 100mg/L. (f) Different growth substances — well known auxins like: Indole — 3 — Acetic Acid (IAA), Indole — 2 — Butyric Acid (IBA), Indole — 3 — Propionic Acid (IPA), o¢ — Naphthalene Acetic Acid (NAA), 2,4 — Dichlorophenoxy acetic acid (2,4-D), 2,45 — Trichlorophenoxy acetic acid (2,4,5-ITPA), source of gibberellin Gibberellic Acid K salt (GA), and source of Kinin — Kinetin (K) were used in the concentrations that ranged from 10° to 10 mg/L. Either all or only certain concentrations were found to be effective. Solid watch glasses (4 x 4 x 1.75 cm) were used to germinate pollinia. The pollinia were cut into small pieces and dispersed in | ml of the medium. A glass cover was used to prevent evaporation of the medium and the watch glass was placed in a humid chamber to prevent evaporation and condensation. Diffuse light of 235 F. C. S. (during day-time) and temperature of 28°C prevailed in the labora- tory. Duration of experiment was limited to 24 hours since most pollen did not germinate or their pollen tubes did not grow after this period. Each experiment was repeated at least twice. At the end of 24 hours, the percentage germination of pollen tetrads and pollen cells, pollen tube length and morphology were studied. Lactophenol with cotton blue was used for staining. At least 225 tetrads were studied for each treatment to assess the percentage germination and 90 tubes for tube length determination. Observations and Results Pollinia and pollen grains: In orchids the pollen tetrads develop into mature structures called compound pollen grains and these are united to form pollinia. In all of them two pollinia are present per flower (Figs. 1-9). The pollinia of different hybrids were excised for comparative study. In size the Aranthera pollinium was smallest among the 9 hybrids studied (Fig. 7). Pollinia of Aranda hybrids were smaller than those of Arachnis or Vanda (Figs. 1-6, 8, 9). The disc was flat in Arachnis, triangular in Aranda, and of irregular shape in Vanda. Even among the Pollen Viability and Germination in some Orchid Hybrids 249 Table 1. Morphology of the pollen tetrads, size and arrangement of pollen cells in them. Morphology of Pollen Tetrads Nittinos of Orchids . Size, ‘ica he grains in tetrads Oo isobi- tetra- . i SLB lateral. | hedeal T-shaped) linear Arachnis Maggie Oei var. Red Ribbon... | 37 x 23 1.61 92 6 1 1 Arachnis Maggie Oei var. Yellow Ribbon | 40 x 27 1.48 90 8 zZ — Aranda Hilda Galistan i! - i |e oy 2&5 dee 82 16 — ys A. Lucy Laycock (long spray) can Sa 2a TAZ 72 Ze 2 4 A. Lucy Laycock (short spray) oe Loe Ok oe ee 88 8 2 2 A. Wendy Scott var. Greenfield vets & 22 eS Most of them are supernumerary spores. Normal tetrad — _ only about 1% Aranthera Lilliput “t ye lA ee 20 Sua 89 9 1 1 Vanda Poepoe .. a ere ~~ 2 b6 68 32 1 1 Vanda Josephine «t Re ee aes ay 72 ad 1 oe L — Length of pollen tetrad B — Breadth of pollen tetrad L/B — Length /breadth ratio. Aranda hybrids the pollinia of each hybrid were distinctly different from the others in having disks and stalks of different shape and size (Figs. 3-6). The pollinia were teased to separate the tetrads. These are diagrammatically represented in Figs. 10-18. Each figure shows the variation in size, shape, arrangement as well as the number of cells in each tetrad or the unit (is one which had more than 4 cells). In addition, the cells of the tetrad or unit also varied in size. Some of them e.g. Vanda Miss Joaquim, Vanda Poepoe, Aranthera Lilliput had one small and three big grains or vice-versa (Figs. 14, 16-18, 22, 28, 29). Two big and two small grains were also common in some (Figs. 13, 22). Supernumerary spores were also common in many (Figs. 12-14, 16, 18) and the individual unit had 5, 6 or 8 cells. In such units, which had supernumerary spores, at least two of them were big and of comparable size with the spores of the majority of the tetrads and the remaining were generally small (Figs. 12, 13, 18). Pollen tetrads show isobilateral, tetrahedral, linear and ‘T’ shaped arrangement. The percentage occurrence of each type is recorded in Table 1. Comparatively, the two latter conditions were rare and the isobilateral type was most common. The average size of pollen tetrad ranged from 28 x 28 » to 42 x 27 uw. The linear tetrads were much smaller in size when compared with the others (Figs. 12, 14, 16). Pollen germination: Positive responses resulted in the pollen germination of Vanda hybrids and these details will be considered first, followed by the results of other hybrids. (a) Distilled water (control medium). In V. Poepoe and V. Miss Joaquim 27 and 30% of the tetrads germinated with an average tube length of 15 and 19 yp respectively (Tables 2, 3; Figs. 19, 26, 30, 31). 250 Table 2. Gardens’ Bulletin — XXVI (1973) different media after 24 hrs. at room temperature. No. of grains germinated per tetrad, in % Vanda Poepoe. Percentage germination and average pollen tube length in Media Concentrations %G pL 1 2 3. jattat Distilled water 18 5 2 2 27 15 (control) | 1 stigma /10 mis. 20 21 14 10 65 57 distilled H2O Stigmatic 2 stigmas /10 mls.| 27 30 13 13 83 68 extract distilled H2O ee “We | =s 3 stigmas /10 mls.| 21 20 8 4 53 44 distilled H20 pe 28 20 4 12 64 56 at 23 25 24 13 85 75 : | Sucrose 6% 8 16 24 41 89 83 8% 15 11 12 | 12 50 | 48 Le) eee seh? oe ia at | tthe 10% 11 9 | 11 8 49 40 | 1x 10—5mg./L| 26 6 2 BO paige bh ag atl iw 320 fi Hiawft 29 pial salle aoe 1x 10—4mg./L 20 12 2 2 | ey pees 'Yivo Soe ee} SAAS Gee oh ee | / 1x 10—3 mg./L 27 10 0 2 39 | 36 Borax 1x 10—2mg./L 20 14 4 2 40 36 1x 10—1mg./L 2a 12 8 2 47 38 1x 10—° mg./L 24 16 8 13 50 55 1x 10—! mg. /L 16 14 8 z | 40 37 Salt-boron medium ZZ 32 10 8 he 72 % G — Perc. germination pa L — Avg. pollen tube length in y, Pollen Viability and Germination in some Orchid Hybrids 251 Table 3. Vanda Poepoe. Percentage germination and average pollen tube length in optimal concentrations of different media, after 24 hrs. at room temperature. No. of cade: 3 per E etrad, in | Media PY le a : 70G pl | 1 2 3 4 Distilled water | 18 5 2 2 ZY 15 (control) 2 TAA 11x 10—6 mg. IL| My We td | taal nk 71 71 IBA \1x 10—5mg./L| 21 25 «| 45 es es hls IPA 1x 10—6mg./L| 24 17 iaab 3 ss | al NAA 1x 10—4mg. 1 10-4 mg. JL 27 26 20 5 78 82 2,4-D fins SED i 10—6mg./L| 31 | 21 15 ala 60 2,4,5-TPA lay 1x t0—Sime/L |. 27 | 25 6 7 65 oll 40 GA ee TX 0? mall 3g 23 5 7 ia 75 Kinetin fix tae n | 24 15 9 | 7 | 55 * 34 G, % G — Perc. germination p. L — Avg. pollen tube length in be (b) Stigmatic extract medium: For both the hybrids the optimal concentration of stigmatic extract was 2 stigmas/10 ml. of distilled water. The percentage germination for V. Poepoe and V. Miss Joaquim was 83% and 79% with an average tube length of 68 and 87 » respectively and these results were better than in the control medium (Tables 2, 3; Figs. 20, 27, 33). The response at other concentrations was variable in terms of percentage germination. (c) Sucrose medium: For V. Poepoe the optimal concentration of sucrose was 6% and 89% of the tetrads germinated with an average tube length of 83 » (Fig. 28). In V. Miss Joaquim 92% of the tetrads germinated in 4% sucrose medium (optimal concentration) with an average tube length of 74 u (Fig. 21a, b). Other variations observed in media with higher or lower concentrations are given in Tables 2, 3 and in all the concentrations used the response was better than in the control medium. (zd) Borax medium: In media with different concentrations of borax better results than control were obtained in V. Poepoe. The optimal concentration for the two hybrids were 1 and 0.1 mg/L respectively (Figs. 24, 34) and the response of V. Miss Joaquim was much better than V. Poepoe (Tables 2, 3). (e) Salt-boron medium: When compared with control or borax medium the values obtained in this medium were better in the case of V. Poepoe (Table 2). In case of V. Miss Joaquim the percentage germination was almost like that of control but tube length improved. These results were poor when compared with those obtained in optimal concentration of borax medium (Table 3, Fig. 22a, b). 252 Gardens’ Bulletin — XXVI (1973) Table 4. Vanda Josephine. Percentage germination and pollen tube length in different media after 24 hrs. at room temperature. | | Media Concentration Distilled H2O (control) stigma /10mls. distilled H2O wI- —_ stigma /10mls. distilled H2O Stigmatic extract 2 stigmas /10mls. distilled H2O Ww stigmas /10mls. distilled H2O 2% 4% Sucrose 6% ft 10—3 mg. /L 1x 10—2mg./L Borax 1 x (10—1 mg. /L | 1x 100 aa 21 | [Ax 10 mg. /L No. of grains germinated per 15 tetrad, in % Gf el 2 3 4 5 on | | 5 2 0 36 | 19 | 11 2-01 x14 | 38 55 9 6 21 | «48 77 | | | 6 20 | | 7 jp YH | | in | 24 2 39 | 47 | ea.) ee | | 15 | 31 21 75 | 62 8 22 | 7-4-2 | 74 21 14 36 80 | 37 MESSE, A at a8! ae A 1 ‘2 1 | 20 | 21 Je AE oS Eee 0 0 0 11 14 17 | 0 | 1 46 | 27 ———__,—___|—____— 8 8 | 0 60 | 41 19 24 | 18 | 85 | 84 | | 13 goin of pu bay | 63 20 OG DONO SNOW Bi? eee 2 1 1 13 | 19 6 5 | 1 38 Se Salt-boron medium | % G — Perc. germination b& L — Avg. pollen tube length in » Pollen Viability and Germination in some Orchid Hybrids 253 (f) Media with different growth substances: Cultures in NAA (104 mg/L) showed the highest percentage germination than in other auxin media, attaining 74% and the longest tube length of 82 ». Effect of IBA was similar of that of IAA giving better results than 2,4-D and 2,4,5-TPA media (Table 4). In case of V. Poepoe for IAA, IPA, 2,4—-D and 2,4,5—-TPA the optimal concentration was 10° mg/L. Of these four auxin media the highest percentage (71%) germination and the longest tubes (71 ») were formed in IAA medium and the lowest percentage germination was obtained in IPA medium (55%); the shortest tubes were produced in 2,4,5-TPA medium (40 yp). Table 5. Vanda Josephine. Percentage germination and pollen tube length in optimal concentration of different media after 24 hrs. at room temperature. No. of grains germinated per | Media Pe iane a ‘va all %oG | pL 1 2 3 4 Distilled water 29 5 2 0 36 19 (control) IAA ee ie ee eee 12 12 7 64. | 43 IBA spe tie 10—5mg./L| 32 sal ab? ae ae 52 | 35 IPA 1x 10—5mg./L| 20 15 i 3 49 59 NAA Piet OO ae ie ie eh AL ee ee ae ae 47 eS. iV 10—Saie (Lee. | a8 it fue ae pea 51 2,4,5-TPA Pe HO— Smet | 3 hl. bk,.6 3 60 45 GA lt 10—Sme. fl |) 29 16 | 5 1 a Sa 39 Kinetin ie 10—6mg./L| 14 15 | 18 | 17 64 64 % G — Perc. germination p. L — Avg. pollen tube length in mM The optimal concentration for GA and K was 10° mg/L (GA), resulting in 61% germination and 75 » tube length and 55% germination and 34 yu tube length respectively (Table 4). The different auxin, gibberellin and kinetin media, at various optimal concentration, gave better results than the control medium. When compared with borax medium, poorer results were obtained in the two auxin media, i.e. IPA or 2,4,5-TPA. In all the others, including GA medium, better results were recorded. Kinetin medium was less favourable than the borax medium and similarly, gibberellin and kinetin media were not so favourable when compared with stigmatic extract or sucrose media (Tables 2, 3). In case of V. Miss Joaquim germination and elongation of pollen tubes in- creased appreciably in auxins, GA and K media, when compared with the con- trol or salt-boron medium. The optimal concentration was between 10° and 10*mg/L. Higher concentrations were inhibitory for germination and tube growth. This was true in all the media except in case of IAA where even at 0.1 mg/L the percentage germination and tube length were very similar to that of the control (Table 5). In IAA and kinetin media better germination resulted when compared with the other auxins or GA media. The media with auxins, GA or K (at optimal concentrations) were not so favourable as stigmatic extract, sucrose or borax media (Tables 4, 5). 254 Gardens’ Bulletin — XXVI (1973) Hybrids other than Vanda The pollen germination studies in case of: (a) Arachnis Maggie Oei var. Red Ribbon, (b) Arachnis Maggie Oei var. Yellow Ribbon, (c) Aranda Hilda Gallistan, (d) Aranda Lucy Laycock (long spray), (e) Aranda Lucy Laycock (short spray), (f) Aranda Wendy Scott var. Greenfield and (g) Aranthera Lilliput, gave very poor results in the different media used. In most of them no germination was seen with the exception of Arachnis Maggie Oei var. Yellow Ribbon the pollen of which showed small protrusions in the control medium. In borax medium there was good germination and pollen tube growth at two lower concentrations i.e. 10 (25% G, 11 p) and 10+ (61% G, 32 ») mg/L. In salt boron medium 52% germination and 29 » tube length resulted. In other media (stigmatic extract with 1, 2 or 3 stigma/10 ml of H, O; sucrose 2-10%) there was no germination. Like- wise, the different hybrids did not germinate in the auxin, GA or K media at various concentrations mentioned previously. Morphology of pollen tubes: Certain variations were noticed. In V. Peopoe a single case of branching was observed in NAA medium (10+ mg/L) and slight swellings of pollen tubes were common in NAA and 2, 4-D media (10° mg/L). Tips of pollen tubes were pointed in sucrose medium. In V. Miss Joaquim oil globules were present in the grains and pollen tubes. Tips or middle portion of pollen tubes were swollen in salt-boron and IAA media (Fig. 22a, 32) and forking of the tube was seen in kinetin medium (10° mg/L). Exudates were common in GA and K media (Figs. 23, 25). Discussion Structural variation of pollinia in different orchids has been described and the present study reveals the fact that hybrids retain the same genetic characters of parents (Adams, 1959). Further study of other hybrids may substantiate this fact since very few hybrids are critically examined from this point of view. Occurrence of pollen tetrads is common in Orchidaceae (Withner, 1959). The presence of supernumerary spores, frequency of their occurrence, and an analysis of their structure has been studied in the diploid and tetradploids of Vanda hybrids (Kamemoto, 1956). In five of the nine hybrids studied presently the supernumerary spores were present and it is interesting to record that only few of them germinated when compared with the regular tetrads. Of the nine hybrids investigated germination was seen only in Vanda Poepoe, V. Miss Joaquim and to a certain extent in Arachnis Maggie Oei var. Yellow Ribbon. Others did not germinate in the different media used. This loss of germinability may result from immaturity of pollen, long preservation, or it can be an inherent property resulting from complicated and forced hybridisation (Miwa, 1937). It is possible that the reason mentioned last is applicable here since fresh pollinia were used in all the experiments. Except Vanda, most of the other hybrids studied presently do not set fruits either by self or cross pollination. In control medium the percentage germination was low and tube lengths shorter in both the hybrids of Vanda, In those cases where the pollen grains germinate with short tubes, it is said that the grains contain a certain amount of nutrients that promote initial growth but this is insufficient for the growth of longer tubes (Brink, 1924a). All the growth adjuncts used presently, both organic Figures 26-35: V. Poepoe. Fig. 26: Germination in dist. water (control). Fig. 27: in stigmatic extract (2 stigmas/10 ml of dist. water), Fig. 28: in 6% sucrose medium, note beak like tubes. Figs. 29-35: V. Miss Joaquin. Fig. 29: Tetrads of different sizes. Figs. 30 & 31: Germination in dist. water (control) and one of the tetrads enlarged to show short tubes in Fig. 31. Fig. 32: Germination in IAA medium (10-5mg/L), showing pollen tubes with bulbous tips. Fig. 33: in stigmatic extract (2 stigmas/10 ml dist. water). Fig. 34: in borax medium (10-1! mg/L). Fig. 35: Aborted grains of different sizes. Pollen Viability and Germination in some Orchid Hybrids 255 and inorganic, were found to be stimulatory when compared with the control medium. Their effects on percentage germination and tube growth were of different magnitude. Better results were obtained for Vanda hybrids in stigmatic extract medium which indicated that the necessary stimulii required for germination and tube growth were present. The stimulatory effect of stigmatic extract on pollen germination was observed in other orchids studied earlier (Curtis and Duncan, 1947; Rao and Lee, 1972). Details pertaining to other angiosperms are also reviewed (Johri and Vasil, 1961). Sucrose undoubtedly had a stimulatory effect on germination and tube growth of Vanda hybrids. The optimal concentration varied from 4-6% and best results were obtained in sucrose media. This is in conformity with earlier findings of other workers (Miwa, 1937; Curtis and Duncan, 1947). Externally supplied sugars do serve as a source of nutrition for pollen germination in most of the species studied so far and the important role of sugars is vividly discussed (Johri and Vasil, 1961; O’Kelly, 1955, 1957; Visser, 1955). In case of V. Miss Joaquim, very good germination and tube growth resulted in boron medium. Vanda Poepoe showed an improvement both in germination and tube growth but the effect was not as good as stigmatic extract or sucrose media. Arachnis Maggie Oei var. (Yellow) Ribbon also showed good germination in the presence of low concentration of borax but the tube growth was poor. Salt-boron medium was more effective than borax medium in V. Poepoe and it was contrary in case of V. Miss Joaquim. In case of the latter the presence of salts reduced both the percentage germination and tube length, though boron was present as part of the medium. In case of V. Poepoe borax alone was not stimulatory but its synergistic action with the salts was obvious. Certain salts, especially calcium (ions), have a stimulatory effect on pollen germination in majority of angiosperms so far investigated (Brewbaker and Kwack, 1964; Mascarenhas and Machlis, 1964; Vasil, 1960, 1964). Negative chemotropic response to calcium ions has also been observed in Lilium, Zea and others (Rosen, 1968, Cook and Walden, 1965). The results obtained in Vanda Miss Joaquim support the above observations. The fact that pollen grains are naturally endowed with auxins is wellknown and in certain cases like Antirrhinum, Bryophyllum the auxin concentration is so high that it is inhibitory to germination (Curtis and Duncan, 1947; Smith, 1942; Vasil, 1960). In the two Vanda hybrids studied, all the auxins (concentration 10-’ to 10+ mg/L) had a stimulatory effect on germination and tube growth. The results obtained presently are similar to the observations made in other angiosperms like Cucumis melo, Eriobotrya japonica, Triticum and others (Loo and Hwang, 1944; Dikshit, 1956; Vasil, 1960). Curtis and Duncan (1947) found that NAA did not increase pollen germination in the three orchids namely, Cyrtopodium punctatum, Cattleya guatmalense, and Phalaenopsis hybrid. However, this could be attributed to the comparatively high concentrations used (0.1 — 10 mg/L) when compared with present studies. Though the effect of gibberellic acid and other gibberellins on germination and elongation of pollen tubes has been extensively studied in other angiosperms, they have not been used before for orchid pollen (Chandler, 1957; Kato, 1955). In both the hybrids of Vanda the percentage germination and tube growth improved in low concentration of GA media. Johri and Vasil (1961) postulated that incorporation of kinetin in nutrient medium together with auxins may hasten the division of generative cell. Kinetin improved the percentage germination as well as tube growth of Vanda Miss Joaquim and the results were much better than in control, salt-boron, or certain auxin media and similar results were recorded in 256 Gardens’ Bulletin— XXVI (1973) V. Poepoe. Thus the present studies would indicate that in the Vanda hybrids the organic nutrients would improve the pollen germination and tube growth to a greater extent than the inorganic or the other growth substances. As for the pollen of other hybrids they are viable as indicated by selective staining. Different physical conditions or growth adjuncts than the ones used presently may be necessary to induce their germination and these are being investigated. Summary A study of pollen morphology, structure and germination is made in the well-known orchid hybrids. They are Arachnis Maggie Oei var. Red Ribbon, Arachnis Maggie Oei var. Yellow Ribbon, Aranda Hilda Galistan, A. Lucy Lay- cock (long spray), A. Luck Laycock (short spray). A. Wendy Scott var. Greenfield, Aranthera Lilliput, Vanda Poepoe and Vanda Josephine. Experiments were con- ducted to germinate them in the laboratory conditions. Of these, only Vanda Poepoe, Vanda Miss Joaquim gave positive results and very limited results were obtained in case of Arachnis Maggie Oei var. Yellow Ribbion. Pollen of other hybrids did not germinate. Different substances that are recognised to promote pollen germination were used at different concentrations and distilled water formed the control medium. Percentage germination, tube length, and morphological variations of such structures are determined and described. In these hyrbids, for pollen germination the organic sources (sucrose, stigmatic extract) appear to be most effective than the inorganic ones (Borax, salt-boron combination, auxins, GA or K). Auxins improved the germination and of all the auxina used IAA was most effective. Similarly GA and K improved the germination in both the hybrids of Vanda. Grateful acknowledgment is made to Mr. John Ede who permitted us to collect the orchids from Mandai garden and Mr. A. G. Alphonso, Acting Director, Singapore Botanic Gardens for the correct identification of the hybrids and some technical information. We are thankful to Mr. R. Tay, Mr. D. Teow and Mrs. M. Goh for their technical assistance. Literature Cited Adams, H. H., 1959: Aspects of variation in the Orchidaceae 73-154 in The orchids — A Scientific survey. (ed.) C. L. Withner. The Ronald Press Co. New York. Brink, R. A., 1924: The Physiology of Pollen I. The requirements for growth. Amer. Jour. Bot., /7, 210-228. , 1924a: The Physiology of Pollen III. Growth in vitro and in vivo. Ibid., 11, 351-364. Brewbaker, J. L. & B. H. Kwack, 1964: The calcium ion and substances influencing pollen growth. Jn Physiology and Fertilisation (ed.) H. F. Linskens, North Holland Co. Amsterdam, 143-151. Chandler, C., 1957: The effect of gibberellic acid on germination and pollen tube growth. Contrib. Boyce Thomp. Inst., 19, 215-223. Cook, E. S. & Walden, D. B., 1965: The male gametophyte of Zea mays L. II. In vitro germination. Can. Jour. Bot., 43, 779-787. Pollen Viability and Germination in some Orchid Hybrids Zar Curtis, J. T. & R. E. Duncan, 1947: Studies in the germination of orchid pollen. Bull. Amer. Orchid Soc., 16, 595-597. Dikshit, N. N., 1956: Effects of hormone on germination in loquat pollen. Curr. Sci., 25, 27-28. Holttum, R. E., 1953, Flora of Malaya — Orchids of Malaya. Govt. Printing Office, Singapore. Johri, B. M. & I. K. Vasil, 1961: Physiology of pollen. Bot. Rev. 27, 325-381. Kamemoto, H., 1956; Cytology of Vanda Nellie Morley and Vanda Emma Van Dercuter. Amer. Orchid Soc. Bull. 25: 234-243. Kato, F., 1955: Responses of plant cells to gibberellin. Bot. Gaz. 117: 16-24. Loo, T. and T. Hwang, 1944: Growth stimulation by manganese sulfate, IAA and colchicine in pollen germination. Amer. Jour. Bot., 3/: 356-367. Masceranhas, J. P. & Machlis, L., 1964: Chemotropic response of pollen of Antirhinum majus. L. to calcium. Plant Physiol. 39: 70-77. Miwa, A., 1937: Test of germinating power of orchid pollen. Orchid Rev. 45: 345-349, O’Kelly, J. C., 1955: External Carbohydrates in growth and respiration of pollen tubes ‘in vitro’. Amer. Jour. Bot. 42: 322-326. , 1957: Boron effects on growth, oxygen uptake and sugar absorption by germinating pollen. Amer. Jour. Bot. 44: 239-244. Roa, A. N. & C. J. Goh, 1970: Progress of orchid research in Singapore, Proc. Seminar on Res. programmes in Singapore. Nanyang Univ. P. 111-113. , & Lee Wai Chin, 1972: Experimental studies in the germination of orchid pollen. La Cellule. 57: 292-310. Rosen, W. G., 1964: Chemotropism and fine structure of pollen tubes. 159-166. in pollen physiology and fertilisation (ed.) H. F. Linskens. North Holland Co. Amsterdam. Smith, P. F., 1942: Studies on the growth of pollen with respect to temperature, auxins, colchicin and vitamin B., Amer. Jour. Bot. 44: 637-650. Vasil, I. K., 1960: Studies on pollen germination of certain Cucurbitaceae. Amer. Jour. Bot., 47: 239-247. , 1964: Effect of Boron on pollen germination and pollen tube growth. Linskens, H. F. (ed.). Pollen physiology and fertilisation. North Holland, Amsterdam. 107-119. Visser, T., 1955: Germination and storage of pollen. Mededeel. Landbouwhoo- geschool (Wageningen), 55: 1-68. Withner, C. L., 1959: The orchids. A scientific survey. Ronald Press Co., New York. rae uerratlog, si-coitolpeteidipaiag Orgun ic mutdents war re isin OVRQRL | extent then the. anecganis ot Md dP ue rica ‘sae ee aetire ! | conditions ‘oF satan adjunety Mine ’ 9 es | — io duelinnm Aiie sernlnating ang these oO sonny Yond i! sa, to ebitiax — = SuMpary 1BE-2EE NS vari of taliog to vmoloigyt?. 219¢ 4) “HiT: “4 ole \ qahdMe sitet ko Mis ie eNOS af LS tas” ts . ' : (wlscy J a. Aranda! ite Li oe railloneddigi chal ae re OGTR ery my | ® in she, Iaborslory, . Hue Bad Pec LEM nc, Geil os ane a lL, Re aaa YO FShiog Yo Wed . oF POLON Bee] oe ee alts ”. contin: = tad Mf 48 : i OSGeT veo ah dencrh og a to ai fe ns 251 on gonmunation the Oneank ® Te, Sipe ne ditcorwe than. toe haceehaas . oh Bor ‘, = ie ri heag Cone ‘ BA Va ’ “ = ~? an oflow Aa nove pat ifr able trishrti sasetbgtont > lation on a4 q a f feutive, & AS + oe os « wots “SeT7y+ oni Tie ~~ tye haa 9 one qu IBY AG i voix ie we) np | 5 baa eet “ Ge “tid “SKA, trot ; soi overphtgndetii si Biro Je wosrgey rer Pe FA Soke b: Sei higatty Aut ayo eate i actiiest q07 29 Si Anish oii ol exihula ilagmeisg 7.4 Bijan sit isW sed: 2. OLESOS 3%. obistie) et .srallog bid 20. iy Id 7 ; snigInde Slt vere beat iqanto 4 die? sean! “by Cc hot aailinsy a ritkhy i, ss : ¥ vey . Co, fa eae Hipsrgmsd ot soeaaon Asim, mllog bs aman, | iy a0 FE a ea ‘tod iia es A fi batho eid .2bc Ri ERA baseline ih dc. it. Shateshahit v “ae a ive. Aollod pas eoRsntniam: athe 3h Ose iasy har, RO iGing > wae ens a 4 North tak ee scabs ‘ _ m eng aii ay) A ai) al a | a e OG tiaees| ely jambs het) mallow tte reente fy . roe, Conta” foe) ar Wena, sexe at in wines romain. — ed ai { ct’ : ‘| - co Lu A * snoring 49 roo f a2 A Review of the Genus Creaghiella Stapf. (MELASTOMATACEAE) by M. P. NAYAR Central National Herbarium, Botanic Garden, Howrah Abstract A new species of Creaghiella is described from Borneo. Anerincleistus philip- pinensis Merr. is transferred to the present genus as Creaghiella philippinensis (Merr.) Nayar. A key to the species so far known is also presented. Introduction Stapf Gin Hook. f. Ic. Pl. 25: t. 2455, 1896) established the genus Creaghiella on the basis of the specimen Creagh s.n. from Maruda Bay, Sandakan in honour of Creagh, Governor of Sabah. The genus Creaghiella is allied to Anerincleistus but differs from it in having subequal or unequal stamens with dorsally gibbose and ventrally 2-lobed connective, whereas in Anerincleistus the connective is inappendiculate ventrally and minutely spurred or tuberculate. The three species C. purpurea Stapf, C. setosa Nayar and C. philippinensis (Merr.) Nayar form a homogeneous group, having similar facies with densely setose or tomentose, usually purplish branches, 4-merous flowers, subequal or unequal stamens with dorsally gibbose and ventrally 2-lobed connective. The purplish branches, leaves and inflorescence give colour to the different species of this genus and this can be introduced as a garden plant. This is an endemic genus with two species (C. purpurea Stapf. and C. setosa Nayar) in Sabah region of Borneo and a single species C. philippinensis (Merr.) Nayar in Palawan Island of Philippines. Palawan Island represents the nearest land-bridge from Borneo to other islands of Philippines. Creaghiella is a genus of densely hirsute shrubs growing in creeks and sand dunes near sea-shore. Key to the species of Creaghiella I. Inflorescence terminal, umbellate. iI. Calyx tube shaggy tomentose, calyx lobes long; inflorescence axis short, stout; flowers and capsules larger C. purpurea II. Calyx tube shortly setose, calyx lobes short; inflorescence axis long, slender; flowers and capsules smaller C. setosa I. Inflorescence axillary, fascicled C. philippinensis 259 260 Gardens’ Bulletin — X XVI (1973) Enumeration of species 1. Creaghiella purpurea Stapf in Hook. f. Ic. Pl. 25: t. 2455, 1896. (Creagh s.n. Holotype K.) Distribution: Endemic to Borneo. Borneo: Sabah, Maruda Bay, Tanjong Batu, Sandakan, 5 May 1895. Creagh s.n. (K) Kg. Malanta, alt. 6 m., 30 May 1933 Olik NBFD 3327 (K, L.); Elopura, Sandakan F.D., alt. 20 m., 18 Dec. 1947, Kadir NBFD A.672 (K); Ibid., alt. 50 m., 20 Jun. 1949, Castro NBFD A478 (K); Ibid., Kadir NBFD A.944 (K); Elopura, Sandakan, Kabili F.R., Austin Cuadra NBFD A.515 (K); Ibid., Keith NBFD 4529 (K, L.); Kabili-seilok F.R., alt. 2 m., 12 Jul. 1937, Enggoh NBFD 7298 (K, L.) Batu Sapi road, Sandakan, 19 Aug. 1960, W. Meijer SAN 21542 (K.). This is a beautiful shrub of 2-4 m. tall with red flowers and pinkish branches and inflorescence. 2. Creaghiella setosa Nayar, sp. nov. Affinis C. purpureae Stapf. sed floribus capsulisque minoribus, lobis calycinis minoribus, calycis tubo minute setoso, pedunculis longioribus tenuibus differt. Frutex Rami teretes, dense hirsuto-tomentosi. Folia ovata, 10-20 cm X 6-13 cm, basi rotunda vel subcordata, apice acuminata, margine denticulata, supra velutino-hispida, subtus hispida, 7-nervia, venulis transversis supra et subtus distinctis, membranacea; petiolus 2-6 cm longus, dense hirsuto-tomentosus. Jnflo- rescentia terminalis, 7-15 cm longa, pseudoumbellata, dense hirsuta. Calycis tubus campanulatus, 5—5.5 mm longus, dense setosus, setis 0.5 mm longis limbus 4 lobatus, lobis triangularibus, 1-2 mm longis. Petala oblonga, 2.5-3 mm X 1-1.5 mm, apice ciliata. Stamina 8, subaequalia, filamentis 6-7 mm longis, antheris lanceolatis 6-8 mm longis, apice rostratis, connectivo non producto, antice 2 lobato, postice gibboso. Ovarium calycis tubo septis 8 adnatum, coronatum. Capsula 4 X 3.5 mm, dense setosa. Semina minuta, 0.3-0.4 mm longa, numerosa. TYPUS: G. Mikil SAN 31422 (Holotype L.) Shrub. Branches terete, densely hirsute tomentose. Leaves ovate, 10-20 cm x 6-16 cm, base rotund or subcordate, apex acuminate, margin denticulate, upper surface velvety hispid, under surface hispid, 7-nerved, transverse venules on the upper and lower surface distinct, membranaceous; petiole 2-6 cm long, densely hirsute tomentose. Inflorescence terminal, 7-15 cm long, pseudoumbellate, densely hirsute. Calyx tube campanulate, 5—-5.5 mm long, densely setose, bristles 0.5 mm long, 4-lobed, lobe triangular, 1-2 mm long. Petals oblong, 2.5-3 mm X 1-1.5 mm, apex ciliate. Stamens 8, subequal, filament 6-7 mm long, anther lanceolate 6-8 mm long, apex rostrate, connective not produced, connective ven- trally 2-lobed and dorsally gibbose. Ovary adnate to the calyx tube by 8 septa, disc present. Capsule 4 X 3.5 mm, densely setose. Seeds minute, 0.3—-0.4 mm long, numerous. Distribution: Borneo: Sabah. Klias, Beaufort Dist., Sept. 1962, G. Mikil SAN 31422 (L); Beaufort, alt. 66 m., 28 Sept. 1932, Bakar NBFD 3399 (K, L.); Padas gorge, south side, Tenom Dist., Sept. 1959, W. Meijer SAN 19827 (K, L.); Pangi, Tenom Dist., alt. 500 m., 5 Sept. 1954, G. H. S. Wood & Wyatt Smith SAN A.4311 (L.). Review of Creaghiella (Melastomataceae) 261 C. setosa is closely allied to C. purpurea, but it is immediately recognisable by its smaller flowers, smaller capsule and short setose hairs in the calyx tube and long slender inflorescence axis and smaller calyx lobes; whereas in C. purpurea the flowers and capsule are larger, calyx tube is shaggy tomentose and inflorescence axis is short and robust. 3. Creaghiella philippinensis (Merr.) Nayar comb. nov. Anerincleistus philippinensis Merr. in Philipp. Journ. Sc. Bot. 12: 337, 1917. Distributon: Philippines: Palawan, Malampuya, Sept. 1910, Merrill 7241 (Syntypes K, BM.); Ibid., May 1913, Merrill 9412 (Lectotype K); Mt. Capoas, Apr. 1913, Merrill 9552 (Syntype K.). In the original description it is mentioned that the stamens are dorsally tuber- culate and ventrally inappendiculate. On careful scrutiny it is seen that the anthers are dorsally gibbose and ventrally 2-lobate; whereas in the genus Anerinceistus the subulate anthers end dorsally in a tubercle and ventrally they are inappendi- culate. Besides the nature of floral parts, the nature of its leaves and its habit indicate that this taxon is allied to Creaghiella purpurea and C. setosa of Sabah, Borneo. In C. purpurea and C. setosa the inflorescence is terminal, whereas in this taxon the inflorescence is axillary. But in all the three species the flower buds in juvenile stage are subsessile and the pedicels prominently elongate during anthesis. C. philippinensis is allied to C. purpurea, but is easily distinguished by its axillary inflorescence, larger petals (1.5 cm X 1.1 cm) and short ventral appendages; whereas in C. purpurea the inflorescence is terminal, petals are much smaller (3.5-4 mm X 2 mm) and the ventral appendages are prominent. Acknowledgements I wish to express my gratitude to Sir George Taylor, Director, Royal Botanic Gardens, Kew, for all facilities during my stay at Kew from 1961-67. I am indebted to the authorities of the following Herbaria for their hospitality during my visits and for the loan of herbarium specimens: Rijksherbarium, Leiden and The British Museum (Nat. Hist.) London. My thanks are also due to Dr. K. Subramanyam, Director, Botanical Survey of India, for his encouragement. Py ees) vd oldseingooer yisisibeny, ai mn iE. a“ boe sdut xyiso odd aie ibe ala sie p iinet - Engemtc to Bota aS. fl Sab: ih. 3 a if | May $895, Ceiam somes gf Sf0r AtE Ay tod 52 iin Gita Aa vinye) TST: CURT” sqpe “eT? 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Sbadgeatir ore odds . yevooy nigral 8 fi yllaztob a jided ai bas eoveol ei} to gaiem od) 2haq Imioh lo srw Bn 3d aisha Peas Kira phe si ay oi bills YO" ca woue innonioed, loom S tiimeribus, calye ot Rp. ud ‘hi’ veo Hw OF giitherid “aP | SAMS ot) mote: . baa s ot ebpdtieswom Crmmarsigd ii SH) Ue He rene? oy 0 uzigodtne gah! sisgkols ylinsitmo ry sevibay els hasty viele Bindu bedelugaiiibvelizes aitud ayostruats . tpagabiaqqenrdtaines soda bien (ind doh DG gang tunped aellame. dourmtrate, aster Jselers) eheetneondndlian, lions | thir campanviatus, 5-SNIGHROIGQ tls gas pikewge teunoy ohne Dre = obit. triangularibes, 22 , onga, 2.5-3 mm bhi: ah Citing 8 7 niin longis, ‘antheris 1 "J is, unio’ meters m orexhocto,; antice 2 \ fe sinstOd [VOT Aototih AMbT Sto HEU Shoe Vir s-spiefhe beri bet eee P wea pac HT WoA 1a Fe VRP RRL BSB) te 10} .v aiziv yor gainoh vivtatiqand tiods tot etahsl gniwollot st to esitine dai. 2A T DAR ple lida oledie5? orededtiogge ausixedted to sane SUE AY, (8G tity Alt Ohm? su Cale one leading hiimaebatdrdaei Ava ~— THING eid. 703 yaad b Jo, gourd tfacdé ver Tapa > ‘ 1 aD hispid, 7-nesved, iran uoper and lower eithabe cd Y. mombrana¢eo rd peti ste usute tomentase Soctlyur ww ue rs rminal, j- is mend or —s dy hirsute, Colyx se Ghanipariniaie, 4-3, ld long, Se bn long, 4-lobed, ice: banesias: +2 umd rag. Pea ob 1-15 mm, apex cities, Sao &, ty wk am lanceolate 6-3 om long, apek somGt, oouie tralty 2-lobed and dores ny pibbow Cvery dive present. Capsule 4% BS rata, Genatsy welsh eh ie jer Tix Cistribuclon: ail Sabah, Kidae, Rew ? (Ly); Beaufort, slt, 66 i, BF gs gorge. ‘south side, Tenom (ist, Sopt, 59, J WM ‘Tenom er $00 my 5 See ad G, AAs Uieteln hae eieaag Sarr, ae ean ' New Plant Disease Records for Sarawak for 1970 and 1971 by KUEH TIONG-KHENG Department of Agriculture, Sarawak, E. Malaysia Lists of plant disease records for Sarawak have been given by Johnston (1960) and Turner (1963, 1964, 1966, 1967, 1969 and 1971). The list below consists of previously unrecorded fungal diseases together with diseases caused by algae and plant parasitic nematodes, observed or collected by the writer from the time of his taking over the office of Plant Pathology Section of Semongok Agricultural Research Centre in mid August, 1970 until the end of 1971. Thirteen of these records appear in the Annual Report of the Research Branch, Department of Agriculture, Sarawak, for the year 1970. The causal organisms are listed under their respective host plants which are arranged in alphabetical order of their botanical names. The frequency of occur- rence is given together with the Commonwealth Mycological Institute Herbarium serial number, where identification has been performed by the Institute. Two species of fungi and one nematode species were identified at the Royal Botanic Gardens, Kew, and the Commonwealth Institute of Helminthology, Herts, respec- tively. In the list that follows, One, Occ., Comm. stand for One record, Occasionally and Common respectively. Alternanthera bettzickiana var. spathulata Wilt Corticium rolfsii Curzi One — Ananas comosus Merr, (Pineapple, Nanas) Leaf rot Curvularia eragrostidis (P. Henn.) J. A. Meyer One 142889 Annona squamosa L. (Sweet Sop) Red rust Cephaleuros virescens Kunze One — Arachis hypogaea L. (Groundnut, Kachang Tanah) Wilt Fusarium solani (Mart.) Sacc. One 159548 Begonia glabra Ruiz. et. Pav. Wilt Corticium rolfsii Curzi One — Bixa orellana L. (Annatto) Leaf blight Pesialotiopsis theae (Saw.) Steyaert One 163728a Leaf blight Colletotrichum capsici (Syd.) Butler & Bisby One 163728b Brassica juncea Coss. (Kai Choy) Leaf rot Choanephora cucurbitarum (Berk. & Rav.) Thaxt. Occ. — 263 264 Gardens’ Bulletin — X XVI (1973) Canangium odoratum Baill. (Kenanga) Velvet blight Septobasidium sp. One — Capsicum annuum L. (Chilli, Chabai) Leaf spot Leptosphaerulina trifolii (Rostrup) Petr. One 153073 Capsicum frutescens L. var. baccatum (Ornamental pepper) Leaf spot Cercospora capsici Heald & Wolf Occ. 163286 Carica papaya L. (Papaya) Leaf spot Sphaerulina sp. One 159602b White root disease Fomes lignosus (Klotzsch) Bres. One — Chrysanthemum sp. Flower blight Cladosporium oxysporium Berk. & Curt. One 160403a Flower blight Fusarium equiseti (Corda) Sacc. One 160403b Leaf spot Cercospora chrysanthemicola Yen One 163291a Leaf spot Septoria chrysanthemella Sacc. One 163291b Citrus microcarpa Bunge (Musk-lime, Limau kesturi) Pink disease Corticium salmonicolor Berk. & Br. One — Cocos nucifera L. (Coconut, Kelapa) From roots of wilting Cunninghamella echinulata (Thaxt.) palm Thaxt, ex Blakeslee One 159555 From roots of wilting Ceratocystis paradoxa (Dade) Moreau One 159557 palm From roots of wilting Fusarium monileforme Sheld. One 159562 palm From fabric bark of Fusarium solani (Mart.) Sacc. One 159556 & 159559 wilting palm Codiaeum variegatum Blume (Garden Croton) Velvet blight Septobasidium sp. One 163287 Coffea robusta Linden (Robusta coffee, Kopi) Leaf blight Glomerella cingulata (Stonem.) Spauld. & Schrenk One 163289 Elaeis guineensis Jacq. (Oil palm) Stem rot ? Thielaviopsis sp. One 148041 Dry basal rot Ceratocystis paradoxa (Dade) Mareau Occ. 159563 Ficus glossularioides Burm. f. Leaf spot Glomerella cingulata (Stonem.) Spauld. & Schrenk One 163726 Ficus uncinata Becc. Leaf spot Cercospora sp. One 163727 Fimbristylis globulosa Kunth (Globular Fimbristylis, Rumput sandang) Inflorescence smut Cintractia exicola (Berk.) Cornu Occ. 163730a & 163729 Inflorescence stalk Stauronema sp. One 163731 blight Inflorescence smut Fusarium heterosporum Nees ex Fr. One 163730b Inflorescence smut Fusarium oxysporum Schlecht. One 163730c New Plant Disease Records, Sarawak, 1970 and 1971 Fimbristylis miliacea Vahl (Lesser Fimbristylis, Rumput tahi kerbau) Wilt Corticium rolfsii Curzi Flemingia congesta Roxb. (Bush cover crop) Thread blight Marasmiellus scandens (Mass.) Dennis & Reid Glycine max (L.) Merr. (Soybean) Leaf spot Phyllosticta glycinea Tehon & Daniels Leaf spot Ascochyta sp. Leaf blight Choanephora circinans (Nag. & Kawak.) Hess. & Benj. Leaf blight Curvularia eragrostidis (P. Henn.) J. A. Meyer Associated with leaf Fusarium equiseti (Corda) Sacc. blight Guizotia abyssinica (L.f.) Cass. (Niger seed) Collar rot Corticium rolfsii Curzi Impatiens balsamina L. (Balsam) Leaf spot Cercospora fukushiana (Mats.) Yam. Wilt Corticium rolfsii Curzi Imperata cylindrica Beauv. (Lalang) Leaf spot Colletotrichum graminicola (Ces.) Wilson False smut Cerebella andropogonis Ces. Inflorescence mould Curvularia pallescens Boedijn One One One One Occ. One One Occ. One Occ. One One One Leersia hexandra Swartz. (Tiger’s tongue grass, Rumput lidah rimau) Leaf spot Pyricularia oryzae Cav. Mangifera indica L. (Mango) Leaf anthracnose Glomerella cingulata (Stonem.) Spauld. & Schrenk Manihot utilissima Pohl. (Cassava, Tapioca, ubi Kayu) Leaf blight Pestalotiopsis sp. Leaf spot Phyllosticta manihot Speg. Tuber rot Sphaerostilbe repens Berk. & Br. Wilt Corticium rolfsii Curzi Michelia alba DC. (White chempaka, Chempaka puteh) Thread blight Marasmiellus scandens (Mass.) Dennis & Reid. Musa sapientum L. (Banana, Pisang) Leaf speckle Periconiella muse M. B. Ellis Leaf spot Pestalotia leprogena Speg. Heart leaf disease Drechslera mus@-sapientum (Hansf.) B. Ellis Myristica fragrans Houtt. (Nutmeg, Buah pala) From roots Gliocladium sp. 265 160402e 160402f 160402a 160402d 160402c 163734 163282 154092a 154092b Comm. — One 163733 Comm. 163292a Occ. One One One One One One One 163292b 153071 153070 163281 159551 266 Gardens’ Bulletin— X XVI (1973) Nephelium lappaceum L. (Rambutan) Fruit blight Fusarium sp. One 163279 Oryza sativa L. (Rice, Padi) Brown grain On grain Wilt Leaf spot Leaf spot Brown grain Pithomyces sacchari (Speg.) M. B. Ellis One 159593b Pestalotiopsis sp. One 159593c Corticium rolfsii Curzi Occ. — Curvularia eragrostidis (P. Henn.) J. A. Meyer One 142890 Pestalotiopsis ? disseminata (Thum.) Steyaert One 142891 Pestalotiopsis palmarum (Cooke) Steyaert One 159549 Pachyrrhizus erosus Urban. (Yam bean) Wilt Colletotrichum capsici (Syd.) Butler & Bisby One 163280 Panicum repens L. (Creeping panic grass, Rumput halia) Wilt Blast Corticium rolfsii Curzi Occ. — Pyricularia oryzae Cav. Comm. — Paspalum scrobiculatum L. (Carpet grass) Seed blight Fusarium heterosporum Nees ex Fr. Comm. 154093b Pennisetum purpureum Schum. & Thonn. (Napier grass) Leaf spot Leaf spot Leaf spot Leaf spot On dead stems & leaves Curvularia eragrostidis (P. Henn.) J. A. Meyer Occ. 163288a Cochliobolus lunatus Nelson & Haasis One 163288d Pithomyces chartarum (Berk. & Curt.) M. B. Ellis One 163288c Septoria sp. One 163288b Volvariella esculenta (Massee) Sing. One — Peperomia pellucida H.B. & K. Wilt Corticium rolfsii-Curzi One — Piper betle L. (Betel, Sireh) Pink disease Corticium salmonicolor Berk. & Br. One — Piper nigrum L. (Black pepper, Lada Hitam) On dead branches On dead spikes, leaves & branches Black spike Leaf spot Pleurotus (Hohenbuehelia) testudo Berk. One Kew Marasmius campanella Holterm. . | One Kew Cochliobolus geniculatus Nelson One 159552 Glomerella cingulata (Stonem.) Spauld. & Schrenk One 159553 Pogostemon cablin Benth, (Patchouli) White root Root knot Wilt Fomes lignosus (Klotzsch) Bres. One — Meloidogyne javanica (Treub., 1885) Chitwood, 1949 Occ. CIH Corticium rolfsii Curzi One — New Plant Disease Records, Sarawak, 1970 and 1971 Ricinus communis L. (Castor oil plant) Leaf spot Fruit blight Leaf spot Fruit blight . Fruit blight Fruit blight Alternaria ricini (Yoshii) Hansford Colletotrichum capsici (Syd.) Butler & Bisby Ascochyta sp. Fusarium oxysporum Schlecht. Botryodiplodia theobromae Pat. Glomerella cingulata (Stonem.) Spauld. & Schrenk Occ. 159600b One 154094a One 159600a One 154094b One 154094c One 154094d On Clypeolella ricini Cicinobella sp. Rosa sp. (Rose) Leaf spot Mycosphaerella sp. Sanchezia nobilis Hook. f. (Kadok gajah) Leaf rot Corticium solani (Prill. & Delacr.) Bourd, & Galz. Sorghum sudanense Stapf (Sudan grass) Leaf spot Fusarium sp. Sorghum vulgare Pers. (Sorghum) Leaf spot Panicle disease Panicle disease Cercospora sorghi Ell. & Ev. Fusarium semitectum Berk. & Rav. Alternaria Iongissima Deighton & One One One One One 159601b 159597b 163278b 153068 153069b Collar rot MacGarvie Corticium rolfsii Curzi Uncaria gambir Roxb. (Gambir) Leaf rot Vanda vars. (Orchid) Leaf spot Corticium solani (Prill. & Delacr.) Bourd. & Galz. Cochliobolus geniculatus Nelson Vanilla planifolia Andr. (Vanilla) Rust Uredo scabies Cke. Zea mays L. (Maize, Jagong) Leaf spot Leaf spot Curvularia senegalensis (Speg.) Subram. Diplodia zeicola Saccas Zingiber officinale Rosc. (Ginger, Haliya) Leaf scorch Hendersonia zingiberi Sawada Leaf spot Coniothyrium fuckelii Sacc. Leaf spot Ophiobolus sp. Leaf scorch Tetraploa aristata Berk. & Br. Leaf scorch Leptosphaeria sp. Leaf blight Pyricularia zingiberi Nishikado Red rust Cephaleuros virescens Kunze Leaf blight Leptosphaeria typhae Karst. Leaf spot Phyllosticta zingiberi Ramakrishnan Leaf blight Cochliobolus geniculatus Nelson One 153069c One — One — One 153074 One 159596 One 163284a One 163284b One 163285a One 163285b One 163285c One 163283b One 163283c One 159594b One 159594c One 159594a Comm. 153072a 163283d Comm. 153072b 163283a 267 268 Gardens’ Bulletin — X XVI (1973) Acknowledgements I am grateful to the Director of Agriculture, Sarawak, for granting permission to publish this list and the Directors and staff of the Commonwealth Mycological Institute, the Royal Botanic Gardens, Kew, and the Commonwealth Institute of Helminthology, Herts, without whose assistance in identifying many of the species, the list could not have been published. References Annual Report (1970). Research Branch, Department of Agriculture, Sarawak. Johnston, A. (1960). A preliminary plant disease survey in Sarawax. F.A.O. Rome (mimeographed). Turner, G. J. (1963). New records of plant diseases in Sarawak for the years 1960 & 1961. Gdns’. Bull. Singapore, 20 285-288. Turner, G. J. (1964). New records of plant diseases in Sarawak for the year 1962. Gdns’. Bull. Singapore, 20 369-376. Turner, G. J. (1966). New records of plant diseases in Sarawak for the years 1963 & 1964. Gdns’. Bull. Singapore, 21 393-402. Turner, G. J. (1967). New records of plant diseases in Sarawak for the year 1965. Gdns’. Bull. Singapore, 22 123-128. Turner, G. J. (1969). New records of plant diseases in Sarawak for the years 1966 & 1967. Gdns’. Bull. Singapore, 24 181-184. Turner, G. J. (1971). Fungi and plant diseases in Sarawak. Phytopath. Papers No. 13, C’wealth Mycol. Inst., Kew, Surrey, England. Jan. 1971. Notes on the Systematy of Malayan Phanerogams XVIII — XXII * by K. M. KOCHUMMEN & T. C. WHITMORE Forest Research Institute, Kepong Abstract Calophyllum austrocoriaceum and C. rupicolum var. elatum are new taxa. C. intramar- ginale is reduced to C. inophylloide var. singapurense. A division of Malayan Garcinia into groups is described. G. burkillii, G. cantleyana and its var. grandifolia, G. cataractalis, G. hendersoniana and G. murtonii are new species. G. dumosa (and its Malayan allies), G. nigrolineata, G. penangiana, G. bancana, G. opaca and G. pyrifera are annotated with several new synonyms proposed; G. opace var. dumosa is proposed as a new variety. In Mesua notes are provided on the alliances of M. assamica, M. elegans, M. kunstleri and M. lepidota; and M, kochummeniana, M. nivenii, M. nuda and M. purseglovei are new species. Barringtonia payensiana is a new species. Crudia viridiflora is a new species, with simple leaves. In Olacaceae, Strombosia maingayi is shown to be the legitimate name for the tree long known as S. rotundifolia; the differences from S. multiflora are discussed. In Proteaceae, Heliciopsis cockburnii, H. montana and H. whitmorei are described as new species. This fourth instalment of our series of notes completes the taxonomic ground- work for volume 2 of the Tree Flora of Malaya, which takes the project over the half way mark. The longest note is on the Guttiferae, mainly Garcinia, the second biggest family (after Euphorbiaceae) treated in volume 2. Yet another new legu- minous species has been discovered, this time a simple-leaved Crudia. Once again we express our thanks to Dr. B. C. Stone for help with the latin descriptions. XVII. GUTTIFERAE T. C. WHITMORE Calophyllum Calophyllum austrocoriaceum Whitmore sp. nov. C. sp. 10 in Henderson & Wyatt-Smith Gdns’ Bull. Sing. 15 (1956) 313, plate VILL, Fig. 3 E. Arbor parva, ramulis crassis, laevibus, leviter complanatis, furfuraceis; gemmis terminalibus crassis, lanceolatis, acutis, subtiliter tomentosis, usque 15 mm longis, foliis valde coriaceis, oblongis vel anguste ovate-ellipticis (7 x 3) - ll x 3 - 16 x 5 cm, apice rotundato vel obtuso, base cuneato, leviter decurrenti, costa lato * continued from Gdns’ Bull. Sing. 26: 49-61. 269 270 Gardens’ Bulletin — X XVI (1973) pagina infera prominenti pagina supra prope basim interdum canaliculato, nervis utrinque conspicuis subdistantibus, petiolo 2-3 cm crasso. Inflorescentiae axillares pauciflorae. Flores ignoti. Fructus subglobosa, usque 23 mm longa apiculata, pericarpio crasso duro siccitate.brunneo sublaevi, pedicello crasso 1 cm, pedunculo crasso 15 mm. JOHORE: G. Panti FRI 7846 (holotype KEP); FRI 7729, 13819, SFN 29969; G. Belumut KEP 98038, FRI 7512, 7789, 8722, 8809. Not seen: SFN 10723, cited by Henderson and Wyatt-Smith. This is close to C. coriaceum of the mountains of the Main and east coast ranges and replaces it on the southermost mountains of the Peninsula. The name is chosen to reflect this affinity. It is also closely related to C. tahanense of G. Tahan. From both of its relatives C. austrocoriaceum conspicuously differs in the nerves being prominent and coarse on both surfaces, the short, few-flowered, axillary racemes and the apiculate unbeaked fruit. Calophyllum imophylloide var. singapurense Henderson et Wyatt-Smith Gdns” Bull. Sing. 15 (1956) 316, plate IX, Fig. 4 D-H. C. intramarginale Henderson et Wyatt-Smith loc. cit. 342, plate XXII, Fig. 13A, B- syn. nov. The latter species is said to differ in its smaller leaves with intramarginal vein, shorter inflorescences with all the flowers apical on the peduncle, and its montane habitat. It appears that Henderson and Wyatt-Smith drafted the monograph when only the type of C. intramarginale was known, a collection from a small tree from 4000 ft on G. Padang; and that KEP 67753, from 1020 ft in Rasau Kerteh F.R., was added later, but neither the key nor notes were amended to allow for its lowland station. I have since made a second collection from G. Padang (FRI 12644) in which the intramarginal nerve is very difficult to see. Three other collections have recently been made which have an intramarginal nerve on one or a few leaves (FRI 41/4, 8839, 10840) and these are all big trees which in leaf shape, bark, exudate colour and the possession of buttresses resemble C. inophylloide var. singapurense. They come from W. Kelantan, N. E. Johore and Trengganu respectively. The flowers of C. inophylloide var. singapurense arise in a cluster from the top of the peduncle and sometimes, on long racemes, there are also 1-2 lateral pairs. In C. intramarginale these lateral pairs are always lacking. I do not believe C. intramarginale can be maintained and therefore reduce it. C. inophylloide var. singapurense differs mainly from the typical variety in its smaller leaves. Calophyllum rupicolum var. elatum Whitmore var. nov. C. rupicolum var. in HENDERSON & WYATT-SMITH Gdns’ Bull. Sing. 15 (1956) 347, plate XXVII, Fig. 13 M, N. A varietas typica elatiore usque 18 m, in foliis maioribus, latioribus, 8 x 2.7 — 11 x 3.5 cm, differt. Systematy of Malayan Phanerogams XVIH-X XII 271 KELANTAN: FRI 2538 (holotype of var. at KEP). TRENGGANU: Ulu Brang FRI 15256. G. Tebu F. R. FRI 2513, 2516. Bt. Rambai F. R. FRI 11401. KEDAH: Ulu Muda F. R. FRI 6757. PAHANG: Rompin KEP 29648. PERAK: Dindings, Telok Kopia F. R. FRI 3105. JOHORE: Kangka Sedili Kechil, Corner s.n. 17. VI. 34; Mawai, SFN 34706, 34747. Henderson & Wyatt-Smith left this as an undescribed variety. More material has now been collected and I feel confident in describing it, but as a variety of the river-bed bush C. rupicolum and not as a full species. Apart from the relatively broader leaves, and the habit and habitat it comes very close to C. rupicolum, and there is some overlap in leaf. All the collections are of small trees away from rivers except FRI 12526, described as a riverside bush, yet with leaves 11 x 3.5 cm, the largest seen. I have studied C. rupicolum on the Sg. Tahan near K. Tenor. There it grows as a small, sinuous tree to 6 m tall on earth river banks at shingle shallows. Leaf size was greatest at the top of the crown and tiny at the crown base, below the level of the highest floods. The collection, FRI 15985, clearly shows the narrow leaves of the typical variety, they are up to 8 cm long. Garcinia I have found it useful to divide Malayan Garcinia into six groups, named A-F, based on the structure of the male flower. These groups are a refinement of. the numbered sections proposed by King in the key to his account in the Materials (J. As. Soc. Beng. 59, 1890, 148-72) and which were more or less copied by Ridley in his Flora (1, 1922, 167-80). To some extent they are natural, though groups B and C very probably are not (species of subgroups Bi and Ci are very similar and easily confusable). A guide to the groups in the form of a key appears in my account for the Tree Flora and this should also be consulted. It must be emphasised that the groups are informal ones, and are based on a study only of Peninsula material; they are published here because I believe they might per- haps provide a useful starting point for other students of this big and intricate genus, but they certainly should not be regarded as definitive. statements on the taxonomy of the whole of Garcinia. The subgroups, where such are defined, are species which are closely similar in one or several respects, i.e. ‘circles of affinity’, and not all species fall into clear subgroups. Further notes on individual species, including the subgroups, can be found in the Tree Flora. In these studies I have not found helpful the groupings proposed by Anderson (in Hk. f. Fl. Brit. Ind. 1, 1884, 259-70) or those proposed by Pierre (For. FI. Cochinch. 1, 1883, i—xlii). The groups are defined on the structure of the male flower. Other characters common to the component species are also given in the descriptions below. There is a final artificial group, G, of 3 incompletely known species. Group A. Sepals and petals 4. Stamens numerous, on 4 fleshy processes, anther cells 2 (4 in G. cuspidata, fide King), pistillode present. Subgroup Ai. Leaves reminiscent of Eugenia, with fine, rather close, parallel, secondary nerves arising nearly at right angles to the midrib. Flowers small, in small inflorescences or clusters, axillary or behind the leaves. 272 Gardens’ Bulletin — X XVI (1973) G. cuspidata King, G. eugeniaefolia Wall., G. merguensis Wight, G. rostrata Wall., G. sp. Al. A closely knit alliance for which I have only been able to give a provisional account; species delimitation needs a critical reappraisal on a monographic basis. See also the notes under G. rostrata below. The following species, dissimilar from each other and from members of the Subgroup Ai, are also in group A: G. diversifolia King, G. holttumii Ridley, G. monantha Ridley, G. murtonii Whit- more, G. sp. A2, G. sp. A3. Group B. Sepals and petals 4. Stamens numerous, joined in a sometimes 4-lobed ring, surrounding a central pistillode; anther cells 2. Flowers terminal, males several, females solitary. Subgroup Bi. Most easily defined as a close knit alliance of spp. extremely similar to the well known mangosteen; virtually indistinguishable when sterile and differing most noticeably in stigma. G. hombroniana Pierre, G. malaccensis Hk. f., G. mangostana L. Species of this alliance occur throughout the East Indies, the best taxonomic treatment of the variation remains to be ascertained from a monographic base. Subgroup Bii. G. costata Hemsley ex King, G. maingayi Hk. f. Two species similar in their coarsely ribbed leaves. Subgroup Biii. Male flowers in a short, terminal raceme, females solitary. Leaves drying thin, green-black. G. atroviridis Griff ex T. Anders. Very distinctive. Group C. Flower structure just as in Group B but with no pistillode, so the stamens as a central mass. Flower position various. Subgroup Ci. Flower position as in Group B. Leaves drying with a distinct reddish or orange tint. G. murdochii Ridley, G. opaca King, G. penangiana Pierre, G. sp. Cl (?). This Subgroup is very close to Bi above. It is discussed further below, under G. penangiana. Subgroup Cii. Flowers mostly behind the leaves, in clusters. Leaves leathery, drying with a distinct reddish tint, or drying thin and green to green-black in G. forbesii, G. griffithii. G. bancana (Miq.) Miq., G. clusiaefolia Ridley, G. forbesii King, G. griffithii King, G. sp. C2. Subgroup Ciii. Leaves with rather open, reticulate nervation, drying brown-green. Flowers minute, in axillary racemes. G. minutiflora Ridley. Not placed: G. montana Ridley. Group D. Sepals and petals 4. Stamens numerous, as a central mass; no pistillode; anther cells 4. Flowers small, in clusters in axils and behind leaves. Leaves drying thin, commonly pinkish grey-brown and with a fine pattern of black dots and dashes. Systematy of Malayan Phanerogams XVIII-X XII 273 G. cowa Roxb., G. nigrolineata Planch. ex T. Anders., G. parvifolia (Miq.) Migq. Corner (Gdns’ Bull. Str. Settl. 10, 1938, 36-8) sorted out the Malayan members. of this small group. Group E. Sepals and petals 4. Stamens not very numerous, connective peltate with the anther cells marginal, often confluent, their dehiscence circumscissile; pistillode absent. Fruits thinly woody. Leaves often drying greenish. Subgroup Ei. Flowers and fruits clustered on small tubercles, axillary and behind the leaves. Leaves usually thin. Often small shrubs. G. dumosa King, G. scortechinii King, G. urophylla Scortechini ex King, Golspy £3 This Subgroup is discussed below, under G. dumosa. Subgroup Eii. Fruits solitary, mainly in leaf axils, some behind the leaves. Leaves: usually leathery. Mainly mountains. G. burkillii Whitmore, G. cantleyana Whitmore, G. hendersoniana Whitmore, G. uniflora King, G: Sp. El (?), G. Sp. E2. This Subgroup shows partly allopatric, partly sympatric distribution patterns in the mountains which are discussed in the Tree Flora. Group F. Sepals and petals 5. Stamens in 5 bundles around a small central pistil- lode. Male and female (and hermaphrodite ?) flowers sometimes mixed. Fruits usually drying shiny brown-black, intricately, sinuously wrinkled. Leaves commonly drying with a greenish or yellowish tinge. G. dulcis (Roxb.) Kurz, G. nervosa (Miq.) Miq., G. prainiana King, G. pyrifera Ridley, G. xanthochymus Hk. f. Long recognised, and at one time gave generic status as Xanthochymus Roxb. Group G. Incompletely known species. G. cataractalis Whitmore, G. sp. G1, G. sp. G2. Garcinia burkillii Whitmore sp. nov. Arbor parva, foliis subcoriaceis, ovatis 7.5 x 4.5 — 15 x 7 cm, acutis, base late cuneato vel rotundato, minime decurrenti, siccitate fuscis, marginibus leviter recur- vatis, nervis lateralibus tenuibus. Flores ignoti. Fructus axillaria solitaria ovoidea, 2 cm longa, usque ad 12 mm diam., siccitate castanea, laevia, tenuiter lignosa, cum sepalis persistentibus ovatis 6 mm latis longisque, stigmatis integro tholiformi 3 mm diam. subsessili subtiliter papillato, axillari, solitario. TRENGGANU: Sg. Loh, K. Datok, FRI 12003. PAHANG: Tahan River, Ridley 2239; G. Tapis FRI 10911 (holotype KEP). In leaf this is close to G. uniflora King of the Main Range, to which I believe it is related (though the flowers remain unknown). The fruiting stigma is, however, quite different. It is also quite clearly a very close relative of G. hendersoniana described below, and differs mainly in the fruit and its stigma. Garcinia cantleyana Whitmore sp. nov. Arbor parva, foliis parvis, crasso coriaceis, ovatis, 5.5 x 3 - 10 x 5 cm, apice late rotundato interdum apiculato, base late cuneato, marginibus valde revolutis; nervis lateralibus tenuissimis, petiolo crasso 14 mm. Flores (femineis ignotis) in 274 Gardens’ Bulletin — XXVI (1973) axillas foliorum delapsorum poxiti, sessiles, sepalis 4 ovatis 4 x 4 mm, petalis 4 quam sepalis aliquantam minoribus, masculi cum staminibus numerosis androecium globosum centralium formantibus, filamentis crassis 1 mm, antheris peltatis 1 mm latis rima lata dehiscentibus, filamentis 1 mm longis spissibus, pistillodio nullo. Fructus subglobosa 3 cm diam., siccitate brunnea, sessilia, calyce persistenti lignoso insidens, stigmatis 3 mm diam. integro tholiformi grosse papilloso sessili. PAHANG: Fraser’s Hill, FRI 5819 (holotype KEP). PERAK: G. Korbu, SFN 16301, KEP 45976; Cameron Highlands, SFN 20603. G. cantleyana var. grandifolia Whitmore var. nov. A varietate typica in foliis maioribus, 17 x 10 cm, fructibus interdum ovoideis differt. PAHANG/SELANGOR: crest of Main Range, road to G. Ulu Kali, FRI 12576 (holotype KEP), FRI 16185. Garcinia cataractalis Whitmore sp. nov. Frutex humilis, foliis lineari — lanceolatis 6 x 0.5 cm. Flores ignoti. Fructus axillaria, carmina, subglobosa, 2—3 lobata, vivo 2.5 cm diam. (siccitate valde vietis, grosse rugosa, 1.5 cm diam.), valde 8 mm rostrata, stigmate nigro 4 — lobato undulato minute papilloso leviter procurrenti, seminibus valde complanatis in pulpa aquosi translucenti inclusis. TRENGGANU: Sg. Kerbat at Jeram Petang: FRI 20350 (holotype KEP); Sg. Nipah at Jeram Gajah: SFN 2591/3. Well known to the local boat men on the Sg. Kerbat as gelugor batu; a highly distinctive rheophyte. | / Garcinia dumosa King G. tenuifolia Ridley Kew Bull. (1928) 72 syn. nov. Garcinia scortechinii King G. gaudichaudii var. minor Ridley Flora 1 (1922) 177 syn. nov. Garcinia urophylla King All King’s three species were described in the Materials (J. As. Soc. Beng. 59, 1890, 166-8). They are only weakly differentiated by the descriptions. The only syntype at Singapore is a sheet of Curtis 1249, G. scortechinii. I am able provision- ally to maintain these species on the weak differential characters given in the Tree Flora but they are undoubtedly close, with apparently little difference in their reproductive parts. A monographer will have to reconsider this group together with its Indo-Chinese relatives. I do not see that Ridley was justified in reducing G. scortechinii to the Indo-Chinese G. gaudichaudii Planch. et Triana, as he did not make the necessary critical comparative studies of this whole alliance, I there- fore do not follow him and, further, I reduce his var. minor of G. gaudichaudii to synonymy, having seen the Johore syntype. G. tenuifolia from the islands off Pahang is simply superfluous, I see no difficulty in reducing this species to G. dumosa. Systematy of Malayan Phanerogams XVIII-X X11 275 Garcinia hendersoniana Whitmore sp. nov. Arbuscula vel arbor parva, foliis percoriaceis, ovatis, 8 x 5.5 - 14 x 8.5 cm, apice acuto, base late cuneato vel rotundato, marginibus revolutis, nervis lateralibus tenuibus attamen manifestis, petiolo 1-2 cm longo. Flores (masculis ignotis) sessiles, solitarii, in axillas foliorum delapsorum poxiti, sepalis 4 ovatis, exterioribus (2) 9 x 8 mm, interioribus (2) 5 x 8 mm, petalis 4 ovatis unusquisque 5 x 5 mm, staminodiis peltatis circulatim dispositis, ovario cylindrico 4 mm longo x 3 mm diam., stigmate sessili tenui integro circulari-discoideo paullo convexo grosse papilloso paullo projectenti 4 mm diam. Fructus (? immaturi) ampulliformia vel rotundata, 3 cm longa, siccitate nigro-cinerea laevia, tenuiter lignosa, sepalis persistentibus. TRENGGANU: G. Padang SFN 31065, 31824; FRI 12718; G. Mandi Angin FRI 10813 (holotype KEP), FRI 10826, 12103. PAHANG: G. Tapis FRI 10994. PAHANG/SELANGOR: G. Ulu Kali FRI 16206. Garcinia murtonii Whitmore sp. nov. Frutex vel arbuscula, vel arbor parva, foliis glabris, coriaceis, ellipticis vel ovato-ellipticis, 5 x 2 - 8 x 3.8 — (8.5 x 4.5) cm, apice acuto, base longe cuneato, leviter decurrenti, nervis lateralibus approximatis (inter se 2 mm) parallelis rectis prope marginem bifurcatis gracilis utrinque prominentibus, costa striato utrinque prominenti. Flores (masculi) in racemis c. 1 cm longis congestibus, spissibus, axil- laribus aggregatis, pedicellis 2 mm longis, sepalis 4 ovatis 2 mm latis caduceis, petalis 4 simillimis, staminibus in 4 fasciculos dispositis, pistillodio fungiformi. Flores feminei ignota. Fructus globosa, 2 cm diam., nigricans, laevia, stigmate disciformi aliquantam quadrato integerrimo grosse papilloso 2 mm diam. promi- nulenti, sepalis nec persistentibus caduceis. JOHORE: G. Belumut, KEP 33830, KEP 33850 (holotype KEP), FRI 7511, 8730, 8737, 17825, SFN 10990. PERAK: G. Korbu, KEP 31486. Garcinia nigrolineata Planch. ex T. Anders. Fl. Brit. Ind. 1 (1874) 263. G. kunstleri King J. As. Soc. Beng. 59 (1890) 165 syn. nov. King cites collections by Wray and Scortechini. Ridley (Flora 1, 1922, 177) adds no others. I have seen at Singapore Wray 828, Scortechini 271, 1852 and s.n., probably all the syntypes. The fruit is not grooved and ridged, the stigma is 2.5 — 3.5 mm wide, papillose and slightly raised, but not on an umbo; the female flowers are 3 mm across on slender, 5 mm pedicels and contain a globose, very young fruit. G. kunstleri has not been recollected. Apart from the very small female flowers it fits well Corner’s diagnosis of G. nigrolineata in Gdns’ Bull. Str. Settl. 10 (1938) 36-8, and I have little hesitation in suggesting that it is but a weakly differentiated form of that species, best reduced to synonymy. On the Scortechini sheets I have seen have no locality notes. I do not know what led Ridley (loc. cit.) to say they came from the big limestone hill at Kuala Dipang. There is no limestone near Tapah, the Wray locality. The G. penangiana group (Group C) Pierre (Fl. For. Cochinch. fasc. 6, 1883, 37) described G. penangiana from Wallich’s collector (Porter) 4852D, a collection with male flowers in bud. He pointed out that Hooker had already mentioned it as distinct from G. cornea L. in a parenthetical addition to Anderson’s account of Garcinia in FI. Brit. Ind. | 276 Gardens’ Bulletin — X XVI (1973) (1874) 260. King saw further collections and amended the description in his account of Garcinia in the Materials (J. As. Soc. Beng. 59, 1890, 158). I follow what I believe to be King’s circumscription, although he does not precisely state which collections he saw, except for Kunstler 3853, which is not at Singapore. I have examined at Bogor, Teysmann s.n. from Riouw, P. Bientang, the type of G. caloneura Boerl. Cat. Hort. Bot. Bogor 68 (2) 1901. It looks very similar to G. penangiana. Dr. Kostermans has for some years now been labelling as G. celebica L. all specimens of this species-group in the herbaria he has visited. All these species are of very similar appearance and I see no prospects of resolving their limits without monographic study. There has been confusion between the G. penangiana group and the G. hombroniana alliance, my Subgroup Bi; this might indicate close affinity, implying that, as I indeed suspect, my species-groups, although convenient, are not entirely natural aggregates. G. bancana (Miq.) Miq. Fl. Ind. Bat. Suppl. (1861) 494. G. bancana var. curtisii (Ridley) Whitmore stat. nov. Basionym of variety: G. curtisii Ridley, Flora 1 (1922) 175, quoad syntypus Curtis 691 = lectotypus. G. curtisii Ridley differs from G. bancana, after which Ridley places it in his Flora, consistently in the fruits with the stigma protruding, more or less lobed and more or less coarsely papillose. The leaves tend to be narrower and are con- sistently pointed. It is undoubtedly very closely related and I reduce it to a variety. The fruits are exactly like those of G. penangiana which, however, always has its flowers borne terminally; in this variety a few are terminal, but most are in fascicles behind the leaves, as in G. bancana. I have selected one of the syntypes as lectotype of my variety. The other, Curtis 240, is typical G. penangiana. Collections seen: PENANG: Curtis s.n. May 1880, s.n. July 1890, 677; SFN 3034, 35322. JOHORE: SFN 25854. SINGAPORE: Ridley 6194. I provisionally also place FRI 4/50 from W. Kelantan here, though it differs in its fruits drying less wrinkled and with sunken stigma. Garcinia opaca King J. As. Soc. Beng. 59 (1890) 160. Apparently a rare species. I have not seen the syntype(s) from which King described the flowers; King’s Collector 5460 at Singapore, which I believe to be a syntype, is in fruit. G. opaca closely resembles G. hombroniana but (fide King) has no pistillode in the male flower, thus it falls in my Group C, whereas G. hombroniana, with pistillode, falls in Group B. As already noted I am not convinced B and C are natural groups, although they are very convenient ones. Both these species have the fruit wall thinly woody, and tending to fracture in the herbarium, this feature is more marked in G. opaca and especially so in its var. dumosa described below. Typically, G. hombroniana has a depressed globose fruit with the stigma on a distinct apical umbo, but sometimes the fruit is slightly elongate. In what I have Systematy of Malayan Phanerogams XVIII-X XII Zve described as variety A (see Tree Flora) the fruit is much longer than broad as in G. opaca, but I am not sure of the true affinity of this variety; its flowers have yet to be found. G. opaca var. dumosa Whitmore var. nov. A varietas typica in habitu plerumque fruticoso, foliis minoribus, ellipticis, 9 x 3.5 - 13 x 5—17 x 7 cm, petiolo graciliore breviore, 8-10 mm, calyce in fructu parvo demum cadenti, differt. PAHANG: Bt. Terom, Ulu Keniyam Mohd. Shah 1560 (holotype SING); G. Benom KEP 97842; Chini F. R. FRI 11066. JOHORE: Bt. Kuing SFN 28653; S. Berassau, Corner s.n. 18.1V.32; G. Chabang Tiga Jumali & Heaslett 4114; Mersing, Sg. Sarah working circle, KEP 77731. SELANGOR: Genting Sempah KEP 1094]. KEP 97842 is a tree, all the others are shrubs, except that the sheet I have seen (at SING) of KEP 1094] has no note on stature. Garcinia pyrifera Ridley J. Str. Br. Roy. As. Soc. 73 (1916) 140 (sphalm ‘pyriferum’). G. densiflora King J. As. Soc. Beng. 59 (1890) 171 (non G. densiflora Vieill. ex Pierre Fl. For. Cochinch. 5 (1883) 25) syn. nov. Ridley noted that his species was close to King’s except for certain foliage differences which he specified. I have compared the type of G. densiflora (King’s Collector 5933) with that of G. pyrifera (Curtis 3094) and all the subsequent gatherings of the latter and can see no. good differences, though two of the three leaves of the Singapore sheet are oblong-ovate, an unusual shape. In Flora 1 (1922) 179 Ridley reduced King’s species to G. glomerata Vesque (in DC. Monog. Phan. 8 (1893) 320), which predates his own G. pyrifera. I have been unable to check this because there is no authentic material of G. glomerata at Singapore or Bogor. Provisionally therefore I recognise G. pyrifera as the name for this tree. The flowers of G. pyrifera remain unknown. The description in the Tree Flora is taken from King’s of G. densiflora flowers, so far as I could confirm it from the poor specimen at Singapore. G. pyrifera is a species with the Gestalt of the other Malayan species of the Xanthochymus group (G. nervosa, G. dulcis, G. prainiana) and quite clearly dis- tinct from them in detail. Garcinia rostrata and its allies (Subgroup Aii) / Garcinia rostrata (Hassk.) Mig. Ann. Mus. Bot. Lugd. Bat. 1 (1864) 209. G. wrayi King J. As. Soc. Beng. 59 (1890) 152, syn. nov. Garcinia eugeniaefolia Wall. ex T. Anders. Fl. Brit. Ind. 1 (1874) 268. I provisionally keep G. rostrata and G. eugeniaefolia apart, not having seen the types of the former, on the stigma differences given by King, which are ‘flat to concave, smooth’ and ‘flat to domed, papillose’ respectively. The two syntypes of G. wrayi at Singapore (Wray 267, 362) show it clearly belongs to this aggregate. Unfortunately all the flowers have fallen; King’s description of the stigma as 278 Gardens’ Bulletin — X XVI (1973) ‘hemispheric, smooth, entire’ (my italics) suggests it belongs with G. rostrata. Ridley (in Flora 1, 1922, 169) reduced G. wrayi to G. merguensis, but I believe that is a fairly distinct species (see key in the Tree Flora) on foliage as well as flowers and that my placing of it is more probably correct. This whole complex, my Subgroup Ai, needs careful monographic investigation. Ca Garcinia merguensis Wight Illustrations of Indian Botany (1831) 122. G. lanceolata Ridley Flora 1 (1922) 170, syn. nov. I have examined the material at Kew, Kepong and Singapore and believe these two to be the same. Mesua The Mesua assamica group Mesua aff. assamica (King et Prain) Kosterm. Kayea assamica King et Prain Ind. Forester 27 (1901) 62. Symington first recorded this for Malaya when, at Kew in July 1939, he matched KEP 4643 (sterile), KEP 5495 (fruiting) and possibly KEP 10600 to H. G. Young s.n. December 1899 and 29 June 1900, two of the syntypes of K. (M.) assamica. Symington (ms. notes preserved at Kepong) observed ‘“‘These, I think, differ [from M. ferruginea]. They are not riparian trees and the fruits are probably smaller than K. (M.) ferruginea. They more closely resemble K. (M.) assamica K. et P. than anything else but the leaves are less cuneate and acuminate and the veination [sic] is less conspicuously dryobalanoid”’. I have placed a whole group of collections here which match KEP 4643 and 5495 in leaf, habit, habitat, and most especially in having the fruit enclosed in two, strongly adpressed, enlarged sepals. The fruits are long-stalked and solitary in 5495 but in open panicles in most of the new collections. King and Prain describe K. assamica as possessing panicles 3-6 in. long. The size of the fruit varies, as does the sepal surface texture and, slightly, the shape. I attach no importance to variation in the degree of splitting of the 2 sepals which I believe to be develop- mental. I have not seen the types of K. (M.) assamica, and following the caution expressed by Symington, name our material as aff. assamica. It is slightly hetero- geneous but is held together by the nature of the fruiting calyx, and, though further collections may reveal it to comprise more than one taxon, I have little doubt all these collections are related. FRI 12033 is an anomalous high altitude collection (780 m,G. Mandi Angin) with possibly galled young fruits which, on present evidence, comes here; and the sterile collection FRI 9347, from 960 m on G. Inas, Kedah matches it in leaf. I have segregated from the above a number of sheets as Mesua sp. D. These puzzle me not a little. They differ in having smaller leaves with more slender stalk, and smaller fruits, also more slenderly stalked. Fuller descriptions are given in the Tree Flora. At the extreme this taxon is rather distinct and KEP 110395, FRI 2617, 8106 approach $21745, 22469, 22507 from Sarawak named by Ashton (unpublished ms.) as M. acuminatissima (Merrill) Kosterm., but it merges into typical Malayan M. aff. assamica. Two collections, FRI 885], 10891, have the fruits solitary, the other collections have panicles. Systematy of Malayan Phanerogams XVIII-X XII 279 The problems of this species-group can only be finally resolved by a mono- grapher who will, I suspect, need more collections than yet exist. Collections seen: M. aff. assamica KEDAH: FRI 9347. KELANTAN: FRI 4054, 4099. TRENGGANU: Corner am. ¢ Now. 35, 12 Nov, 35, FRI 12033. PAHANG: KEP 4643, 5495, 77710. JOHORE: Kiah s.n. 19 Oct. 35, SFN 25973, 26140, 29272, Lake & Kelsall s.n., -28 Oct. 1892. re M. sp. D. | ; v 4 4 PAHANG: G. Tapis FRI 10071, 10115, 10891; FRI 8106. JOHORE: KEP ie FRI 2617, 8851. alt Y ih the fuer of flowers or fruits collections of the M. assamica group from Malaya are very difficult to distinguish from M. ferruginea on morphology. On the basis of size and habitat the following collections of fallen leaves are put under the M. assamica group: M. aff. assamica TRENGGANU: KEP 81070, 93779. KEDAH: KEP 81329. NEGRI SEMBILAN: KEP 105202. JOHORE: FRI 5088. M. sp. D. SELANGOR: KEP 76378. Mesua sp. E. Possibly this is another relative of M. assamica from which it differs con- spicuously in the shiny leaves with very faint nervation and not or scarcely reticulate. KEP 94924 has remarkable fruits enclosed in 2 big woody sepals plus, alternating with them, 2 lanceolate ones. In other respects this collection seems similar to KEP 76086 with fruits with the type of calyx found in M. aff. assamica. FRI 8125 is again similar in leaf. Its flowers are larger than in the collections of M. aff. assamica. The taxon is known from the east coast as follows: TRENGGANU: Bt. Bauk: KEP 76086. PAHANG: Bt. Beserah: KEP 94924; Ulu Sg. Anak Endau: FRI 8125. Mesua elegans (King) Kostermans Reinw. 7 (1969) 427. Kayea elegans King J. As. Soc. Beng. 59 (1890) 183 basionym Kayea caudata King loc. cit., syn. nov. Mesua caudata (King) Kostermans loc. cit. 426, syn. nov. The leaves of M. elegans, based on one flowering collection (King’s Collector 7346, K!), differ from those of M. caudata, based on one fruiting collection (King’s Collector 7937, K!). They are lanceolate, tapering gradually into a pronounced drip tip, the secondary nerves are faint, and a fine, faint reticulum can be made 280 Gardens’ Bulletin — X XVI (1973) out with the aid of a lens on the undersurface. The leaves of the type collection of M. caudata are much broader, taper suddenly into a drip tip, have the secondary nerves prominent, and the reticulum is almost invisible. In both, the leaves dry khaki-brown, the twigs are slender and tiny, and subulate ‘stipules’ and hypso- phylls are present. The two species are close, but on these collections they can be distinguished, as indeed King does distinguish them in his key. The numerous gatherings which have been made since 1885 show a complete range of intermediates, and I do not believe that more than one species exists. I have chosen to call it M. elegans because other species of Mesua in Malaya also have caudate leaves. M. elegans is in many ways a delicate version of M. kunstleri. Collections seen (all at KEP and SING except the types): PERAK: Batang Padang district: King’s Collector 7937 (type of K. caudata), G. Bubu: King’s Collector 7346 “(type of K. elegans); Keladang Saiong F. R.: FRI 5998; Parit F. R.: KEP 39492; Tapah: Ridley 14085. SELANGOR: Ulu Langat: FRI 12203; Klang Gates: KEP 532, 33224, 47132, Ridley 13521, 13527. PAHANG: Kemasul F. R.: KEP 10608, 10876. JOHORE: G. Panti: KEP 99232. SINGAPORE: Garden’s Jungle: Furtado s.n. 13 I 1931, Ahmad s.n. 10 III 1926. Mesua kochummeniana Whitmore sp. nov. Arbor magna, usque ad 36 m alta, 3 m diametro, ligno duro (ferreo); ramulis crassis angulosis subsparse squamatis, foliis ovato-oblongis, c. 22 x 9 cm, base rotundatis, apice rotundato-apiculatis, marginibus revolutis, laminis infra glaucis, supra nitide castaneis, nervis lateralibus approximatis parallelis, ex costa subper- pendicularibus, leviter curvatis, distaliter incurvato-approximatis arcuatis, petiolo crasso rugoso c. 12 mm longo. Inflorescentia racemosa, racemis 3—10 terminalibus, 4-7 cm longis, 3-6 floriferis, floribus subapicalibus, pedicellis 5 mm _longis, bracteolis 1 mm longis oppositis dettoideis; sepalis 4 ovatis 6 mm longis, petalis caducis (non vidi), staminibus caespitosis exsertis, filamentis tenuibus 7 mm longis. Fructus (probabiliter immaturi) c. 3 cm diametro, includentibus in sepalis duobus globosis truncatis dilatatis sublignosisque in base connatis, in sicco rugosis, parte basali styli crassi (2 mm longo) protrudento. Type: Corner SFN 29430, 14 May 1935, Johore, 134 mile Mawai-Jemaluang Road. Holotype in SING. Also examined: JOHORE: 13 mile Mawai-Jemaluang Road. Corner s.n. 31 Dec. 1934; 21 mile Mersing-Kota Tinggi Road. KEP 70811; Jemaluang F. R. FRI 2773, KEP 69992, KEP 71895. SOUTH PAHANG: G. Lesong F. R. KEP 83464, 86903. Probably the huge woody fruits 7 cm across (at KEP), collected without leaves as KEP 79193 from G. Lesong F. R. compt. 20, are this. All the collections cited except the type are at KEP only and all are sterile twigs or fallen leaves except KEP 69992 in flower. I name this fine tree in honour of the Forest Botanist at Kepong Mr. K. M. Kochummen. It is common in east Johore and the adjacent part of Pahang. Corner. Systematy of Malayan Phanerogams XVIII-X XII 281 in making the collection I have chosen as type, noted ‘24 hrs of sheer cutting to fell this tree ... wood exceedingly hard, like iron, as “‘penaga”’’ and that it is common on hillocks and drier parts of swamp around Sg. Kayu Ara and Sg. Buloh Kasap. Mesua kunstleri (King) Kostermans Reinw. 7 (1969) 427. Kayea kunstleri King J. As. Soc. Beng. 59 (1890) 182; basionym. Kayea rivulorum Ridley J. As. Soc. Str. Br. 54 (1909) 22. syn. nov. Mesua rivulorum (Ridley) Kostermans loc. cit. 429, syn. nov. ‘King described his Kayea kunstleri in the Materials from four collections: King’s Collector 3301, 6850, Curtis 1419 and Maingay 176. I have seen sheets of the first three of these syntypes at Singapore. I have no doubt that Ridley’s K. rivulorum belongs here, although it is known only from two syntypes, both in fruit, whereas all the collections I have seen of M. (K.) kunstleri are sterile or in flower. The argument for this conclusion is deferred until after the emended description of M. kunstleri var. curtisii below. In describing K. kunstleri in the Materials King noted ‘a shrubby form of this occurs in Penang (Curtis, Nos. 805, 1418) and in Perak (King’s Collector, No. 1345) in which all the parts are smaller and the leaves are less acuminate at the apex, and rounded instead of much attenuated at the base’. I have seen (also at Singapore) the two Curtis numbers and confirm King’s description. In 1894 in another publication, King raised the Penang form to species status as Kayea curtisii based on one of the sheets cited in the Materials, Curtis 805, plus another collection Curtis 748, which I have also seen at Singapore, and which is a good match. He made several serious mistakes in his description in firstly saying ‘leaf . . . base acute’ whereas it is rounded, secondly ‘flowers . . . leaf opposed’, whereas they are actually terminal, and thirdly ‘petals slightly smaller than the sepals’, whereas actually they are twice as long. The earlier note in the Materials and the syntypes clearly show however what was intended by K. curtisii. Ridley in Flora 1 (1922) 191 sunk K. curtisii into K. kunstleri commenting that Curtis 1418 (of King’s original note) is ‘obviously the same plant’ (? taxon) as K. curtisti, yet retained under K. kunstleri by King. Since 1890 numerous additional collections have been made, all from Penang and Kedah. King’s original note is amply confirmed, that there are two distinguish- able taxa, the extreme forms being very distinct. There are however few substantial differences, and the placing of a few collections has to be rather arbitrary. Never- theless I think it convenient and useful to give recognition to the ‘shrubby Penang form’. I therefore accordingly reduce K. curtisii to varietal status. The necessary formulation is set out below with a new latin diagnosis, necessary because of King’s mistakes. Mesua kunstleri (King) Kostermans var. curtisii (King) Whitmore stat. nov. Kayea curtisii King Ann. Bot. Gard. Calcutta 5 (1896) 144, t. 174B basionym. Mesua curtisii (King) Kostermans Reinw. 7 (1969) 426. syn. nov. A varietas typica in foliis minoribus, basi rotundato, nervis lateralibus abaxialis obscurioribus, differt. Lectotype Curtis 805. 282 Gardens’ Bulletin— X XVI (1973) Although, as noted above, I have seen no fruiting collections of M. kunstleri I have seen several of M. kunstleri var. curtisii. These have the fruit wall thin but not brittle, smooth, and irregularly covered with tiny, fawn, corky scales; the fruits are mostly round, tipped by the 1-2 mm long style base, the largest is 3.3 cm across. The fruit is seated on the persistent sepals which are not enlarged. Both syntypes of Kayea rivulorum Ridley are in fruit, young in Ridley 7349, and, in Goodenough 1976, 4.5 cm long and ellipsoid, including a stout 1 cm apical beak. The wall is wrinkled and bears the same tiny scales typical of M. kunstleri var. curtisii. These fruits differ from those of M. kunstleri var. curtisii in drying wrinkled and in their shape. King, in his description of M. kunstleri and later as plate 173 in Ann. Roy. Bot. Gdn. Calcutta 5 (1896), depicts fruits of this shape, and there is a single Curtis collection of M. kunstleri var. curtisii at Singapore with neither number nor date which has them (this sheet I have labelled X). I consider the resemblance in fruit and leaf adequate to justify reduction of K. rivulorum. The types of K. rivulorum have the leaf base slightly rounded, whereas in the types I have seen of K. kunsileri it is long cuneate (and is thus described in the type description and depicted in plate 173 in Ann. Roy. Bot. Gard. Calcutta 5, 1896). There are 3 collections of Curtis 557] at Singapore, of these 2 have cuneate and one slightly rounded leaf bases. All are dated March 1901 and all bear flowers. I think in this case we can assume they come from a single gathering of one tree, though Curtis often did place different gatherings under one number. Amongst the other abundant material at Kepong and Singapore this character breaks down. I do not think the shape of the leaf base is useful in this case to distinguish two taxa. M. kunstleri is in many respects a coarse kind of M. elegans with bigger leaves and flowers. It differs from M. nervosa (in the circumscription of which | have been assisted by manuscript notes made by Symington at Kew) in the lack of harsh red tomentum. Monographic study is necessary to delimit more closely these two species. Mesua lepidota Anderson FI. Brit. Ind. 1 (1874) 278. KING J. As. Soc. Beng. 59 (1890) 185. Anderson based M. lepidota on Griffith (K.D.) 845 from Malacca. King added Scortechini 183b and King’s Collector 4551, 5881, recording that these differ in smaller leaves, more slender twigs and slightly thinner pericarp. He went on to note that these collections might be a second species, also possessing the very remarkable woody reflexed fruiting calyx which had prompted Anderson to suggest his species was ‘probably a new genus between Kayea and Mesua’. The abundant gatherings which have now been made show there clearly are two taxa, but a few collections are intermediate (KEP 32163, 71830, 76734, FRI 11321). It is convenient to give a name to the delicate form especially as it is very similar in the sterile state to M. nuda; the two are common together in the Ulu Gombak F. R. None of the cited material is represented in the Far East. Symington however has matched KEP 12906 to Griffith 845 and KEP 37290 to the slender variety. In fact, I believe, from examination of Burn-Murdoch (=CF=FMS=KEP) 382, the single sheet extant of Kayea parviflora Ridley (kindly loaned to me at Kepong by Kew) that this little known taxon is the same thing in young flower bud, and it therefore provides the epithet for the variety, as follows: Mesua lepidota var. parviflora (Ridley) Whitmore stat. nov. Kayea parviflora Ridley J. As. Soc. Str. Br. 82 (1920) 170, basionym. Systematy of Malayan Phanerogams XVIII-X XII 283 Mesua parviflora (Ridley) Kostermans Reinw. 7 (1969) 428 syn. nov. Type: Burn-Murdoch 382-(at K). Also examined (at KEP and SING): PERAK: FRI 11937. TRENGGANU: Corner s.n. 9 XI 35, SFN 30480. SELAN- GOR (all Ulu Gombak): KEP 10929, 13064, 12633, 12855, 37290, 93292, 94717, 94718, FRI 1154, Hume 9270. MALACCA: Derry 63, 902, collection o¢ (at SING, no name or date). JOHORE, G. Blumut: SFN 10790. In the Perak collection the fruit is young and has not developed a woody calyx. The Johore collection has the leaves 5 X 1 cm and broadest near the base with a long narrowed upper portion. These collections may represent slightly divergent populations. The Trengganu collection, SFN 30480, is the only one I have seen of the variety with the flowers in racemes. The collections I have examined suggest the typical variety inhabits plains, and var. parviflora is a tree of the hills. The four intermediate collections cited all come from hilly places. Mesua nivenii Whitmore sp. nov. A M. wrayi valde simillime sed in floribus sessilibus et foliis infra conspicue reticuloso-venulosis differt. Arbor parva, foliis ovato-oblongis, c. 9 X 4 cm, basi late-cuneatis, apice acutis, coriaceis, nitidis, nervis lateralibus subtus tenuibus prominulentibus; pagine infra reticulato, nervis tertiariis valde approximatis, petiolo 1 cm longo ruguloso. Inflorescentia racemoso-paniculata, racemis ramificatis, pedunculis 1 cm _ longis, pedicellis obsoletis (floribus sessilibus) trifloriferis, sepalis ovatis rotundatis con- vexis in sicco nigris rugosis 6 mm longis, petalis c. 10 x 6 mm obovatis rotun- datis, staminibus exsertis caespitosis. Fructus (submaturus?) includentibus in sepalis duobus paribus, sepalis maturitate incrassatis coriaceis rugosis turbinatis, parte basalari stylo 2 mm protrudento. Type: Whitmore FRI 12100, 12 July 1968. Trengganu, G. Mandi Angin, Summit ridge north of birop (trig. point), 1425 m. Holotype in KEP, isotypes K, L, SING, ARN. In flower. Also examined: a, TRENGGANU: leading E. ridge G. Mandi Angin: FR/ 1081/5; G. Padang SFN 31064, 33909 (in fruit). | First collected by Moysey and Kiah on their pioneering journey up the Ulu Brang to G. Padang in 1937. Found in full flower by myself, and a delightful sight, on the summit ridge of G. Mandi Angin in 1968. Named for the part-time Superintendent of the first Singapore Botanic Garden who ‘had a fine eye for design’ (Purseglove). Mesua nuda Kostermans ex Whitmore sp. nov. Arbor, ramulis gracilibus teretibus plerumque griseis, foliis ellipticis (5 x 1.5) — 6 x 2—8 x 3 cm, base cuneatis, apice cuspidatis, subcoriaceis, nervis laterali- bus utrinque tenuissimis approximatis parallelis leviter arcuatis ex costa subper- pendicularibus, petiolis gracilibus 3 mm longis. Flores (in alabastro) cum pedicellis 2 mm longis. Fructus solitarius, axillaris, pedunculo incrassato (a 5 mm), pedi- cello 2 mm longo, maturitate oblongo-ovoideis 4 cm longis 2.5 cm latis, apice interdum rostellatis, base rotundatis, paricarpio fructi tenuoso-fragile bubalino laeve squamulis minutis suberosis obtectis, sepalis caducis. Semen unicum in fructu, testa fusco nitido. 284 Gardens’ Bulletin — XXVI (1973) Type: Whitmore FRI 3440, 24 March 1967. Pahang, G. Benom, Ulu Sg. Krau, ridge crest 270 m. Holotype in KEP, isotypes K, L, SING, ARN. Also examined: KELANTAN: KEP 98807, FRI 2333, 2339, 7255. TRENGGANU: SFN 30390, FRI 10682. PAHANG: KEP 4059, 29682, 100105, 108974, FRI 15333. Baloh F. R.: KEP 3149, 97913, FRI 3798. KEDAH: KEP 105354. PENANG: Curtis 748. PERAK: KEP 29928; s.n. Wyatt-Smith 25 Jan. 1953. SELANGOR: Hashim s.n. (SING), Burn Murdoch (KEP ?) 278 = 14248. Ulu Gombak F. R.: KEP 10942, 11220, 18163, 18165, 98401, FRI 2555. MALACCA: (illegible) 85/, Alvins 1066, Derry 1003 no locus; Symington KEP 18163. NEGRI SEMBILAN: KEP 104871, FRI 11347. JOHORE: KEP 69996. Mesua nuda is a very common species, apparently endemic, which unfor- tunately has never been found in full flower. It is highly distinctive in its brittle, buff, naked fruits. The name is in wide usage but as far as I can discover Koster- mans has never published it. Young fruits are sometimes globose, leaf size is slightly variable as is the clarity of the secondary nerves, but I have little doubt that all the collections cited belong to this species. Mesua purseglovei Whitmore sp. nov. Arbor excelsa, foliis coriaceis lanceolatis, 13 x 4 cm, lamina prope basin quam apicem latiore, base rotundato, apice acuminatissimo; nervis lateralibus majoribus ex costa subperpendicularibus parallelibus leviter curvatis distaliter incurvato con- junctatis subtus conspicuis supra inconspicuis subdepressis, nervis lateralibus secundariis minoribus (1-3 aggregatis cum majoribus alternantibus) subtus incon- spicuis, nervis tertiariis (reticulationis) evidentibus. Flores ignoti. Fructus solitarius (variter geminatus), axillaris, pedicello crasso 1 cm longo, pedunculo 4 cm longo, in sepalem '+ inclusus (sepalis 4 maturitate valde dilatatis ovatis subcoriaceis furfuraceis depresso globosi diametro 3-3.5 cm). Type: Holttum SFN 31233, 14 May 1936, Pahang, Cameron Highlands, mile 47 Telom Rd. Holotype in SING. This is a high mountain species and I think likely to prove to be endemic, I therefore describe it, despite its being known from only one collection, in fruit. M. purseglovei is similar to M. wrayi but conspicuously differs in leaf characters. XIX. LECYTHIDACEAE T. C. WHITMORE Barringtonia payensiana Whitmore sp. nov. Species nova, a speciebus ommibus generis Barringtoniae in fructibus pyrami- dalibus differt. Arbor parva, foliis obovatis acuminatis chartaceis vel subcoriaceis, c. 21 x 6.5 — 50 x 14.5 cm, base longe decurrente, marginibus leviter revolutis, apicem versus leviter dissitiserratis, nervis lateralibus in paribus c. 26, petiolo 15 mm. /nflores- centiae et flores ignotae. Fructus pyramidata usque ad 4.5 cm lata, c. 4.5 cm longa, base omnino plano, plus minusve quadrata in sectione, superficie ruguloso, calyce persistenti usque ad 15 mm diametro. SELANGOR: Ulu Batang Kali FRI 4550 (holotype SING). PERAK: Pangkor Island SFN 31658. Named in honour of the recent monographer, Dr. J. P. D. W. Payens. Systematy of Malayan Phanerogams XVIII-X XI] 285 XX. LEGUMINOSAE (d) T. C. WHITMORE Crudia viridiflora Whitmore sp. nov. Arbor, foliis simplicibus subcoriaceis, glabris, oblonge-ovato, 15 X 6 — 22 X 8 cm, base cuneato, apice rotundato, apiculato, costa supra canaliculata, nervis lateralibus c. 9 paribus, pagina inferiore valde prominentibus, pagina superiore obscuris, nervis lateralibus secundariis reticulisve similiter, petiolo crasso rugoso 5 mm. Flores congesti, viridi, in racemos terminales, solitarios vel — 4 fasciculatos, 7-12 cm longi, rhachidi aureo pubescenti, bracteis minutis aureo tomentosis, pedi- cello 3 mm aureo tomentoso, sepalis ovoideis 6 x 4 mm exteriore ad basim leviter tomentosis, staminibus 8 exsertis, filamentis filiformibus 10 mm, ovario 5 mm longo aureo lanato, stylo gracili 5 mm. Fructus ignoti. PAHANG: (G.) Lesong F. R., Sg. Jekatih FRI 15949 (holotype KEP); Samsuri 540. This distinctive new species was discovered too late for inclusion with the Leguminosae in vol. | of the Tree Flora and will have to go into a supplement later. It comes near to C. gracilis Prain apud King and C. subsimplicifolia Merr. in De Wit’s monograph (Bull. Jard. Bot. Buitenz. sér 3, 18, 1950 407-34) but differs from both. It can easily be distinguished from all the other Malayan species from its simple leaves. I did not see any compound leaves on the limbs which were cut for me from the crown of the tree from which the type collection was made. XXI. STROMBOSIA (OLACACEAE) T. C. WHITMORE Strombosia maingayi (Masters) Whitmore comb. nov. Anacolosa maingayi Masters FI. Brit. Ind. 1 (1872) 580 basionym. Strombosia rotundifolia King J. As. Soc. Beng. 64 (1895) 103 syn. nov. Both names are based on the same type sheet, Maingay (Kew Distribution No. 1019), and the former clearly has priority. When King transferred Masters’ species to Strombosia he incorrectly coined a new epithet for it. Ridley (Flora 1, 1922. 425) cites A. maingayi as a synonym of King’s species. Strombosia multiflora King J. As. Soc. Beng. 64 (1895) 102: syntypes Perak, King’s Collector 7824; Penang, Curtis 859. King described this and his S. rotundifolia (see above) at the same time, noting that they are closely related. The name S. multiflora has never entered currency. The abundant material at Kepong and Singapore appears very homo- geneous. I can see in it no significant difference in the three differential characters given by King, namely the number of flowers per cymose fascicle (the pedicels, it should be noted, are brittle and flowers easily break off), flower size (all have tiny flowers only c. 3 mm long, on a 1.5 mm pedicel), or leaf shape and size (King’s contrast of rotund, 2.5 x 2 in., versus 2.5 x 1.5 — 4 x 2 in. seems in any case to be weak). There is variation between collections with sessile, round to oblong, fruits and those with stalked, round fruits. The former group has small, very woody fruits which might perhaps be galled. 286 Gardens’ Bulletin— X XVI (1973) Mr. L. L. Forman has kindly made a very thorough examination of the Kew material of S. maingayi (rotundifolia) and S. multiflora. Not one collection is in fruit. He could see no leaf or twig differences and the material was too incomplete to study the buds. He did however find a substantial difference in the flowers, along the lines indicated by King; to quote ‘the Kew isotype of Str. multiflora has dense bunches of flowers with petals 5 mm long and styles 3.5 mm. Srr. rotundifolia on the other hand has petals only half that length and a very short style less than 1 mm... These species are surely distinct’. He later reported that in both species the flowers have 1.5 — 2.0 mm pedicels. The Singapore isotypes which had been mislaid were subsequently rediscovered, examination of them confirmed Forman’s careful observations. The problem facing us in Malaya is thus a very tricky one. On the one hand, the two species appear to differ substantially in flower but the types are inadequate to discern any other differences. On the other hand, all the collections im the East (many of them duplicated at Kew) appear to represent one species, except Curtis 1371 from Pangkor Island, but they are mostly sterile or in fruit. That some collections have sessile fruits does not help as both species have pedicellate flowers, and in addition such fruits are suspected possibly to be galled. Since King described S$. multiflora no further collections have been added to the species, except Curtis 1371, just mentioned, which was misnamed ‘? Anacolosa sp.’ Ridley in his Flora cites just the two syntypes. All the abundant material at Kepong and Singapore has been named S. rotundifolia (=maingayi). The few flowering collections amongst this abundance have the tiny flowers and short stigmas of S. maingayi, and are a good match for the fruiting and sterile collections. I conclude that the pragmatic solution is to name all these collections S. maingayi. Although S. multiflora differs conspicuously in flower more collections than have been assembled in the last 76 years will be needed to discover if these two species can be differentiated in any other way. Thus, in summary, I propose to follow the practice since 1895, in putting our material into S. rotundifolia, but have had to revert to the earlier epithet for this taxon which becomes therefore S. maingayi. Curtis 1371 from Lumut, Pangkor (Perak) at Singapore might represent S. multiflora in that the style is very long (4 mm) and thread-like, but the flowers are only 2 mm long. XXII. HELICIOPSIS (PROTEACEAE) K. M. KOCHUMMEN Three new species of this small genus, only differentiated from Helicia in 1955, have been discovered in preparing the account for the Tree Flora. Heliciopsis cockburnii Kochummen sp. nov. Arbor ad 24 m alta, 60 cm diam., ramulis striatis junioribus ferrugineis, foliis (foliis pinnatisectis adhuc ignotis) ellipticis vel anguste obovatis, 9.5 x 2.8 — 21 x 7 cm, apice obtuso, base angustato decurrenti, nervis lateralibus 6-11 paribus arcuatis, petiolo 1-4 cm longo base incrassato. Flores adhuc ignoti. Infructescentiae 10 cm longae rhachidi striata rubiginosa gracili 2 mm diam., bracteis linearibus 6 mm longis rubiginosis sub-persistentibus, bracteolis bracteis similibus 3 mm Systematy of Malayan Phanerogams XVIII-X XII 287 longis, pedicellis 4 mm longis 2 mm crassis. Fructus oblonga nitida siccitate brun- neola 2.5 cm longa 2 cm diam., stylo vestigio, exocarpio tenuissimo, mesocarpio radiato fibroso, 3 mm crasso, endocarpio tenuissimo. Semina leaves, testa brunneola. PAHANG: G. Tapis 800 m FRI 10934 (holotype KEP). PERAK: Keladang Saiong F. R. KEP 2900, KEP 32181. UPPER PERAK: FRI 594. Heliciopsis montana Symington ex Kochummen sp. nov. Arbor 26 m alta, 60 cm diam., ramulis striatis cum cicatricibus foliorum delapsorum notati, foliis simplicibus coriaceis, ellipticis vel anguste oblongis, (9 x 3.5) — 15 x 6— 21 x 10 - (40 x 20) cm, apice obtuso, base cuneato raro inaequilatero, nervis lateralibus 6 — 10 (- 14) paribus arcuatis, pagina supra nitida, petiolo 2 — 4.5 cm longo, ad base gibboso, foliis pinnatisectis 48 x 24 — 54 x 27 cm, lobis 3-5 paribus oblongis 17 x 3 — 19 x 7 cm, apice acuto, nervis lateralibus 9-11 paribus arcuatis, petiolo 10-13 cm longo. Flores adhuc ignoti. [nfructescentiae rhachidi crassa 5 mm diam. striata ferruginea, bracteis adhuc ignotis (? cadicis), pedicellis crassis 6 mm longis 3 mm diam. Fructus siccitate purpurea vel brunneola oblonga 2.8 — 3.8 cm longa x 2.5 — 3.5 cm diam., exocarpio coriaceo tenuissimo, mesocarpio fibroso 3 mm crasso, endocarpio tenuissimo leavis. PAHANG: Fraser’s Hill FRI 16172 (holotype KEP). Cameron Highlands KEP 27194, 30966, SFN 32871. PERAK: G. Korbu KEP 32248. A species of mountain forests at 300 m and above. Endemic. Symington named this Helicia montana in manuscript. Heliciopsis whitmorei Kochummen sp. nov. Arbor 18—21 m alta, trunco usque ad 40 cm. diam., ramulis striatis, junioribus ferrugineis, foliis simplicibus coriaceis ellipticis vel oblongis 7.5 x 2.5 -— 21 x7 cm, siccitate viridiflavis, apice acuto, base acuto, nervis lateralibus 9-12 paribus, arcuatis, utrinque inconspicuis, costa supra conspicua, petiolo 2-5 cm longo base incrassato, foliis pinnatisectis 46 x 28 — 60 x 30 cm, lobis c. 11 paribus, oppositis, 12.5 x 2—15 x 3 cm, petiolo 9-14 cm longo. Flores adhuc ignoti. /nfructescentiae 15 cm longae, rhachidi striata glabra 3-4 mm diam. Fructus cylindrico-truncata 18-20 mm longa 8-11 mm lata, stylo persistenti 1 mm longo, exocarpio tenuissimo. mesocarpio radiato fibroso 1 mm crasso, endocarpio 0.3 mm crasso. TRENGGANU: G. Mandi Angin FRI 12062 (holotype KEP). Ulu Sg. Trengganu 900 ft FRI 10544. Ulu Bendong Kemaman, 700 ft SFN 30/93. va -custdeluligole ditriatce oteptenodscd penne. vo , insealine ine power on is UES le dud Nowe has zeebaladh 1 hA Fale (SEL, oepgiaiorny hup ete LA Pie la Ae J ' 3 : re ‘a1 { -* +s fas toro? s neck kor Lone aignoldda:dtuges dod a eloitetls ae oget: beanie — coeutdt ets mat FF sdoinny yi nithe binky cane tts OF @ apie my yoyo = eX eG) ganeo iy : ae dinatio’3 ah Sty JA EGET Coda at UVR clue oned” onbSh” shite ave sac) Oghol ins SSO Misigedos "Brahe ‘Bieoy eit pont glizoqgy dike 1) 9 vor ae GE eo): - 230") OP vis iil SOUNSAZST Ue onl oudba weil * Oat A mp S10 oloitag mat porte “epiibnd yd) umoalnuinalip agin 4.2 cp alet chanel Dinero Th ORAS 0 gro! mie iy dasmigocratyia: ve) ef! eee ondaty cum £.0 cigrssobes .ozes7s pum | ozordéo vaggonstT 9é wll Bib sqviolod) SOOC\ \A4 aia oft ibn | 3 i. peat SF ER ae KM. Kieceoseeaia’ a a4 7 ant ee e535. Aa bo oewerad GH Mpeg she Avex mnt * ae oven ae a 7 e er oP t . co ) aT T ’ ios my Ret a P as drm ag 1 ales, -qewallgtiias, cs ‘Colic meumfleectee ecu hercis) tipticig } eng, yplee. obtuse, base angreinge, dee wt vere i, petiole Ion, lenge By eee chactn Sie | i : , pices ya! en) ce A survey of the occurrence of epiterranean soil algae in Singapore Island by ANNE JOHNSON Department of Biology, Nanyang University Singapore Epiterranean soil algae found in the upper layers of soil are of considerable importance in soil economy since they increase the organic content of the soil by photosynthesis, disintegrate soil particles to increase the water-holding capacity of the soil and secrete mucilaginous material binding surface particles together to form a matrix; while heterocystous blue-green algae fix atmospheric nitrogen increasing the nitrogen content of the soil (Johnson, 1962). Subterranean soil algae which live in complete darkness and may lose their photosynthetic pigment (Tiffany, 1951) are less important to soil economy. Few investigations have been made on the soil algae of tropical areas and include those of Fritsch (1970) in Ceylon; Prasad (1949), Gonzales and Valavigi (1960), Singh (1940-41), Sulaiman (1944) and Ghose (1923) in India; Bristol (1919) - in Malaysia and Johnson (1962, 1969) in Malaysia and Singapore. Over a large area of Singapore Island the soil is derived from sedimentary parent rocks of the Triassic or Carboniferous period, with small areas derived from igneous rocks of acid or basic composition, as well as alluvial soils in swamps or valley bottoms. Most areas in the West of the Island the soil was derived from argillaceous series including red iron-rich strongly lateritic soil, light coloured to white iron-poor shale-derived soil, as well of soils mixed composition (Panton, 1955). Sedimentary soils of arenaceous series are commoner in the East of the Island including both coarse and find-grained series. The red colour of laterized soils is due to their high iron content since laterisation leads to an accumulation of iron and aluminium in the upper layers (Becking, 1961). Most of the iron is unavailable to the plants being in the form of insoluble iron-aluminium sequioxides or ironstone concretions. Most soluble cations such as calcium, magnesium, potas- sium and sodium have been leached away, and Singapore soils are extremely deficient in cations. The soils are also low in available phosphate due to loss by leaching and have a rather low pH. The rapid decomposition of organic matter under tropical conditions and subsequent leaching make them typically moderately low in nitrogen. Singapore Island was originally covered with tropical lowland forest with dipterocarps as the dominant type of trees. During the second half of the nineteenth century considerable areas of the Island were cleared for the planting of gambier and pepper (Jackson, 1968). The soil rapidly became exhausted leaving a very infertile soil. When gambier and pepper cultivation was abandoned early this century, the land was allowed to revert to grassland and scrubland (belukar). In other areas the top soil has been removed in reclamation operations and exposed 289 290 Gardens’ Bulletin — XXVI (1973) subsoil (B layer) remains. Forest is extremely limited in extent. Cultivated areas support vegetable crops, and estates have rubber, coconuts, rambutans and other fruit. Disturbed areas are city dumps, abandoned building sites, army camps etc. Materials and Methods Surface soil was collected from 100 habitats in the rural area of Singapore to a depth of 2 cm. Two series of cultures were set up for each type of soil: — (i) Plate culture, consisting of soil placed to a depth of 1 cm. in sterile petri-dishes, moistened with sterile distilled water and (ii) Liquid culture, consisting of small quantities of soil introduced into sterile Knop’s solution in conical flasks under sterile conditions. Both types of culture were exposed to 12 hours natural light per day. Observa- tions on the algae appearing in the cultures continued for two months. After setting up the cultures the remaining soil was air-dried and sieved through a 2 mm. sieve before analysis. The soil type was characterised into clay, clay-loam, loam, sandy-loam, loamy-sand, sand and sandy-clay according to the method of Leamy and Panton (1966). The soil was analysed for percentage of water, percentage of organic matter, ppm. nitrate, phosphate, calcium, ammonium nitrite, magnesium, iron, aluminium, sulphate, chloride, potassium and carbonate using the methods of Jackson, 1958 and those of the Edwards Laboratory, Ohio. pH was determined by use of a pH meter on soil paste. Colour was determined using standard Munsell Colour Charts to give hue, colour value and chroma on both wet and dry soils. Results 1. Soil type The soil type characterised according to Leamy and Panton (1966) in different habitats is seen in Table I. Surface soil consisted predominately of clay (24%) or clay loam (39%). Sandy loam and loamy sand amounted to 10% and 13% respectively. Sands (6%). sandy clays (2%) and loams (6%) are rare. Forest soil is usually clay loam (66.7%) but clays and loams occur. Alluvial soil is usually clay (57.2%) with some occurrence of clay loam, loam and sandy loam. Exposed subsoil (B-layer) is usually clay or clay loam with rare occurrence of loamy sand and sand. Grassland may be found on all types of soil. Scrubland (belukar) occurs on all except sand and sandy clay. Estate soil (rubber, banana, rambutans etc.) included no example of clay or sandy clay; while disturbed soil (dumps etc.) may be found on clay, clay loam or sandy loam. Note. In the tables the following abbreviations are used for different habitats : — F = forest soil; A = alluvial soil; E = exposed sub-soil (B-layer): G = grassland: S = scrubland (belukar); C = cultivated soil: Est. = estate soil; D = disturbed soil (city dumps etc.). Samples were collected in the Jurong, Bukit Panjang, Serangoon and Katong districts of Singapore. Epiterranean soil algae in Singapore 291 TABLE | Percentage samples with different types of soil CLAY ss mae J «Sor JE de ee A 333 Bs 16.7 24 coamysann |. |. | se[ ital wo | na faes] . | ool a EES CS 5 ae a 2. Soil Analysis The percentage water in air-dried soil, percentage organic matter; and nitrate, phosphate, calclum, ammonium, nitrite, magnesium, iron, aluminium, sulphate, chloride and potassium in ppm. are given in Table 2. No soluble manganese was detected by the methods used. The percentage of soils containing carbonate is given in Table 3. The water holding capacity of dry soil is closely correlated with the organic content being high in forest soils and alluvial soils but low in all other soils. Nitrate and calcium levels were normally at moderate to moderately low levels especially in alluvial or cultivated soils but all other ions were at extremely low levels. The values for phosphate, sulphate and potassium are critically low in most souls. Most soils are acid, forest soils having an average pH of 3.83 with the pH somewhat higher in other types of soil especially in disturbed soil where an average pH of 6.45 was observed. This is correlated with the higher levels of potassium, calcium and carbonate in this type of soil. Carbonate was detected in only 17% of the samples. It is completely absent from alluvial and estate soil, but occurrs in 83.3% of disturbed soil (dumps etc.). There are no calcareous rocks on Singapore Island. 292 Gardens’ Bulletin — XXVI (1973) TABLE 2 Soil Analysis. Mean values for different soils °% water (dry soil) .. | 6.29 |10.81 | 1.47] 1.54 | 233 | 192] 109] 145 2.53 % organic matter .. | 18.09 |12.62 | 4.95 | 7.66 | 6.29| 7.17| 5.19 | 647| 7.18 _ NO3 (ppm.)__.. g 8.6 | 14.16 | 6.00] 9.56 |11.00 |21.11 | 13.60 | 17.50 | 10.04 PO, x .. | 0.16 | 1.08 | 0.23 | 0.56 | 0.70| 1.33 | 1.50| 0.33 | 0.66 ca [1000/1166 |14.12 [1437 | 8.00 |13.30 12.70 /1330 13.08 NH4 2.33 | 0.67 | 0.70 | 1.87 | 2.40 | 3.50 | 3.60 | 233 | 1.95 pH 3.83 | 4.88 | 4.95] 5.80| 5.00] 5.67) 5.40, 6.45| 5.26 NO» EEE 0.09 | 0.17 0.08 Mg 0.66 | 4.83 | 1.88 | 2.31 1.80 | 2.66 | 3.70 | 3.00) 2.41 Fet* 1.66 | 1.00 | 1.65 | 0.65 | 0.80 | 0.89 | 1.80 | 0.33 1.02 Al = o4a7 oa; Pia ome Mn Lee, H162 url lt arshly af tiuecSlelvmecia |satinal aanaeen nnn SO, t, vol T8667) 333 [Poss Plo) oo go gee nae Cl ‘ |... 3.33 | 1.76 | 003 | .. | O01) 400 ee K pay 1.50 | 1.67. 5.00 | 0.60 Colour value (dry) is 80.1 53 167 (ee 1 oe Tae 6.1 63 | 5.9 (wet) | 33 | 40°R4N 42 | 43 | 42 |. 460) aOul ee Chroma (dry) .. -- 28 | 43 | 49 | 36 | 26) | ae) ) aie 3.5 (wet) .. Serer ee eee ne me pe 4. | 3.2 TABLE 3 Percentage Samples Containing Carbonate | of = f | | fF OC + CO3 Sa ee ey 7 11.8 9.4 | 30 35.3 is $3.3-], AE —'CO3 ne .- | S30 | 100 | "66.2 | 506) 70 66.7 | 100 | 16.7 83 3. Soil Colour The colour values in Table 2 reflect the somewhat lower values of alluvial and forest soil, and the relatively higher values for exposed and estate soil. The chroma is lowest in forest soils. The hue of wet soils all occurred on the YR colour range of the Munsell colour charts (Table 4). In spite of the uniformity of hue with 60% of the soils in the 10 YR range, Singapore soils are very variable in colour (Table 5). Epiterranean soil algae in Singapore Hue 10 YR 5 YR 5 YR 2.5 YR TABLE 4 Distribution of hue in 100 samples (wet soil) ———— | fl | | | | EE TABLE 5 293 Number of soils with different surface colour (Standard Munsell terminology-wet soil) Strong brown Dark brown Brown Pale brown Very pale brown Very dark greyish brown Dark greyish brown Greyish brown .. Dark Reddish brown Reddish brown .. Light reddish brown Dark yellowish brown Yellowish brown Pale yellowish brown Black Very dark grey .. Dark grey Red Yellowish red Pink Reddish yellow .. Brownish yellow Light yellowish brown ee Nee 2 1 Z ibe to pdranfuegedeino>| 2 | 3 | 2 hartge Yo <0g Ipr0t g eajamege Tp og | voqeifrog | 4 —}—}—- +--+ ————}— mandrel treat siete dort ee bo le per Oh Bent hops Hr aiibeoo Pay (pS ogy Te ge gee a mela ge a yea ind ||} eee Te incr sats athe Ud ieee dedinad —- +--+ — —}—|—} |_| — —- — Ae a a I + —— sj} meme iataiiih } | ||| + oe ee ae a ——}—}——_ —— | 2 ae ae ee | | — —_-—__—_ com ee | Goan a Total —) pl 294 Gardens’ Bulletin — X XVI (1973) 4. Soil Algae (a) constancy of occurrence. The appearance of algae in sterile culture indicates the presence of these algae, at least as resting spores, in the soil. As has been reported frequently, the frequency of algal species in culture does not necessarily reflect their frequency in the soil since the conditions of culture may favour some species more than others. Although frequency estimates of growth in culture were made, these were considered to have no direct relationship with the actual frequency in the soil and these results are not reported here. The constancy rather than the frequency of occurrence of algal species was considered to be of greater significance since it gives a measure of the number of soil samples in which the algal species occurs. 80 species of algae were recorded in the cultures and they were distributed with constancies shown in Table 6. The soil examined show a great diversity of algal flora with very few constant species. The typical J-shaped curve of constancy / no. of species is common in tropical floras (Johnson, 1966). TABLE 6 Contancy and number of species (Constancy = no. of samples; total no. of samples = 100) Constancy ..| 85 | 29 | 19 | 17 7 6 5 4 a] 2 | 1 mi | | | LE LN TT EE No. of species .. 1 1 1 1 5 1 1 5 7 Total no, of species = 80 Table 7 gives the habitat occurrence of species with a constancy of more than 5% (only 11 species have a constancy of more than 5%). Although the list includes 5 species of blue-green algae (with asterisk) none of these are capable of fixing nitrogen under aerobic conditions. TABLE 7 Number of Occurances of Algal species with a constancy of more than 5% a | F A EB G Ss * Est. D Total DESMOCOCCUS VULGARIS Brand. 74 a 5 5 15 30 a 8 10 3 85 HORMIDIUM FLACCIDUM (Kitz.) Br; 0 1 52 13 0 1 1 1 19 * OSCILLATORIA JASORVENIS Vouk. ori antares per pa po ae *OSCILLATORIA OBSCURA | PALMELLA MINATA Nag. .. 7 1 7 10 3 2 4 29 Briihl. et Biswas Epiterranean soil algae in Singapore 295 TABLE 7 — continued. Number of Occurances of Algal species with a constancy of more than 5% Roo” E G S Cc Re om | Tent *SYNECHOCOCCUS AERUGINOSUS | | Nag. a2 ars 0 0 4 2 0 1 OOP rian 7 TROCHISCIA ASPERA (Reinsch) Hansg. uO eh 1 0 0 1 ee 1 i 7 PINNULARIA sp. bis 0 2 0 3 0 0 2 0 7 NAVICULA sp. .. "a 0 0 2 1 0 1 1 a 7 * CHROOCOCCUS MINUTUS (Kiitz.) Nag. & 0 ys Z 0 0 0 2 0 6 (Kiitz.) Nag. he 0 0 1 i} * CHROOCOCCUS TURGIDUS (b) Blue-green algae. As in other tropical soils, blue-green algae (Cyanophy- ceae) form a conspicuous part of the algal flora. In this investigation 41 species of blue-green algae are reported from Singapore soils. These may be divided into the non-heterocystous algae which do not normally fix nitrogen under aerobic conditions, and the heterocystous algae which are capable of nitrogen fixation. Of the 26 species of non-heterocystous blue-green algae (Table 8) none are found in forest and only 3 in estates. They are characteristic of open areas such as exposed sub-soil, grassland and scrubland where the light-intensity is high. TABLE 8 Number of Occurrences of non-heterocystous blue-green algae ——- F A E G S . Est D Total | CHROOCOCCUS MINUTUS (Kiitz.) Nag. 2% 2 Z | a 6 C. TURGIDUS (Kiitz.) Nag. .. 1 3 1 5 ee ee ee C. SCHIZODERMATICUS West 1 | 1 CHROOCOCCUS sp. 3% 1 1 | 2 MICROCYSTIS AERUGINOSA | Kiitz. ee “fe 1 E 2 CHAMAESIPHON FUCUS | (Restaf.) Hangs. a 1 | | 1 OSCILLATORIA JASORVENSIS Vouk. ae a 2 11 3 1 17 296 Gardens’ Bulletin — X XVI (1973) TABLE 8 — continued. Number of Occurrences of non-heterocystous blue-green algae | | or ae E G S C | Est. , D | Total O. osscurRA Briihl. et Biswas 1 1 ‘a ah | | 1 7 | eee Se SS | | | O. ANNAEA Van Goor ae | 1 1 zm | | 4 Ipeut | reel os esd Sites cx at 220 O. CHALYBYA (Mertens) Gom. | | 1 3 | | ga vr or ah O. FREMYI De Toni 2 1 3 O. MINIATA (Zanard) Hauck 1 1 1 3 : a as oe i nn eM —_— es O. TEREBRIFORMIS Ag. | 1 1 | ) ene ere 05 | a O. CHLORINA Kiitz. ). 1 1 2 | i ees ee | ee Sad } | O. OKENI Ag. .. a 1 1 | | | 2 vr O. Limosa Ag. .. on ] 1 | eek: O. SUBTILLISSIMA Kitz. .. 1 | 1 O. GEITLERIANA Elenkin. .. 1 | | | 1 ideiaccunctisiriwe-satdedinaieiein waeeltealat ane SYNECHOCOCCUS AERUGINOSUS | Nag. re J Z | 1 | 7 ARTHROSPIRA TENUIS Stikenb. 1 | 1 | 2 ret ei | SPIRULINA LABYRTHIFORMIS | (Menegh.) Gom. | 1 | | 1 PHORMIDIUM TRUNICOLA Ghose | 1 | 1 PH. SUBINCRUSTATUM Fritsch and Rich 4: fh 1 1 LYNGBYA POLYSIPHONAE a a 1 | 1 Frémy a | y | 1 | L. BIRGEI Smith .. a 1 | TOTAL NO. OF SPECIES 0 7 15 10 | > Epiterranean soil algae in Singapore 297 Table 9 shows the occurrence of 16 heterocystous blue-green algae which are capable of nitrogen-fixation. They occur in just over a quarter of the soil samples. None are found in forest soil but at least one species of nitrogen-fixing algae has been found in all other types of habitat. Their occurrence is sporadic in these habitats and nearly three quarters of the samples showed no evidence of the presence of nitrogen-fixing algae. TABLE 9 The Occurrence of heterocystous blue-green algae ANABAENA FERTILISSIMA Rao ANABAENA SPIROIDES Kleb. .. ANABAENA FUELLEBORNII Schmidle ANABAENA ORYZAEA Fritsch ANABAENA ANOMALA Fritsch ANABAENA SP. NosTOC MICROSCOPICUM Carm. N. MUSCORUM Ag. N. | PUNCTIFORM (Kiitz.) Hariot ie id TOLYPOTHRIX PHYLLOPHILA West : SCYTONEMA HOFMANNI Ag. CYLINDROSPERMUM SPHAERICA Prasad : -_ AULOSIRA AERIGMATICA Frémy HAPALOSIPHON WELWITSCHII West bs NOSTOCOPSIS RADIANS Bharadn. WESTIELLOPSIS PROLIFICA Janet : TOTAL NO. OF SPECIES | F A | E G S c Est. D Total 1 1 1 3 1 1 2 | 1 1 1 1 1 1 1 1 1 7 ee 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 2 1 3 0 3 z 7 1 3 2 3 16 298 Gardens’ Bulletin — XXVI (1973) (c) Green algae. The occurrence of green algae (Chlorophyceae, Xanthophy- ceae, Euglenophyceae) is seen in Table 10. The highly constant species Desmococ- cus vulgaris, Palmella miniata and Hormidiun flaccidum are widely distributed, but most of the 31 species are very sporadically distributed and include some species as Gonium pectorale and Euglena elongata which are characteristic of freshwater habitats. TABLE 10 (Cholorphyceae, Xanthophyceae and Euglenophyceae). DESMOCOCCUS VULGARIS Brand. PALMELLA MINIATA Nag. HORMIDIUM FLACCIDUM (Kiitz.) Br. my TROCHISCIA ASPERA (Reinsch Hansg. a * MURIELLA TERRESTRIS Petersen EUGLENA ELONGATA Schewiakoff .. CHLAMYDOMONAS Sp. SCENEDOSMUS FLEXUSOS (Lemm.) Ahlstrom CYLINDROCYSTIS sp. BUMILLIERA EXILIS Kleb SPONGIOCHLORIS (Vischer) Starr SPONGIOSA MICROSPORA SP. GONIUM PECTORALE Miill .. OEDOCLADIUM Sp. EUASTRUM TORUM BINUCLEARIA TECTORUM (Kiitz.) Beger LEPTOSIROPSIS TORULOSA Jao DICTYOCHLORIS FRAGRANS Vischer oe ZYGOGONIUM Sp. TRENTEPOHLIN AUREA (L.) Mart. by ji STIGEOCLONIUM Sp. ANKISTRODESMUS FALCATUS (Corda) Ralfs ; eae es ae Re en Peete Je 8 ee: ia 278 Ldn dreabard 1 C20) fee | 4 | 1 1-1—4 1 a 3 3 1 404.22 Pon. ie nee ey es eee A a 1 . 1 | a ee 1 1 2 1 1 1 1 1 ated ee 1 Total Epiterranean soil algae in Singapore 299 TABLE 10 — continued (Cholorphyceae, Xanthophyceae and Euglenophyceae) i zing ida a fs a a Ned a D | Total CEDERCUREUTZIELLA SAVONEN- sis Vischer .. of a 1 ie big de: ae i 1 DACTYLOCOCCUS INFUSIONUM Nag. my i ‘ay 4 1 s mT oes ae = 1 MACROCHLORIS DISSECTA Korch. 7 fy AES > a Fe a 1 ie) i. 1 PENIUM sp. ? 7 ” << sts od “. 1 | 3 | 3 | 3 | 0 | 1 | 2 | 1 7 300 Gardens’ Bulletin— X XVI (1973) (e) Diversity of algal flora. Table 12 shows the occurrence of different kinds of algae in the different habitats. Tree communities tend to lack diversity in algal flora. In forest only 4 species are present, none of which is blue-green. Scrubland and disturbed land have 15 and 16 species respectively. Algal flora is best deve- loped in exposed sub-soil (29 species) and grassland (35 species). As mentioned above nitrogen fixing blue-green algae may occur in all kinds of habitat except forest. TABLE 12 Summary Table for number of species of different groups of algae = 2 Fy, | A’: |) Be ae Ss |) te | ee Total a a bogeeay Non-heterocystous blue-greens| 0 ‘4 15 10 9 7 2 | ot | 26 Heterocystous blue-greens .. 0 3 2 7 1 3 4 3 | 16 Green algae. te ap [i el Sa iy | | eee Diatoms { SPs i se Ss i Total no. of species = 5 23 2B) 35 15 18 | 16 | 10 | 80 Discussion Although 80 species of algae have been found in the soil of Singapore Island in this investigation, most of these show low constancy of occurrence in the Island as a whole or in an individual type of habitat. The phenomenon of a large diversity of species distributed with low frequency is familiar amongst tropical organisms and occurs amongst such diverse groups as lowland tropical forest angiosperms, beetles (Wallace, 1969) and tropical fish. When the natural vegetation is completely cleared and the top soil removed by man or erosion, an exposed sub-soil is devoid of any type of higher plant vegetation. This is subject to excessive leaching by torrential rain, strong isolation from the tropical sun and considerable temperature fluctuation day and night (Hill, 1966). Despite these severe conditions 29 species of soil algae were found some of which may act as first colonisers of the bare ground before the establishment of a higher plant flora. Non-heterocystous blue-green algae predominate in this habitat. When colonisation by higher plants commences, grassland becomes established. The soil surface becomes protected from excessive leaching and there is an increase in available ions for plant growth (Table 2.) Light intensity is still high but there is less fluctuation in temperature day and night. 35 species of soil algae were found including 7 species of heterocystous blue-green and 15 species of green algae. Grassland progressively develops into scrubland (belukar) with colonisation by woody species such as Adinandra dumosa Jack and Melastoma malabathricum Linn. These species increase the shading of the ground layer. The light intensity Epiterranean soil algae in Singapore 301 decreases and there is an increase in available nitrate. The algal flora loses some of its diversity and only | species of heterocystous blue-green algae was found. In the undisturbed forest the organic matter content is high but most cations and anions remain low, since the ionic reserve in tropical forest is directly absorbed by the shallow feeding roots of the trees and does not persist in the soil (Jackson, 1968). The light intensity is very low being only about 0.1% of the illumination outside the forest (Johnson, 1970). No blue-green algae are found here and there are only four species of green algae and one diatom. Cultivated soil in Singapore has been subjected to various manures and organic fertilisers and usually has a relatively high nitrate level. The algal flora is limited but heterocystous blue-green algae can occur. Estates are also subject to disturbance by man. Since these are predominately tree crops (e.g. rubber, banana, coconut, rambutans, durian, citrus etc.), the light intensity is usually lower than in cultivated soil; but, since such estates are quite open, about 10% light penetrates. The algal flora is limited but less so than in undisturbed forest. Disturbed soil has been subject to all kinds of dumping and building operations by man, including industrial pollution and general devastation. The occurrence of such areas near a city of over two million population is to be expected. Chemical analysis of the soil gives very erratic results often with high figures for one or more ions. Ten species of algae were found in this kind of habitat. Summary The surface soil was collected in 100 localities in the rural area of Singapore. The samples were cultured by liquid and plate culture to determine the algae occurring, and the soil was subjected to physical and chemical analysis. The surface soil consisted predominately of clay and clay-loam, with some sandy loam and loamy sand, other types of soil being rare. The water-holding capacity of dry soil is closely correlated with organic content being high in forest and alluvial soils but low in all other types of soil. Nitrate level was normally at moderate to moderately low levels. but the values for phosphate, sulphate and potassium are critically low in most soils. Most soils are acid, forest soils having an average pH of 3.83 with pH somewhat higher in other types of soil. Caroonate was detected in only 17% of the samples. Although the soils were somewhat uniform in hue with 60% soils in the 1OYR range, they were very variable in colour. 80 species of soil algae were found in the soil. Tree communities lack diversity in algal flora, which was best developed in exposed subsoil and grassland. Most species show low constancy of occurrence in the Island as a whole or in individual habitats. 302 Gardens’ Bulletin — XXVI (1973) Literature cited Becking, J. H. (1961) Studies on the nitrogen-fixing bacterium of the genus Beijerinckia If Mineral nutrition and resistance to high levels of certain elements in relation to soil type. Plant and Soil 14: 297-322. Bristol, B.M. (1919) On a Malayan form of Chlorococcum humicola (Naeg.) Rabenh. Jour. Linn. Soc., Bot., 44: 473-482. Edwards Laboratory, Norwalk, Ohio: Soil Testing, in Technical Bulletin No. 162, Michigan Agric. Exp. Station. Fritsch, F.E. (1907) A general consideration of the sub-serial and freshwater algae of Ceylon. Proc. Roy. Soc., Lond. B 79: 197-254. Ghose, S.L. (1923) A systematic and ecological account of a collection of blue- green algae from Lahore and Simla. Jour. Linn. Soc., Bot., Lond. 46: 333-346. Gonzales, Ella A., and Valavigi, V.S. (1960) Algae in the rhizosphere of some crop plants. Proc. Symp. Algol., Indian Council Agric. Research 333-342. Hill, R.D. (1966) Microclimate observation at Bukit Timah Forest, Singapore. Malayan Forrester 29: 78-89. Jackson, H.L. (1958) Soil Chemical Analysis. Constable and Co., Ltd., London. Jackson, J.C. (1968) Planters and Speculators. University of Malaya Press. Kuala Lumpur. Johnson, Anne (1962) Precursory studies on the epiterranean soil algae of Singapore and Malaya. Gdns. Bull., Singapore, 19: 379-384. ——. (1966) B-values and the phytoplankton community. Verh. Internat. Verein. Limmol. 16: 619-626. . (1969) Blue-green algae of Malaysian rice-fields. Jour. Sing. Nat. Acad. Sci. 1: 30-36. —. (1970) Determinative factors in the distribution of lowland vegeta- tion in Singapore and Malaysia. Nanyang Univ. J. 4: 165-174. Leamy, M.L. and Panton, W.P. (1966) Soil Survey Manual for Malayan Conditions. Division of Agric. Bull No. 19, Ministry of Agric. and Co-operatives, Kuala Lumpur, Malaysia. | Munsell Soil Colour Charts (1971) Munsell Colour Company Inc., Baltimore, U.S.A. Panton, W.P. (1955) Soil survey reports No. 2, A part of west Singapore Island. Malayan Agric. J. 38: 1-26. Prasad, S. (1949) Nitrogen recuperation by blue green algae in soils of Bihar and their growth on different types of soil. J. Proc. Inst. Chemists (India) 21: 135-140. Singh, R.N. (1940-41) The relation of algae to the nitrogen economy of the soil. Chron. Bot. 6: 202-203. Sulaiman, M. (1944) Effect of Algal growth of activity of Azotobacter in rice soils. Ind. J. Agric. Sc. 14: 277-283. Tiffany, L.H. (1951) in G.M. Smith, Manual of Phycology Ch. 15: 303-306. Wallace, A.R. (1869) The Malay Archipelago, Macmillan and Co., London. Pollination in Dioecious Figs Pollination of Ficus fistulosa by Ceratosolen hewitti By J. GALIL Department of Botany, Tel Aviv University, Israel ABSTRACT Pollination in the dioecious fig Ficus fistulosa was studied in Singapore. The pollinator, Ceratosolen hewitti (Agaonidae), possesses closed thoracic pockets and coxal combs. Pocket filling with pollen from detached anthers previously cut by the male wasps occurs in the almost ripe male figs. Pollen is lifted from the anther to the underside of the thorax by means of the arolia of the fore legs and then shoveled backwards into the pockets by the combs. In the male figs, pollination of the stigmata occurs at the end of oviposition prior to the withdrawal of the ovipositor. Pollen is taken directly from the pockets by the fore- legs and is carried downwards onto the stigmata. The behaviour of the female wasps in the female syconia is very similar except that the oviposition act does not culminate with actual egg-laying. The significance of pollination in short-styled, gall-producing flowers and of Ovipositor insertion into the styles of long-styled, seed-producing flowers is discussed. Introduction It is known that the symbiotic wasps (Agaonidae, Chalcidoldea) which serve as pollen vectors in the genus Ficus carry out in the figs two types of activities namely, pollination of the long-styled flowers and oviposition into the pistils of the short-styled ones. Hence, the former produce seeds whereas the ovaries of the latter develop into galls and give rise to the next generation of wasps. Both activities are indispensable for the existence of fig symbiosis. There is no evidence of the presence of apomixis in Ficus so that pollination, too, appears to be essential for seed production. The question arises as to the occurrence of pollination in the short-styled flowers and oviposition into the long-styled ones. In the common fig (F. carica L.)., the most extensively studied Ficus sp. (Grandi, 1929), the situation is comparatively simple. Here, pollen is passively carried on the body of the female wasp Blastophaga psenes L. from one fig to another so that pollination of both long- and short-styled flowers takes place indiscriminately; on the other hand, the long styles, which so markedly differ in structure from the short styles, do not appear to be suitable for oviposition. The situation is not always as simple in other Ficus spp. There is considerable variation in structure and behaviour of the pollinators of different species of Ficus, in spite of an overall resemblance. Unique organs used by various spp. of wasps as containers for transporting pollen from male figs (phase D*) to young receptive female figs (phase B*) were recently discovered (Ramirez, 1969; Galil and * According to Galil and Eisikowitch (1968) the developmental phases of figs were named as follows: Phase A (Pre-female): Young syconium prior to the opening of the ostiole. Phase B (Female): Ostiolar scales loosen, female flowers ripen, sycophilous wasps penetrate into the syconium and oviposit into the ovaries. Phase C (Interfloral): Wasp larvae and fig embryos develop within their respective ovaries. Ovaries occupied by the larvae are transformed into galls. Phase D (Male): Male flowers mature, wasps reach the imago stage, fertilized female wasps leave syconia via channels bored by the males. Phase E (Post-floral): Both syconia and the seeds inside them ripen. 303 304 Gardens’ Bulletin — X XVI (1973) Eisikowitch 1969; Galil and Snitzer-Pasternak, 1970; Galil et al., 1974). In these figs, the pollination process consists of “deliberate” loading of the containers and, upon entering the young figs, extracting the pollen and pollinating the female flowers. Actual pollination takes place at the end of the oviposition act, before the wasp withdraws her ovipositor from the pistil. Hitherto, pollination by wasps with specialized pollen containers was exclusively studied in monoecious figs that contain both long- and short-styled female flowers (the so-called seed- and gall- flowers, respectively) in the same syconium. Because of the dense packing of the fiowers within such figs, it is very difficult to observe the behaviour of the wasps in regard to the different flower types. Nevertheless, there is some information available. Johri and Konar (1956) discovered fig embryos up to the octant stage in the short-styled ovaries of F. religiosa L. This is clear evidence of fertile pollination of the short styled flowers. “Seed figs” in which both long- and short- styled flowers bear normal seeds that are occasionally found in various species (Galil and Eisikowitch, 1971), provide further evidence of normal pollination in both types of flowers. On the other hand, the blackening of the upper part of the style in long-styled flowers of F. religiosa, observed by the same authors, indicates that the wasps insert their ovipositors into such pistils as well. Direct observations of the behaviour of the wasps in long- and short-styled flowers in regard to pollination and oviposition were hitherto carried out only in the monoecious figs of F. sycomorus L. (Galil and Eisikowitch, 19769). However. since the two flower types in that species are very similar and cannot be distinguished from the above without destroying the fig, only a few sporadic observations could be made. Dioecious fig species, in which long- and short-styled female flowers develop in different syconia, lend themselves more easily to detailed observations on wasp behaviour with regard to the two flower types. For such observation Ficus spp. whose pollinators are equipped with specialized pollen containers which are actively loaded and emptied must be chosen. The dioecious figs of F. fistulosa Reinw. ex Bl. (Fig. 5) which belong to the Sycocarpus section of subgenus Ficus and are pollinated by Ceratosolen hewitti Waterston, possessing thoracic pockets for pollen transport, prove suitable for the present study. Materials and Methods The study was carried out in June-July, 1971 in Singapore. The chief object, F. fistulosa, is a small tree, 3 to 4 m. high, that grows abundantly along road edges in the suburbs of the town and on the borders of the jungle. Generally, the study of pollination in dioecious figs is more complicated than in monoecious species since it is necessary to find wasp-releasing D-phase male figs and receptive B-phase figs of both sexes simultaneously in the same vicinity. The month of June proved to be very suitable since it is a period of overlapping of the two main crops of figs in F. fistulosa. Hence, all of the required develop- mental phases of the figs were available (for Hong Kong, see also Hill, 1967). During the study, no ripening female figs were found, but these were not indis- pensable for carrying out the necessary observations. The cauliflorous figs of F. fistulosa develop in great numbers, provide abundant and easily accessible material. The timing of observations was adjusted to the activities of the wasps in the male and female phases of the fig. Hence, the Pollination of Ficus fistulosa 305: observations on pocket filling were carried out in the early morning, between 3 and 5 a.m. For observing the behaviour of the wasps in young figs during oviposition and pollination, B-phase figs, male and female separately, were enclosed in plastic vials covered with insect-proof netting together with wasp-releasing D-phase male figs. The emerging wasps which penetrated the receptive B-phase figs provided good material for observation of wasp behaviour during oviposition and pollination. Observations Syconia and pollinators The figs develop on small lateral shoots on the trunks and branches (Fig. 6). The receptive (phase B) figs of both sexes are pear-shaped with a short neck and are borne on distinct peduncles that are up to 25 mm long (Fig. 7). The male figs. are somewhat larger than female figs, 14 x 16 mm as compared to 11 x 13 mm. The ostiole is depressed and is sealed by imbricate scales. The structural features of the fig are especially prominent when bisected figs are viewed from the inner side (Figs. 8, 9). The different components have different colours: the ostiolar scales are orange and the pale stigmata stand out on the background of crimson ovaries. The internal cavity of the fig is wide and distinct. The male figs contain a large number of short-styled female flowers (“gall flowers’) covering almost the entire inner surface of the fig and 30 to 35 ostiolar male flowers arranged in one to two rows (Fig. 8). The male flowers are relatively prominent and easily discernable, but are still closed and unripe. The “gall flowers” are almost hidden within the fused perianth lobes (Fig. 1, IJ). The short (about 0.3 mm) and relatively thick style and the funnel-shaped stigma at its end project out of the perianth. The broad and dense stigmata form a continuous surface around the fig cavity but do not coalesce into a single “synstigma”. The long-styled female flowers (‘‘seed flowers”) which cover the entire inner surface of the female figs are quite different from the short-styled ones. The perianth is much reduced thus forming a narrow ring at the base of the ovary (Fig. 1, IIL). Consequently, the pistils are naked and visible for their entire length in bisected figs. The ovary is smaller and the style is narrower and much longer (about 1 mm) than their counterparts in the short-styled flowers. The stigma is cylindrical, thick and long (0.16 x 0.3 mm). At the wasp-releasing stage (phase D), the male figs are almost globose (Fig. 10), somewhat broader than high (20 x 25 mm). The holes in the empty gall-ovaries from which the wasps have already emerged have rounded lids which are shifted to the side (Fig. 12). Upon ripening of the male flowers, the filament of the single stamen elongates so that the anther is pushed through the top of the fused perianth (Fig. 1, I). At this stage, the males of the pollinating wasps approach the male flowers and cut off some of the anthers causing them to be scattered within the cavity of the fig. Thus, the emerging female wasps encounter cut anthers in addition to the intact stamens remaining in situ around the ostiole (Fig. 11). At the end of phase D, a large hole is cut through the scales in the centre of the ostiole by the male wasps (Fig. 12). The pollen loaded females emerge from the fig through this hole. 306 Gardens’ Bulletin — XXVI (1973) After the liberation of the wasps, the fig grows rapidly, and at ripening (Phase E) it reaches about 30 mm in diameter. It becomes yellowish and attains a characteristic smell which is typical for bat-dispersed fruit. As noted above, no ripe female figs of F. fistulosa were found throughout the period of the present study. Hill (1967) states that the ripe female figs are somewhat smaller than the male figs, but are more succulent and their yellow colour is more pronounced. The pollinator, Ceratosolen hewitti Waterston (Wiebes, 1963, 1966; Grandi, 1963) is a small agaonid wasp about 1.5 mm long (Fig. 2). Only those characteristics of the female wasp that pertain to pollination will be described below. Of the two types of specific pollen containers hitherto described in pollinators of Ficus, namely “coxal corbiculae” and “thoracic pockets” (Ramirez, 1969; Galil et al., 1974), the female of C. hewitti possesses the latter only. In whole mounts of the ventrum of the body, the pockets appear on both sides of the mesothorax as triangular depressions that are entirely covered by continuous flaps (Fig. 3). Generally, the structure of the pockets is quite similar to that of the corresponding organs described in C. arabicus Mayr, the pollinator of F. sycomorus L. in East Africa (Galil and Eisikowitch, 1969). It appears that the pockets open and close along their inner suture facing the median axis of the body through changing in the position of the covering flaps. Coxal corbiculae such as described by Ramirez (1969) and Galil et al. (1974) were not found on the fore coxae of C. hewitti. There are no depressions along the coxae: only the comb, namely the row of stiff bristles which usually fringe the depression on its inner side, is found here. In C. hewitti, the comb is a distinct organ formed from a straight row of about 18.38y-long bristles (Fig. 4). The length of the ovipositor of the pollinating wasp is of decisive importance in fig symbiosis. As seen from the side, the ovipositor of C. hewitti scarcely projects beyond the apex of the body. In fact, the ovipositor is not short but arises at some distance forwards, on the under side of the abdomen. Its real length may be seen during egg-laying when its shaft is released from the protecting valves and is directed towards the stigmata. Pocket loading Detailed observations of the filling of pockets of C. hewitti in D-phase male figs of F. fistulosa are not easy. Figs with ostiolar male flowers are usually suitable for such observations. However, in F. fistulosa the situation becomes more complicated because of the early emergence of the female wasps from their gals and the filling of the pockets prior to the tunnelling of the syconial wall by the male-wasps. For successful observation of pocket loading, the figs must be cut and studied during a comparatively short lapse of time when the anthers are still attached but sufficiently ripe, and the fertilized females have not yet left their galls. When the figs are cut at the appropriate time, the female wasps become active, emerge from the galls, approach the male flowers and start filling their pockets from the anthers which protrude out of the floral perianth. Lifting of the pollen from the anthers to the side of the thorax is accomplished by rapid alternating movements of the forelegs. Similar movements were observed in the pollinators of monoecious figs previously studied. Clumps of pollen are Pollination of Ficus fistulosa 307 Fig. 1. Ficus fistulosa, flowers: I— male flower, Il — short-styled female flower (°); III — long-styled female flower °. Fig. 2 Ceratosolen hewitti (° and ¢). Fig. 3. Mesothorax of Ceratosolen hewitti (from below) showing pollen pockets. Fig. 4. Fore leg of Ceratosolen hewitti, with comb. Pocket 308 Gardens’ Bulletin— XXVI (1973) placed between the body wall and the fore coxae which move up and down when the pollen is lifted by the bending legs. Shovelling movements of the fore coxae and combs by which the pollen is brushed backwards from the underside of the thorax to the thoracic pockets occur after each sequence of pollen liftings. The coxal comb is clearly a unique organ and even in the absence of the coxal depression (‘‘corbicula’’), it has a particular role in pocket filling. The shovelling movements are rapid but can be clearly seen. The role of the combs in the brushing of the pollen is evident from the pollen grains which are found on the bristles. When the quantity of pollen in the anther does not suffice for the filling of the pockets, the wasp moves to a second anther and begins a new series of liftings and shovelling movements. Pocket emptying: pollination in male figs The observations on the behaviour of the female wasps during oviposition and pollination were especially successful in the early morning when B-phase receptive figs were enclosed in plastic vials together with wasp-releasing D-phase male figs. As in the monoecious Ficus spp. previously studied (Galil and Eisiko- witch, 1969; Galil et al, 1974), the impregnated and pollen-carrying female ‘wasp starts to oviposit and to pollinate the stigmata immediately after its entrance into the receptive fig. In C. hewitti, as usual in the agaonid wasps, the ovipositor shaft is released from the protecting valves at the beginning of oviposition act and is supported and guided to the stigmata by the sternites that fold lengthwise to form a triangular, gutter-shaped, body. The total length of the ovipositor is about 0.8 mm., i.e. its length is intermediate between that of the styles in long- and short-styled flowers. After several probing movements for an appropriate oviposition site, the ovipositor sinks into one of the funnel-shaped stigmata of the short-styled flowers. It remains in the style only for 15 to 18 seconds which is a very short time compared to the duration of the act in pollinators of other figs previously studied, i.e. about 60 seconds in C. arabicus, and 3 to 3.5 minutes in Blastophaga estherae Grnd., the pollinator of F. costaricana (Liebm.) Miq. (Galil et al., 1974) Almost immediately after the insertion of the ovipositor into the pistil of the fiower, the female wasp begins to perform typical pollination movements. The forelegs fold upwards upon themselves one after the other intermittantly, thereby bringing the arolia into contact with the pollen pockets on the under side of the mesothorax. The pollen grains are taken up by the legs directly from the pockets; in any case, no shovelling of pollen from the pockets by means of the combs of the fore coxae were observed. The pollen-carrying legs open downwards one ofter the other and touch the stigmata beneath. It is apparent that normal pollination of the short-styled flowers takes place systematically at the end of each oviposition act. During oviposition and pollination, the head of the female wasp remains motionless and no biting of the stigmata takes place, as observed in C. arabicus (Galil and Eisikowitch, 1969). Pocket emptying: pollination in female figs In spite of the distinct differences in the structure of the pistils in the female flowers of the male and female figs, the behaviour of the wasps was almost alike. As in the male figs, the female wasps moved on the surface of the stigmata, inserted the ovipositors into them one by one and performed the characteristic pollination movements at the end of each insertion. The retention of the ovipositor Plate 1. Figs. 5-12 Ficus fistulosa Fig. 5. Leafy branches. Fig. 6. Fruiting stem. Fig. 7. Short branch with male figs at Phase B. x 0.7. Fig. 8. Cross section through male fig at phase B. (°) short-styled female flowers. ¢ —vunripe male flowers . x 3.7. Fig. 9. Cross section through female fig at phase B. ° — long-styled female flowers. Fig. 10. Male figs at phase D. x 0.6. Fig. 11. Longitudinal section through male fig after release of wasps. $ — attached anthers; A — detached anthers; x 2.3. Fig. 12. Cross section through male a ey RE De gat) A ee) Ee eee ae > ee ar ey 2, a. me. | Pollination of Ficus fistulosa 309 within the style was shorter than in the short-styled flowers and lasted barely 8 to 10 seconds. Here also pollen is transferred directly from the thoracic pockets onto the stigmata of the long-styled female flowers by the arolia of the fore legs. During these observations, several cases of unusual behaviour of the wasps in the female figs were encountered. Exhausted female wasps, which have already completed their usual activities, continued to move their fore legs in vain efforts to extract pollen from the pockets and to deposit it on the stigmata. In order to observe this phenomenon more closely, pollen loaded female wasps were taped in a supine position on a glass slide and observed through the dissecting microscope immediately after they emerged from the D-phase male figs. Most of the wasps made vain attempts to get off the glass; however, in a few cases, the wasps started to perform the characteristic pollination movements during which they folded the fore legs onto the thorax and inserted the arolia into the widely gaping pollen pockets. Pollen grains could be clearly seen on the arolia. No such movements have hitherto been seen in the pollinators of the monoecious figs in which pollination has been studied. Discussion In spite of numerous structural differences between the syconia of dioecious F. fistulosa and monoecious F. sycomorus (Galil and Eisikowitch, 1969) which belong to different subgenera of Ficus, namely subgenus Ficus and subgenus Sycomorus respectively, the two have several biological features in common. Of particular significance is the relatively small number of male flowers which reflects the efficiency of the pollen vectors in pollen transfer. In both Ficus taxa, the pollinating wasps are species of Ceratosolen possessing closed thoracic pollen pockets of similar structure and coxal combs without corbiculae, and behave very similarly in relation to the figs. In both species, the male wasps cut the stamens and the females emerge from their galls, fill their pockets with pollen from the detached anthers and gather in the fig cavity before the syconial wall is tunnelled by the males. Also the behaviour of the female wasps during pocket filling and emptying, i.e. pollination, is similar. Such likenesses indicate that physiological conditions within the figs are probably also similar in both cases (Galil et al., 1973). In F. fistulosa, the behaviour of the female wasps in the short- and long-styled female flowers (in different figs!) is almost identical in spite of the structural differences between them. In the two types of flowers, both insertion of the ovipositor into the style and deposition of pollen on the stigmas take place. Essentially, it is the same process although eggs are not likely to be laid within the long-styled pistils. In F. sycomorus, serial microtome sections through long- styled pistils in pollinated figs failed to reveal any remnants of Ceratosolen ova. The pistils of the long-styled flowers of F. fistulosa have not been similarly investigated but there are some indirect indications that here too, no real oviposition takes place. The relatively much shorter time that the ovipositor remains in the long styles is significant. It appears that for effective oviposition, the stimulus which the flower ovary exerts on the tip of the ovipositor is indispensable. Consequently, when the style is too long and the ovary is beyond the reach of the ovipositor, there is no egg-laying. 310 Gardens’ Bulletin — X XVI (1973) The behaviour of the wasp in the male and female figs raises two questions which may be of fundamental importance for the understanding of fig biology: 1. What is the biological purpose of pollination of the short-styled pistils which are in any case destined to develop into wasp-galls? and 2. Is there any significance to the futile insertions of the ovipositor into the long-styled pistils? In F. religiosa, where the first problem was studied more thoroughly, the selective advantage of pollination of the gall-producing short-styled flowers is evident. In unpollinated short-styled flowers, the biological equilibrium is destroyed because of paucity of food, resulting from the failure of normal endosperm development, causes high mortality in female wasp larvae (Galil and Eisikowitch, 1971). It is likely that in F. fistulosa the situation is similar. As yet, there is no satisfactory answer to the significance of the ovipositor insertion (‘‘stinging”’) into the pistils of the long-styled flowers. Perhaps the changes caused by wounding of the tissue has some impact on pollen germination or pollen tube growth. It should be borne in mind that attraction of the wasps to the female figs and the activities of the wasps within the female figs are of no apparent immediate advantage to the insect. Identical chemo-attractants, produced by both types of figs, could cause the wasps to mistake the female figs for the male figs, which provide the necessary brooding place for the new generation. Thus, it is also possible that insertion of the ovipositor into the long-styled pistils has nothing to do with the biological processes in the female figs but represents a link in an automatic, instinctive chain of activities, orignally concerned with short-styled flowers in male figs. It should be noted that a strict coordination between the pollination movements and oviposition in both male and female figs is not always sustained. The numerous pollen-extraction movements observed in dying female wasps and the repeated pollen withdrawals from the pockets observed in taped wasps hint at a possible disruption of such coordination. Consequently, normai intermittent pollination which is regularly interrupted by search for new sites of oviposition and insertions of the ovipositor into the stigmata may be replaced by a continuous act, consisting of mere emptying of the pockets. Acknowledgements The author acknowledges with many thanks Professor E.J.H. Corner, Botany School, Cambridge, and Professor J.T. Wiebes, Rijksuniversiteit, Leiden, for their helpful information on the fig spp. and the fig wasps of Singapore. The author is also indebted to Professor A.N. Rao, Head, Professor H. Keng and Dr. P.N. Avadhani of the Department of Botany, University of Singapore and to Mr. A.G. Alphonso, Director, and Dr. Chang Kiaw Lan of the Botanic Garden of Singapore for their assistance and hospitality. Thanks are also due to S. Bullowa and Dr. A. Horovitz for reviewing the manuscript, to S. Schaeffer for the illustrations, and to J. Cohen and A. Shub for the photographic work. Literature Cited Corner, E.J.H. 1958. An introduction to distribution of Ficus. Reinwardtia 4: 15-45. . 1965. Check-list of Ficus in Asia and Australia with keys to identification. Gard. Bull. Sing. 21: 1-186. Pollination of Ficus fistulosa 311 Galil, J. & D. Eisikowitch, 1968. On the pollination ecology of Ficus sycomorus in East Africa. Ecology 49: 259-269. & . 1969. Further studies on the pollination ecology of Ficus sycomorus. Tijdschr. Entomol. 112: 1-13. —— & . 1971. Studies on mutualistic symbiosis between syconia and sycophilous wasps in monoecious figs. New Phytol. 70: 773-787. , M. Zeroni & D. Bar Shalom, 1973. Carbon dioxide and ethylene effects on the coordination between the pollinator Blastophaga quadraticeps and the syconium in Ficus religiosa. New Phytol. 72 (5). , W.B. Ramirez, & D. Eisikowitch, 1974 Pollination of Ficus costaricana and F. hemsleyana by Blastophaga estherae and Blastophaga_ tonduzi (Agaonidae) in Costa Rica. Tijdschr. Ent. In Press. , & Y. Snitzer-Pasternak. 1970. Pollination in Ficus religiosa L. as connected with the structure and mode of action of the pollen pockets of Blastophaga quadraticeps Mayr. New Phytol. 69: 775-784. Grandi, G. 1929. Studio morfologico e biologico dell Blastophaga psenes L. Boll. Lab. Ent. Bologna. 2: 1-147. . 1963. Catalogo regionato degli agaonidi de mondo. Boll. Ins. Ent. Univ. Bologna 26: 319-373. Hill, D.S. 1967. Figs (Ficus spp.) of Hong Kong. Hong Kong Univ. Press, Hong Kong. Johri, B.M. & R.N. Konar, 1956. The floral morphology and embryology of Ficus religiosa Linn. Phytomorph. 6: 97-111. Ramirez, W.B. 1969. Fig wasps. Mechanisms of pollen transfer. Science 163: 580-81. — . 1970. Taxonomical and biological studies of neotropical fig — wasps (Hymenoptera, Agaonidae) Univ. Kansas Sci. Bull. 49: 1-44. Wiebes, J.T. 1963. Taxonomy and host preferences of Indo-Australian fig — wasps of the genus Ceratosolen (Agaonidae). Tijdschr. Ent. 106: 1-112. ——— . 1966. Provisional host catalogue of fig wasps (Hymenoptera, Chalcido idea). Zool. Verh. 83: 1-44. ; rT sy oo Pre . | - +¢ Gar ony ay ; arrdenanee wii bilo ywoloon qoieatilogr | 7a em fTumdarae ntai ITAportanc Dac ooltinasi . } rick A he by ‘to ‘auloas sovenion, ee in yt St t it a brs sinboy2 aSswied sionals oteitapan ia ETAT TS POR Nona wet wae er qi" I) ge et . De f eres Ra sialydse bas. aeate ‘modue bay] od}. Dae a " S udene's apianillog ot, ag (0) SY lord “* Subaor ype ll fay por gD ne A aed pul in, . ee 4 err: ara ri" epee ata OTM ORR P Picadas ‘ *tg6q"" nlis4 dif To Hose % Shorts ‘wrurouTheol tama aRLRNR eee ae Sony WEVA TREE 20 | waeos to mnta Alo at ee 0 " ngnaasneshBl, ithe cnigoloid oe cepa ou d . + Ata : u T) ove i\ MIGET ts viet) 208 al dof ‘obrom ‘ab tbinonaé ifgsb ie ira ad id : , t ; : iWiLies, D enoH ast wild anol an0H yr" ynoH io (.qq@ cal i eRe as | re tor 4 logan ba a terol oar, ‘deh 504. a ALT-T@. 22 otlengmengasl rol some: tolenet? qolldq: t . omni frst 39 7M. “aqeew on TR BE MLD). FOL ow ait oft thi may be replaced bw i> Ipdigotiosa Ls eaibyie | bs piaoleid: bos {poirronoxe , Ab :@b Ile@ .io@ 2oznmaad vial (sebinosgA .é nN ;c2aW — ae nsileveuA-obnl 0 «soneagtoig.dand. ans umonoxeT £2 Sti-f. 2901 nF ‘woebRT, (osbinos aap) uslorom : ah wiese. 4) <0 faints DB valgoa aca eae ph 10 unoleleg ; decrt Innciaivent ate fort on the tig ig pnp ase o it | TOrCRSO! 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