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Cie ie GARDENS’ BULLETIN SINGAPORE
Index to Vol. 44 (Part 1) June 1992 Vol. 44 (Part 2) Dec 1992
Published by National Parks Board Singapore Botanic Gardens Cluny Road Singapore 1025
CONTENTS
Volume 44
PART 1 — Ist June 1992
TAN, H.T.W., CHUA, K.S. and TURNER, IM: tie papiospern Plota Of Singapore Project ......<......-.cccdaseceseeeesscescenssesscoeees 1-2
CORLETT, Richard T.:: The Angiosperm Flora of Singapore 1. Introduction ...............cccceeeeeeees 3-21
KIEW, R.: Five New Species of Didymocarpus (Gesneriaceae) from RR ra sec cudh ck wngcet ae iRE eRe dhe pho sense noaden coven ace 23-42
mORNER, E.J.H.: | Notes on the Development of the Fruit-bodies of Four Malayan Species of Amanita (Basidiomycetes) ...........::cccseeseeeees 43-45
AZIZ, Bidin and RAZALI, Jaman: Notes on the Rare Fren, Pteris holttumii C. CHM. ...................00eccecccceeeeeeeee 47-50
TURNER, I.M., TAN, H.T.W. and CHUA, K:S:.: OE SS BOT Tes id So ES 0 ot eee 51-71
PART 2 — Ist Dec 1992
WAH, T.T., WEE, Y.C. and PHANG, S.M.: Diatoms from Marine Environments of Peninsular Malaysia Bi eer k as Sopa idl ss eth e ca reasons ovcOe nani instn ons cnynedves seseeye 73-125
TAN, H.T.W., IBRAHIM, Ali bin and CHUA, K:S.: Piasmotis to be PlOTa OF SINGaPOTe, T ...n..s..0cn.20cs.censsessarnecscrecsersaseneseess 127-133
OGINUMA, Kazuo, LUM, Shawn K.Y., LEE, Y.H. and TOBE, Hiroshi: Karyomorphology of Some Myrtaceae from SingapoTe .............:006: 135-139
Basionyms and synonyms appear in italics. Page numbers in italics indicate the presencve of illustrations.
Abrus precatorius 59 Abutilon indicum 68 Acacia auriculiformis 59 Acacia mangium 59 Acalypha indica 67 Acanthaceae 54
Acanthus ebracteatus 54 Acanthus volubilis 54 Acaules 38
Aclisia secundiflora 66 Acrostichum 47 Acrostichum aureum 47,54 Acrostichum speciosum 54 Actinodaphne macrophylla 68 Adenostemma lavenia 66 Adiantum flabellulatum 65 Adiantum latifolium 52,54 Adinandra dumosa 16,64 Aegiceras corniculatum 69 Agavaceae 54,66 Ageratum conyzoides 56 Aglaonema simplex 66 Aizoaceae 54
Albizia retusa 68 Allamanda cathartica 55 Allophyllus cobbe 63 Alocasia macrorrhizos 55 Alstonia angustiloba 55 Alysicarpus vaginalis 59,68 Amanita 43
Amanita princeps 43,44 Amanita sp. aff. A.fritillaria 44 Amanita virginea 43,44,45 Amaranthaceae 55,66 Amaranthus lividus 66 Amaryllidaceae 66 Ampelocissus elegans 65 Anacardiaceae 55 Anacardium occidentale 55 Ananas comosus 56 Andira inermis 59 Andrographis paniculata 54 Annonaceae 15,55,66 Antidesma velutinosum 57,67
INDEX
Volume 44 (Part 1)
Apocynaceae 66
Araceae 55,66
Araliaceae 55,66 Archidendron clypearia 59 Ardisia crenata 61
Ardisia elliptica 61
Ardisia singaporensis 53,69 Ardisia villosa 69
Arenga pinnata 62 Arthrophyllum diversifolium 55 Artocarpus dadah 68 Artocarpus heterophyllum 61 Artocarpus integer 61 Asclepiadaceae 55,66 Asplenium macrophyllum 65 Asplenium nidus 54 Asystasia nemorum 54 Avicennia alba 55
Avicennia officinalis 55,66 Avicennia rumphiana 55 Avicenniaceae 55,66 Axonopus compressus 17,58
Baccaurea motleyana 57 Bambusa glaucescens 58 Basidiomycetes 43 Bidens pilosa 56 Bignoniaceae 55,66 Blechnum orientale 54 Blumea balsamifera 56 Blumea riparia 66 Boeopsis 25,32,34,38,39,40 Bombacaceae 55 Boraginaceae 55
Borreria alata 63
Borreria articularis 63,70 Borreria laevicaulis 63,70 Borreria setidens 63 Bouea macrophylla 55 Bougainvillea x buttiana 17 Breynia coronata 57 Bridelia stipularis 57 Bromeliaceae 56 Bromheadia finlaysoniana 62
141
142
Brucea javanica 64 Bruguiera cylindrica 63,70 Bruguiera gymnorrhiza 14,63 Buchanania arborescens 55 Bulbophyllum medusae 69 Bulbostylis barbata 57
Caesalpinia bonduc 14 Caesalpinia crista 60 Calophyllum inophyllum 59,67 Calophyllum pulcherrimum 59 Calophyllum spp. 16 Calopogonium mucunoides 60 Calotropis procera 66 Campanulati 38,41 Campnosperma auriculatum 11,55 Canavalia cathartica 60,68 Canna indica 56 Cannaceae 56 Capparaceae 56 Cardiospermum halicacabum 70 Carica papaya 56 Caryota mitis 62,69 Cassia alata 60 Cassia lechenaultiana 60 Cassia mimosoides 60 Cassytha filiformis 59 Casuarina equisetifolia 56 Casuarinaceae 56 Catharanthus roseus 55 Celastraceae 66 Celosia argentea 55 Centella asiatica 65 Centotheca lappacea 58 Centrosema pubescens 60 Ceratopteris thalictroides 54 Cerbera odollam 17,55 Ceriops tagal 63 Champereia manillana 62 Cheilanthes tenuifolia 65 Chirita 38 Chisocheton erythrocarpus 68 Chisocheton macrophyllus 53,68 Chloris barbata 58,67 Chrysopogon aciculatus 58 Cinnamomum iners 59 Cissus hastata 65 Claoxylon indicum 57 Claoxylon longifolium 53,67 Claoxylon longifolium var. brachystachys 53 ‘ Clausena excavata 70 Cleome rutidosperma 56
Gard. Bull. Sing. 44 (1992)
Clerodendrum inerme 65,70 Clerodendrum laevifolium 65 Clerodendrum paniculatum 65 Clerodendrum philippinum 65 Clerodendrum villosum 65 Clidemia hirta 61
Clitoria laurifolia 60
Cnestis palala 56
Cocos nucifera 62
Codiaeum variegatum 57 Codonoboea 23,25,32
Coix lacryma-jobi 58 Colocasia esculenta 55 Colubrina asiatica 63 Combretaceae 56 Commelina diffusa 56 Commelinaceae 56,66 Commersonia bartramia 64 Compositae 56,66,71 Connaraceae 56,67
Connarus planchonianus 67 Convolvulaceae 56
Cordia cylindristachya 55 Cordyline fruticosa 60 Corymborkis veratrifolia 69 Crassocephalum crepidiodes 14 Crinum asiaticum 66 Crotalaria mucronata 60,68 Croton hirtus 58
Cucumis hirtus 57 Cucurbitaceae 57
Curculigo orchioides 59 Cyathostemma viridiflorum 53,66 Cyathula prostrata 55,66 Cyclea laxiflora 68 Cymbidium finlaysonianum 69 Cynodon dactylon 17,58 Cyperaceae 57,67,71
Cyperus aromaticus 57 Cyperus compactus 57 Cyperus compressus 57 Cyperus cyperinus 57 Cyperus halpan 57
Cyperus javanicus 57
Cyperus kyllingia 57
Cyperus pilosus 57
Cyperus trialatus 57 Cyrtococcum accrescens 58
Dactyloctenium aegyptium 58 Dalbergia candenatensis 60,68 Dalbergia junghuhnii 68 Datura candida 70
Index to Volume 44 (1)
Davallia denticulata 54 Dendrobium crumenatum 11,62 Dendrophthoe pentandra 60 Dendrotrophe varians 63 Derris elliptica 60 Derris heptaphylla 60,68 Desmodium heterocarpon 60 Desmodium heterophyllum 60 Desmodium umbellatum 60 Desmos dasymaschala 55 Dianella ensifolia 60 Dicranopteris linearis 54 Didymanthus 25,32,38 Didymocarpus 229,202.02 Bye Noe, 9;5455) 37,38,39,40,41,42 Didymocarpus anthonyi 23,24,25,26,27,28,31,32,34,39,40 Didymocarpus azureus 41 Didymocarpus battamensis 38 Didymocarpus caelestis 23 Didymocarpus caeruleus 41 Didymocarpus codonion 32,39,40 Didymocarpus cordatus 23,41 Didymocarpus corneri 25,26,27,29,32,33,34,35 Didymocarpus crinitus 23 Didymocarpus densiflorus 41 Didymocarpus densifolius 23,41 Didymocarpus falcatus 23,32,38 Didymocarpus fasciatus 23 Didymocarpus filicifolius 41 Didymocarpus flavescens 32 Didymocarpus flavobrunneus 23,32,38 Didymocarpus floribundus 36,39,40 Didymocarpus geitleri 23,25,26,27,29,32,34,35 Didymocarpus grandiflorus 38 Didymocarpus heterophyllus 24,25,31,36,38,39,40 Didymocarpus holttumi 41 Didymocarpus kompsoboea 37,38 Didymocarpus lacunosus 38 Didymocarpus leiophyllus 23,25,28,30,31,32,39,40 Didymocarpus leucanthus 23,25,26,27,28,31,32,33,34 Didymocarpus leucocodon 23,25,26,27,28,32,33,34 ~Didymocarpus lithophilus 23,37,38 Didymocarpus longipes 23,38 Didymocarpus oreophilus 39 Didymocarpus parviflorus 23,31,32,34
143
Didymocarpus pectinatus 41
Didymocarpus perditus 38,39
Didymocarpus platypus 23,25,29,32,33
Didymocarpus polyanthoides 32
Didymocarpus primulinus 23,38
Didymocarpus pumilus 38,39,40,41
Didymocarpus puncticulatus 31,37,38,39,40,41
Didymocarpus pyroliflorus 32,34
Didymocarpus quinquevulnerus 25,26,27,28,29,32,33,34,35
Didymocarpus reptans 23
Didymocarpus ridleyanus 38
Didymocarpus robinsonii 32
Didymocarpus rubiginosus 39,40,41
Didymocarpus salicinoides 23,35,36,41
Didymocarpus salicinus 36,41
Didymocarpus salicinus var. major 23
Didymocarpus serratifolius 41
Didymocarpus soldanella 38,39,40,41
Didymocarpus sp. nov. 39,40
Didymocarpus stoloniferus 23,36,37,42
Didymocarpus tahanicus 41
Didymocarpus tiumanicus 41
Didymocarpus venustus 23
Didymocarpus violaceus 38
Didymocarpus violascens 24
Didymocarpus yongii 25,26,27,28,32,33,35,39,40
Dieffenbachia seguine 55
Digitaria ciliaris 58,67
Digitaria violascens 67
Dillenia suffruticosa 57
Dilleniaceae 57
Diodia ocymifolia 63
Dioscorea glabra 57
Dioscorea laurifolia 57
Diplectria viminalis 68
Dipterocarpaceae 14,15,16,67
Dipterocarpus cf. sublamellatus 53,67
Dischidia major 55
Dischidia nummularia 55
Dissochaeta gracilis 61
Dolichandrone spathacea 66
Dracaena elliptica 66
Dracaena porteri 66
Drynaria quercifolia 54
Drynaria sparsisora 65
Durio zibethinus 55
Dysoxylum cauliflorum 68
Echinochloa colona 58 Eclipta prostrata 56
144
Elaeocarpaceae 16,57 Elaeocarpus ferrugineus 57 Elaeocarpus pedunculatus 57 Elaeocarpus spp. 16 Elephantopus scaber 56 Eleusine indica 58 Eleutheranthera ruderalis 67 Embelia ribes 61
Emilia sonchifolia 56 Entada spiralis 60 Eragrostis pilosa 58 Eragrostis tenella 67 Erigeron sumatrensis 56,67 Eriocaulaceae 57 Eriocaulon longifolium 57 Erycibe tomentosa 56 Erythroxylaceae 57 Erythroxylum cuneatum 57 “Eugenia grandis 17
Eugenia jambos 61
Eugenia leucoxylon 69 Eugenia longiflora 61 Eugenia palembanica 61 Eugenia spicata 61
Eugenia spp. 16
Euodia roxburghiana 63 Euphorbia hirta 58 Euphorbiaceae 15,53,57,67,71 Eurya acuminata 64 Excoecaria agallocha 14,58
Fagaceae 67
Fagraea acuminatissima 60 Fagraea auriculata 68 Fagraea fragrans 11,60 Fagraea racemosa 68 Fibraurea tinctoria 61 Ficus aurata 61
Ficus fistulosa 61,68
Ficus grossularioides 61 Ficus heteropleura 61 Ficus kerkhovenii 69
Ficus laevis 69
Ficus microcarpa 61
Ficus obscura var. borneensis 69 Ficus pellucido-punctata 69 Ficus variegata 61
Ficus virens var. glabella 61 Fimbristylis cymosa 57 Fimbristylis dichotoma 57 Fimbristylis ferruginea 67° Fimbristylis griffithii 57 Fimbristylis polytrichoides 57
Gard. Bull. Sing. 44 (1992)
Fimbristylis schoenoides 57 Flagellaria indica 58,67 Flagellariaceae 58,67 Forrestia gracilis 53,66
Gaertnera viminea 70 Galearia fulva 67
Garcinia eugeniaefolia 67 Garcinia hombroniana 67 Garcinia mangostana 59 Garcinia nervosa 67 Garcinia nigrolineata 59 Garcinia spp. 16 Gesneriaceae 23,32 Glochidion microbotrys 67 Glochidion superbum 58 Gloriosa superba 60 Gmelina asiatica 65 Gnetaceae 58
Gnetum macrostachyum 58 Gomphia serrata 69 Gomphrena globosa 55 Goodeniaceae 58167 Gramineae 58,67,71 Grammatophyllum speciosum 69 Guettarda speciosa 63 Guioa pleuropteris 63 Guttiferae 16,59,67 Gynotroches axillaris 63 Gynura procumbens 56
Hedyotis corymbosa 63 Heliotropium indicum 55 Hemigraphis primulifolia 52,54 Heritiera littoralis 64 Heteroboea 25
Hevea brasiliensis 7,58 Hibiscus rosa-sinensis 17,61 Hibiscus surattensis 68 Hibiscus tiliaceus 61
Homo erectus 6
Homo sapiens 6 Hornstedtia leonurus 70 Hoya parasitica 55
Humata heterophylla 65 Hypolytrum nemorum 57 Hypoxidaceae 59
Hyptis brevipes 59
Hyptis capitata 59
Hyptis suaveolens 59,68
Imperata 16 Imperata cylindrica 7,58
Index to Volume 44 (1)
Indorouchera griffithiana 60
Intsia bijuga 60
Ipomoea aquatica 56 Ipomoea batatus 56 Ipomoea cairica 56 Ischaemum indicum 59 Ischaemum muticum 59,67 Ixonanthaceae 59 Ixonanthes reticulata 59 Ixora congesta 63,70
Khaya spp. 17
Knema glaucescens 69 Knema globularia 69 Knema sp. 61 Kompsoboea 38
Labiatae 53168
Languas galanga 65 Lantana camara 65 Laportea interrupta 65 Lasianthus cyanocarpus 70 Lauraceae 15,16,59,68 Leea indica 59
Leeaceae 59 Leguminosae 53,59,68,71 Lemna perpusilla 60 Lemnaceae 60 Lepisanthes rubiginosa 70 Leptaspis urceolata 67 Leucas zeylanica 59 Licuala spinosa 62,69 Liliaceae 60
Limacia scadens 61 Limnophila sessiliflora 64 Linaceae 60
Lindernia antipoda 64 Lindernia crustacea 64 Lindernia sessiliflora 64 Lindsaea divergens 65 Lindsaea ensifolia 54 Linostoma pauciflorum 64 Lithocarpus elegans 53,67 Lithocarpus wallichianus 67 Litsea spp. 16
Litsea umbellata 68 Loganiaceae 60,68 Loranthaceae 15,60 Loxocarpus 41
Ludwigia hyssopifolia 62 Lumnitzera littorea 56 Lumnitzera racemosa 56 Lycopodium cernuum 54
Lygodium circinnatum 54 Lygodium microphyllum 54
Macaca fascicularis 11 Macaranga conifera 58 Macaranga gigantea 58 Macaranga griffithiana 58 Macaranga heynei 58 Macaranga hypoleuca 58 Macaranga triloba 58,67 Macrosolen cochinchinensis 60 Magnoliaceae 60
Mallotus paniculatus 58 Malpighiaceae 60,68 Malvaceae 61,68 Mangifera indica 55 Manihot esculenta 58 Manihot glaziovii 58 Melastoma malabathricum 61 Melastomataceae 61,68 Meliaceae 61,68 Memecylon edule 61 Menispermaceae 61,68 Merremia tridentata 56 Michelia champaca 60 Mikania cordata 56 Mimosa invisa 60
Mimosa pigra 60
Mimosa pudica 60 Mischocarpus sundaicus 63,70 Mnesithea glandulosa 59,67 Moraceae 15,61,68 Morinda citrifolia 63 Morinda umbellata 63,70 Muntingia calabura 57 Murraya koenigii 63
Musa acuminata 61
Myrica esculenta 61 Myricaceae 61 Myristicaceae 61,69 Myrsinaceae 61,69 Myrtaceae 15,16,61,69
Nenga pumila 69 Neolitsea zeylanica 59,68 Nepenthaceae 62 Nepenthes gracilis 62 Nephelium lappaceum 63 Nephrolepis biserrata 54 Neptunia plena 60
Nypa fruticans 62
145
146
Ochnaceae 69
Ocimum basilicum 59 Ocimum tenuiflorum 68 Olacaceae 62
Olea brachiata 69 Oleaceae 69
Onagraceae 62 Oncosperma tigillarium 62 Ophiorrhiza singaporensis 70 Opiliaceae 62
Orania sylvicola 69 Orchidaceae 15,62,69,71 Oxalidaceae 62
Oxalis barrelieri 62
Oxalis corniculata 62 Oxyceros longiflora 63,70
Palmae 15,62,69
Pandanaceae 62,69
Pandanus amaryllifolius 62 Pandanus odoratissimus 62,69 Pandanus parvus 69
Pandanus yvani 62
Panicum maximum 59 Paraboea 38,39,41
Paraboea caerulea 41
Paraboea salicinia var. major 35 Paspalum conjugatum 59 Paspalum orbiculare 59 Passiflora foetida 62,69 Passiflora laurifolia 62 Passiflora suberosa 62 Passifloraceae 62,69
Pectinati 41
Pedaliaceae 62
Peltophorum pterocarpum 17,60 Pennisetum polystachyon 52,59 Peperomia pellucida 62 Phoebe grandis 53,68 Phyllanthus acidus 58 Phyllanthus amarus 58 Phyllanthus debilis 58 Phyllanthus urinaria 58 Phymatosorus scolopendria 54,65 Physalis minima 64,70
Piper betle 63
Piper caninum 69
Piper sarmentosum 63 Piperaceae 62,69 Pithecellobium ellipticum 68 Pityrogramma calomelanos 54,65 Planchonella obovata 64. Plantaginaceae 63
Gard. Bull. Sing. 44 (1992)
Plantago major 63
Platycerium coronarium 54 Pluchea indica 56 | Pogonanthera pulverulenta 68 Pogonatherum paniceum 59 Poikilospermum cordifolium 70 Poikilospermum suaveolens 70 Polyscias fruticosa 55
Pongamia pinnata 68 Portulacaceae 63
Premna corymbosa 65
Psidium guajava 62
Psychotria griffith 70
Psychotria malayana 70 Psychotria rostrata 70
Pterideae 47
Pteridium caudatum ssp. yarrabense 54 Pteridophytes 52
Pteris 47,50
Pteris ensiformis 65
Pteris vittata 54
Pterocarpus indicus 11,17,60 Pterospermum diversifolium 64,70 Pyrrosia lanceolata 65
Pyrrosia longifolia 54
Pyrrosia piloselloides 54
Reissantia indica 66 Renanthera elongata 69 Rhamnaceae 63,69 Rhizophora apiculata 63 Rhizophora mucronata 63 Rhizophoraceae 63,70 Rhodamnia cinerea 11,16,62 Rhodomyrtus tomentosa 62 Rhynchelytrum repens 59 Ricinus communis 58 Rubiaceae 15,63,70,71 Rutaceae 63,70
Saccharum arundinaceum 59 Salacia chinensis 66 Salacia grandiflora 66 Salicini 32,34,36,39,41 Samanea saman 17 Sandoricum koetjape 68 Sansevieria trifasciata 54 Santalaceae 63 Sapindaceae 63,70 Sapium discolor 58 Sapotaceae 64
Scaevola sericea 58,67 Schefflera cephalotes 66
i a
Index to Volume 44 (1)
Schefflera elliptica 55 Schefflera lanceolata 66 Schizaea digitata 54 Scleria corymbosa 57 Scleria levis 57
Scleria oblata 67
Scoparia dulcis 64 Scrophulariaceae 64 Scurrula parasitica 52,53,60 Scyphiphora hydrophyllacea 63 Sesamum radiatum 52,62 Sesuvium portulacastrum 54 Sida rhombifolia 61 Simaroubaceae 64 Smilacaceae 64
Smilax megacarpa 64 Solanaceae 64,70
Solanum melongena 64 Solanum torvum 64,70 Sonneratia alba 64 Sonneratia ovata 64 Sonneratiaceae 64
Sparganophorus sparganophora 56
Spathoglottis plicata 52,62,69 Spilanthes iabadicensis 56 Sporobolus indicus 59 Stachytarpheta indica 65,70 Stenochlaena palustris 54 Sterculia coccinea 70 Sterculiaceae 64,70 Streblus elongatus 61 Suregada multiflora 58,67 Swietenia macrophylla 17 Symplocaceae 64 Symplocos fasciculata 64 Synedrella nodiflora 56,67
Taenitis blechnoides 54 Talinum paniculatum 53,63 Tarenna costata 63
Tarenna fragrans 63
Tectaria semipinnata 53,65 Terminalia catappa 17,56 Tetracera indica 57
Theaceae 64
Thespesia populnea 14,61,68 Thoracostachyum bancanum 57
Thrixspermum amplexicaule 52,62
Thrixspermum calceolus 69 Thunbergia fragrans 54 Thunbergia grandiflora 54 Thymelaeaceae 64 Thysanolaena latifolia 53,59
Tiliaceae 64
Timonius wallichianus 11,63 Trema cannabina 64
Trema tomentosa 64 Tridax procumbens 56,67 Tristaniopsis whiteana 62,69 Tristellateia australasiae 60,68 Triumfetta rhomboidea 64 Turnera ulmifolia 64,70 Turneraceae 64,70
Typha angustifolia 53,64 Typhaceae 64
Ulmaceae 64
Umbelliferae 65
Uncaria gambir 7
Uncaria glabrata 70
Urceola lucida 66
Urena lobata 61
Urnularia flavescens 66 Urophyllum streptopodium 70 Urticaceae 65,70
Vanilla griffithii 62,69 Ventilago malaccensis 69 Verbenaceae 65,70 Vernonia cinerea 56 Vitaceae 65
Vitex pinnata 65
Vitex trifolia 65,70
Wedelia biflora 56,67 Wedelia trilobata 56 Willughbeia coriacea 66 Willughbeia grandiflora 66
Ximenia americana 62 Xylocarpus granatum 61
Youngia japonica 56 Zingiberaceae 65,70
Zoysia 17 Zoysia matrella 59
147
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INDEX
Volume 44 (Part 2)
Basionyms and synonyms appear in italics. Page numbers in italics indicate the presencve of illustrations.
Acanthaceae 128
Achnanthes 77
Achnanthes brevipes var. intermedia 77,80
Achnanthes hauckiana 77, 80
Achnanthes lewisiana 77,80
Achnanthes longipes 77, 80
Actinocyclus 77
Actinocyclus ehrenbergii var. sparsa 77,80
Actinocyclus octonarius 77, 80
Actinocyclus platensis 78, 80
Actinoptychus 78
Actinoptychus senarius 78, 80
Adiantaceae 127
Adiantum fructuosum 127
Adiantum latifolium 127
Aeschynomene americana 130
Ammania baccifera 131
Amphora 77,80
Amphora acutiuscula 78,80
Amphora angusta var. eulensteinii 78,80
Amphora angusta var. oblongella 78,80
Amphora angusta var. ventricosa 78,80
Amphora coffeiformis 78,80
Amphora crassa 79,80
Amphora decipiens 79, 80
Amphora graeffi var. minor 79,80
Amphora holsatica 79,80
Amphora ostrearia 79,80
Amphora proteus 79,80
Amphora turgida 79,80
Amphora valida 79,80
Amphora wisei 81
Asclepiadaceae 128
Asterionella 81
Asterionella japonica 81,83
Bacillaria 81
Bacillaria paradoxa var. tumidula 81,83 Bacteriastrum 81
Bacteriastrum delicatulum 81,83 Bacteriastrum elongatum 81,83 Bacteriastrum hyalinum 81,83
Balsaminaceae 128
Biddulphia 81
Biddulphia biddulphiana 81,83,85 Biddulphia mobiliensis 82,83 Biddulphia petitiana 82,83 Biddulphia recticulata 82,85 Biddulphia vesiculosa 82,85
Callistemon 135,136,138 Callistemon citrinus 135,136,137 Callistemon lanceolatus 138 Caloneis 82 Caloneis alpestris 82,85 Caloneis bacillum 82,85 Caloneis egena 82,71 Caloneis liber 82,85 Caloneis linearis 82,85 Campylodiscus 84 Campylodiscus fastuosus 84,85 Campylodiscus hypodromus 84,85 Campylodiscus ralfsii 84,85 Canna indica 128 Cannaceae 128 Celastraceae 128 Chaetoceros 84 Chaetoceros danicum 84,87 Chaetoceros lorenzianum 84,89 Chaetoceros peruvianum 84,87 Chaetoceros tetrastichon 84,87 Climacosphenia 84 Climacosphenia moniligera 84,87 Cocconeis 86 Cocconeis dirupta 86,87 Cocconeis disculoides 86,87 Cocconeis heteroidea 86,87 Cocconeis pelta 86,87 Cocconeis placentula var. euglypta 86,87 Cocconeis pseudomarginata var. intermedia 86,87 Cocconeis speciosa 86,87Cocconeis sublittoralis 87,88 Compositae 129 Convolvulaceae 129 Coscinodiscus 88
150
Coscinodiscus argus 88,89 Coscinodiscus decipiens 88,89 Coscinodiscus decrescens 88,89 Coscinodiscus marginatus 88,89 Coscinodiscus nodulifer 88,89 Coscinodiscus oculus-iridus 88,89 Coscinodiscus radiatus 88,89 Coscinodiscus wailesii 88,89 Coscinasira 90
Coscinosira oestrupii 89,90 Cyclotella 90
Cyclotella kutzingiana 90,93 Cyclotella menghiniana 90,93 Cyclotella operculata 90,93 Cyclotella striata 90,93 Cyclotella stylorum 90,93 Cymatosira 90
Cymatosira lorenziana 90,93 Cymbella 90
Cymbella pusilla 90,93 Cyperus papyrus 132
Delphineis 91
Delphineis surirella 91,93 Denticula 91
Denticula subtilis 91,93 Desmanthus virgatus 130,132 Diploneis 77,91 Diploneis bombiformis 91,93 Diploneis bombus 91,93
Diploneis bombus var. densestriata 91,93
Diploneis chersonensis 91,93 Diploneis coffaeiformis 92,93 Diploneis crabro 92,93 Diploneis interrupta 92,93
Diploneis exemta var. digrediens 92, 93
Diploneis gravelleana 92,93
Diploneis incurvata 92,93
Diploneis interrupta 92,93
Diploneis interrupta var. gorjanovicil 92,93
Diploneis littoralis 92,97
Diploneis nitescens 94,97
Diploneis notabilis 94,97
Diploneis oculata 94,97
Diploneis puella 94,97
Diploneis smithii 94,97
Diploneis smithii var. rhombica 94,97
Diploneis subovalis 94,97
Diploneis vetula 94,97
Diploneis weissflogii 95,97
Dipterocarpaceae 129
Dipterocarpus sublamellatus 129
Gard. Bull. Sing. 44 (1992)
Donkinia 95 Donkinia recta 95,97
Eucalyptus citriodora 138 Eugenia caryophyllata 135,136,138 Eugenia caryophyllus 135 Eunotogramma 95 Eunotogramma laeve 95,97
Fragilaria 95
Fragilaria cylindrus 95,97 Fragilaria lapponica 95,97 Fragilaria leptostauron var. dubia 95,97 Fragilaria oceanica 96,97 Fragilaria schulzi 96,97
Gramineae 130
Grammatophora 96
Grammatophora hamulifera 96,97
Grammatophora marina var. adriatica 96,97
Grammatophora oceanica 96,97
Grammatophora undulata 96,97
Gyrosigma 96
Gyrosigma balticum 96,100
Gyrosigma distortum 98,100
Gyrosigma exile 98,100
Gyrosigma fasciola var. sulcata 98,100
Gyrosigma grovei 98,100
Gyrosigma simile 98,100
Hantzchia 98
Hantzchia amphioxys var. capitata 98,100
Hantzchia virgata 98,100
Hedyotis pumila 131
Hemigraphis primulaefolia 128
Huttoniella 99
Huttoniella reichardtii 99,100
Hydrocera triflora 128
Impatiens balsamina 128 Ipomoea obscura 129 Ipomoea pes-tigridis 129 Isthmia 99
Isthmia enervis 99,100
Leguminosae 130
Licmophora 99
Licmophora abbreviata 99,100 Licmophora ehrenbergii 99,100 Licmophora flabellata 99,100 Licmophora gracilis 99,100
Index to Volume 44 (2)
Loranthaceae 130 Lythraceae 131
Mangifera foetida 131 Mastogloia 99
Mastogloia angulata 99,104 Mastogloia binotata 101,104 Mastogloia citrus 101,104 Mastogloia fimbriata 101,104 Mastogloia ovata 101,104 Mastogloia quinquecostata 101,104 Maytenus emarginata 128 Melaleuca 135,136,138 Melaleuca cajuputi 135,136,137 Melaleuca dealbata 135,136,137 Melaleuca genistifolia 135,136,137 Melosira 101
Melosira granulata 101,104 Melosira nummuloides 101,104 Myrtaceae 135
Navicula 73,77,102 Navicula brasiliensis 102,104 Navicula clavata 102,104 Navicula cuspidata 102,104 Navicula distans 102,104 Navicula forcipata var. suborbicularis 102,104 Navicula glacialis 102,104 Navicula grundleri 102,104 Navicula halophila 102,104 Navicula lyra 103,104 Navicula menaiana 103,104 Navicula monilifera 103,104 Navicula monilifera var. constricta 103,104 Navicula nicaeensis 103,104 Navicula pennata 103,104 Navicula pi 103,104 Navicula platessa 103,108 Navicula platyventris 105,320 Navicula plicata 105,108 Navicula praetexta 105,108 Navicula pusilla var. jamalinensis 105,108 Navicula radiosa 105,108 - Navicula ramosissima 105,108 Navicula ramosissima var. caspia 105,108 Navicula reichardtii 105,108 Navicula rhaphoneis 106,108 Navicula transfuga 106,108 Navicula transitans 106,108 Navicula yarrensis 106,108 Navicula zostereti 106,108
151
Neuropeltis racemosa 129
Nitzschia 73,77,106
Nitzschia amphibia 106,108
Nitzschia apiculata 106,108
Nitzschia bilobata var. minor 106,108
Nitzschia brebissonii var. borealis 107,108
Nitzschia cocconeiformis 107,108
Nitzschia commutata 107,108
Nitzschia constricta 107,108
Nitzschia granulata 107,108
Nitzschia ignorata 107,108
Nitzschia longissima 107,108
Nitzschia navicularis var. typica 107,108
Nitzschia panduriformis 109,112
Nitzschia parvula var. terricola 109,112
Nitzschia punctata 109,112
Nitzschia punctata var. coarctata 109,112
Nitzschia sigma var. rigida 109,112
Nitzschia sigmoidea 109,112
Nitzschia subtilis 109,112
Nitzschia tryblionella var. victoriae 109,112
Nitzschia vermicularis 110,112
Odontella 110
Odontella aurita 110,112
Odontella aurita var. obtusa 110,112 Oldenlandia pumila 131
Opephora 110
Opephora martyi 110,112 Opephora schwartzii 110,112
Paralia 110
Paralia sulcata 110,112 Pedaliaceae 131
Pennisetum polystachyon 130 Pennisetum purpureum 130 Pennisetum setosum 130 Plagiodiscus 110
Plagiodiscus nervatus 110,112 Plagiogramma 111 Plagiogramma staurophorum 111,112 Pleurosigma 77,111
Pleurosigma aestuarii 111,112 Pleurosigma delicatulum 111,114 Pleurosigma elongatum 111,114 Pleurosigma formosum 111,114 Pleurosigma intermedium 111,114 Pleurosigma majus 111,114 Pleurosigma marinum 113,114 Pleurosigma normani 113,114
152
Pleurosigma nubecula var. mauritiana 113,114
Pleurosigma salinarum 113,114
Pleurosigma salinarum var. boyeri 113
Podocystis 113
Podocystis adriatica 113,114
Polygonaceae 131
Polygonum orientale 131
Porophyllum ruderale 129
Portulacaceae 131
Psammodiscus 113
Psammodiscus nitidus 113,117
Pyxidicula 113
Pyxidicula africana 113,117
Rhabdonema 115
Rhabdonema adriaticum 115,117
Rhaphoneis 115
Rhaphoneis amphiceros 115,117
Rhaphoneis amphiceros var. tetragona 115,117
Rhaphoneis castracanii 115,117
Rhodomyrtus tomentosa 138
Rhopalodia 115
Rhopalodia gibba var. ventricosa 115,117
Rhopalodia gibberula 115,117
Rhopalodia gibberula var. vanheurckii 125, 107
Rubiaceae 131
Scurrula ferruginea 130,131
Scurrula fusca 130
Scurrula parasitica 130
Secamone elliptica 128
Secamone micrantha 128
Sesamum radiatum 131
Setaria barbata 130
Stauroneis 116
Stauroneis membranaceae 116,117 Stephanopyxis 116
Stephanopyxis turris var. polaris 116,117 Striatella 116
Striatella unipunctata 116,117
Surirella 116
Surirella amoricana 116,117
Surirella fastuosa 116,117
Surirella fastuosa var. recedens 116,117 Synedra 116
Synedra amphicephala 116,119 Synedra crystallina 118,119
Synedra demerare 118,119 |
Synedra fasciculata var. truncata 118,119 Synedra formosa 118,119
Gard. Bull. Sing. 44 (1992)
Synedra gaillonii 118,119
Synedra hennedyana 118,119
Synedra provincialis var. tortuosa 118,119
Synedra tabulata var. grandis 118,119
Syzygium 135,136,138
Syzygium aromaticum 135,136,137,138
Syzygium aromaticuum 135.138
Syzygium iambolanum 138
Tabellaria 118
Tabellaria fenestrata 118,119
Talinum paniculatum 131 Thalassionema 120
Thalassionema nitzschioides 120,121 Thalassiosira 120
Thalassiosira eccentrica 120,121 Thysanolaena latifolia 130
Trachyneis 120
Trachyneis antillarum 120,121 Trachyneis antillarum var. kurzii 120,121 Trachyneis aspera 120,121
Trachyneis aspera ar. intermedia 120,121 Trachyneis aspera var. pulchella 120,121 Triceratium 122
Triceratium broecku 121,122. Triceratium dubium 121,122 Triceratium zonulatum 121,122 Tropidoneis 122
Tropidoneis maximan 121,122 Trybliophychus 122
Trybliophychus cocconeiformis 121,122 Typha angustifolia 132
Typhaceae 132
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JUN 15 1993 aRmpBorRETUM ISSN 0374-7859
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SINGAPORE
| VOL. 44 (Part 1)
THE GARDENS’ BULLETIN June 1992 x ()
® | CONTENTS j Y TAN, HTW., CHUA, K.S. and TURNER, IM: . Rae eer at Ora Of SINGAPOTe PLOJECt: ... ze. soos. cs cecavesescensseneyrnesanvenneseceonces 1-2 - CORLETT, Richard T.: The Angiosperm Flora of Singapore 1. Introduction ..............ccceeseseseeeteeetetsees 3-21 _ KIEW, R:: Five New Species of Didymocarpus (Gesneriaceae) from a AY Sol, Se ene Dy. tad siseies set enneatnttanaies 23-42 ‘Xy) =©©CORNER, E.J.H.: Da Notes on the Development of the Fruit-bodies of f . Four Malayan Species of Amanita (Basidiomycetes) ............:::::cceseeseeseseeeessesteees 43-45 6 - AZIZ, Bidin and RAZALI, Jaman: K _ Notes on the Rare Fern, Pteris holttumii C. CMI. 1.0... see ec ec eee tee tees seen 47-50 ¢ _ TURNER, I.M., TAN, H.TW. and CHUA, K.S.: ‘j - A Botanical Survey of Pulau Ubi ...00......... cece eect eeeeeeeeeeeeeeseteeteeseneeneteeeneniens 51-71 4}
Published by National Parks Board Singapore Botanic Gardens Cluny Road Singapore 1025
Bi | ; PIE OE mp EOD EOE A aa
:
GARDENS’ BULLETIN
EDITORIAL COMMITTEE
Chairman: Bernard T.G. Tan, B.Sc.Hons. (Singapore); D.Phil. (Oxon.)
Editor: S.Y. Geh, B.Sc.Hons., M.Sc. (Singapore); Dip.Hort.Sc. (Massey) ©
Members: Syed Yusoff Alsagoff | 1
C.J. Goh, B.Sc.Hons. (Singapore); Ph.D. (Newcastle upon Tyne)
Leo W.H. Tan, B.Sc., Ph.D. (Singapore)
W.K. Tan, B.A. (Wms Coll., Mass.); M.Sc. (M.S.U., Mich.); | Ph.D. (U.M., FI.)
Lawrence L.C. Leong, B.Sc., M.Sc., Ph.D. (Malaya) |
TW. Foong, B.Sc.Hons., Ph.D. (Cant.)
Jennifer Ng-Lim Cheo Tee, B.Sc., M.A. (Dublin)
Reviewers: The late R.E. Holttum Hsuan Keng B.L. Burtt
The Gardens’ Bulletin is published twice yearly by the National Parks Board, Singapore. Neither the National Parks Board nor the Editorial Committee is responsible for the opinions ~ or conclusions expressed by the contributing authors. :
ee
The Gardens’ Bulletin is priced at Singapore $16.00 cts. excluding postage. Overseas sub- scribers are required to make payment in the form of bank drafts or international money orders in Singapore currency payable to the Executive Director, National Parks Board, Singapore.
Instructions for contributing authors are found behind the Contents Table.
THE GARDENS’ BULLETIN
SINGAPORE
VOL. 44 (Part 1)
CONTENTS
TAN, H-TW., CHUA, K.S. and TURNER, I.M.:
The Angiosperm Flora of Singapore Project .................cccecececceeeeeeees
CORLETT, Richard T.:
The Angiosperm Flora of Singapore 1. Introduction ......................
KIEW, R.: Five New Species of Didymocarpus (Gesneriaceae) from
SE ke allah Repeal» Dept il ae See aE
CORNER, E.J.H.: Notes on the Development of the Fruit-bodies of
Four Malayan Species of Amanita (Basidiomycetes) ................:+
AZIZ, Bidin and RAZALI, Jaman:
Notes on the Rare Fern, Pteris holttumti C. Chr. ...............000cc0eeeeeeees
TURNER, I.M., TAN, H.TW. and CHUA, K:S:.:
= Botanical Survey of Pulau Ubim ......-.....22......2..10c...scc0ceccessseneeeeeeees
Published by National Parks Board Singapore Botanic Gardens Cluny Road Singapore 1025
June 1992
Eps Fi 47-50
INSTRUCTIONS TO AUTHORS
Manuscripts: The Editorial Committee of the Gardens’ Bulletin will be glad to receive and consider for publication original research findings and reviews of progress in the fields of botany, horticulture, and allied subjects. Contributions must be original and the material must not have been submitted or, if accepted, be submitted for publication elsewhere.
Two copies of the manuscript should be submitted, typed or typeprinted, and if typed, then the top copy must be one of the two. Type or print on one side only, with double-line spacings and a margin of at least 4 cm. Do not type all the letters of any word in capitals. Underline only in pencil: with a straight line for italic type face and wavy line line for bold type face. Authors should see the layout of other papers recently published in this journal to ensure that papers submitted conform as closely as possible to the accepted pattern. Numerical data should only be included if it is essential to the argument and this can be presented either in the form of tables or diagrams.
Titles and authors: The title should give a concise description of the content of the paper. The name(s) and affiliation(s) of author(s) must be given below the title. Lengthy papers and those of a complex nature must have the contents listed at the beginning of the paper.
Scientific names: The complete scientific name — genus, species, authority, and cultivar where appropriate — must be cited for every organism at time of first mention. The generic name may be abbreviated to the initial thereafter except where intervening references to other genera with the same initial could cause confusion.
Tables: All tables should be numbered and carry headings describing their content. These should be comprehensive without reference to the text.
Abbreviations: Standard chemical symbols may be used in the text (e.g. IAA, IBA, ATP), but the full term should be given on the first mention. Dates should be cited as: 3 May 1976. Units of measurement should be spelled out except when preceded by a numeral where they should be abbreviated in standard form: g, mg, ml, etc. and not followed by stops.
Literature citations: Citations in the text should take the form: King and Chan (1964). If several papers by the same author in the same year are cited, they should be lettered in sequence (1964a), (1964b), etc. When papers are by three or more authors they should be cited as e.g., Geesink et al. (1981). All references must be placed in alphabetic order according to the surname of the (first) author and in the following form:
Singh, H. (1967). Sclereids in Fagraea. Gard. Bull. Sing. 22, 193-212.
Abbreviations of titles of journals should be those of the World List of Scientific Periodicals (4th Edition) or the Selected Abbreviated Titles of Biological Journals (London: Institute of Biology).
References to books and monographs should be cited according to the following form: Ridley, H.N. (1930). The Dispersal of Plants Throughout the World, L. Reeve; Ashford, Kent; 242-255.
For literature citations in taxonomic papers the following style is required: Medinilla alternifolia Bl., Mus. Bot. Lugd.-Bat. I:2 (1849) 19. Sterculia acuminatissima Merr., Philip. J. Sci. 21 (1922) 524.
Illustrations: Drawings should be done in Indian ink. Authors should indicate where individual illustrations receive first mention in the text.
Offprints: Authors will be given 50 offprints gratis. Additional copies must be ordered and paid for, prior to publication.
Manuscripts should be sent to: THE EDITOR, GARDENS’ BULLETIN, SINGAPORE, NATIONAL PARKS BOARD, SINGAPORE BOTANIC GARDENS, CLUNY ROAD, SINGAPORE 1025.
per cr
The Angiosperm Flora of Singapore Project
H.TW. TAN, K.S. CHUA and I.M. TURNER Department of Botany National University of Singapore Lower Kent Ridge Road Singapore 0511 Republic of Singapore
EFFECTIVE PUBLICATION DATE: 15 MAR 1993
The Angiosperm Flora of Singapore Project is long-term and was initiated on Ist November, 1988. A piecemeal, family by family approach is to be adopted. The approximately 200 families containing an estimated 2,400 native and naturalized species are to be studied by either experts willing to contribute to the project or the researchers of the project. Family treatments will be published in the Gardens’ Bulletin, Singapore as and when ready. It is hoped that the project will be completed within ten years from the publication of the first family treatment. Ultimately, when all family treat- ments are completed, a monograph compiling all the updated family treatments will be published.
The main objective of this National University of Singapore funded project is to produce an Angiosperm Flora of the Republic of Singapore. So far, only annotated species lists have been written (Ridley, 1900, 1901; Sinclair, 1953, 1956; Keng, 1973, 1974a, 1974b, 1976, 1978, 1980, 1982, 1983, 1985, 1986, 1987, 1990). During World War II, staff members of the Singapore Botanic Gardens Herbarium compiled the Flora of Syonan but this consisted only of a species list and is extant only in typescript (Keng, 1990). Recently, Keng (1990) published the Concise Flora of Singapore but this included only about 1,200 species, about 44 per cent of the estimated total flora, and excluded the monocotyledons. A checklist of all vascular plants recorded to occur in Singapore has recently been compiled (Turner, Chua and Tan, 1990).
The target audience for this Flora are scientists and informed laymen who require precise information for the identification of angiosperm specimens from Singapore. Thus, this flora will be of a traditional format using concise and precise language.
Collections of plants from all accessible parts of the main island and islands of the Republic of Singapore are being carried out. All plants extant and previously found in Singapore will be catalogued, described and distinguished by keys. Illustration of representative taxa by line drawings and photographs will also be included. Expert systems for the identification of the taxa are planned. A database is currently being developed.
The advisors of this project are Drs Hsuan Keng and Tan Wee Kiat (Executive Director, National Parks Board). The researchers include Drs Hugh Tan Tiang Wah (editor), Ian Mark Turner and Mr Chua Keng Soon.
Authors for family treatments are welcome and advised to contact the editor to discuss choice of families and to obtain the format for the Flora. Please address correspondence to:
2 Gard. Bull. Sing. 44(1) (1992)
Dr Hugh TAN
Editor, Angiosperm Flora of Singapore Department of Botany
National University of Singapore Lower Kent Ridge Road
Singapore 0511
REPUBLIC OF SINGAPORE
References
Keng, H. (1973). Annoted list of seed plants of Singapore (1). Gard. Bull. Sing. 26, 233-237.
_____. (1974a). Annotated list of seed plants of Singapore (II). Gard. Bull. Sing. 27, 67-83.
. (19746). Annotated list of seed plants of Singapore (III). Gard. Bull. Sing. 27, 247-266.
_____. (1976). Annotated list of seed plants of Singapore (IV). Gard. Bull. Sing. 28, 237-258.
_____. (1978). Annotated list of seed plants of Singapore (V). Gard. Bull. Sing. 31, 84-113.
. (1980). Annotated list of seed plants of Singapore (VI). Gard. Bull. Sing. 33, 329-367.
. (1982). Annotated list of seed plants of Singapore (VII). Gard. Bull. Sing. 35, 83-103.
. (1983). Annotated list of seed plants of Singapore (VIII). Gard. Bull. Sing. 36, 103-124.
. (1985). Annotated list of seed plants of Singapore (IX). Gard. Bull. Sing. 38, 149-174.
____. (1986). Annotated list of seed plants of Singapore (X). Gard. Bull. Sing. 39, 67-95.
. (1987). Annotated list of seed plants of Singapore (XI). Gard. Bull. Sing. 40, 113-132.
____. (1990). The concise Flora of Singapore: gymnosperms and dicotyledons. Singa- pore Univ. Press; Singapore.
Ridley, H.N. (1900). The Flora of Singapore. J. Straits Branch Roy. As. Soc. 33, 27-196.
_____. (1901). Supplementary notes on the Flora of Singapore. J. Straits Branch Roy. As. Soc. 35, 84-90.
Sinclair, J. (1953). Additions to the Flora of Singapore and new localities in Singapore for some plants thought to be extinct. Gard. Bull. Sing. 14, 30-39.
. (1956). Additions to the Flora of Singapore and new localities in Singapore for some plants thought to be extinct — Part II. Gard. Bull. Sing. 15, 22-30.
Turner, I.M., Tan, HW. and Chua, K.S. (1990). A checklist of the native and natural- ized vascular plants of the Republic of Singapore. J. Sing. Nat. Acad. Science 18 & 19, 58-88.
The Angiosperm Flora of Singapore 1. Introduction
RICHARD T. CORLETT Department of Botany University of Hong Kong Hong Kong
EFFECTIVE PUBLICATION DATE: 15 MAR 1993
The Republic of Singapore is an independent state of 2.7 million people at the southern tip of the Malay Peninsyla, 137 km north of the equator (Fig. 1). It consists of the island of Singapore and more than 50 smaller islands. The main island is 42 km by 23 km at its widest points and has an area of 574 km?, of which more than 30 km? has been added by recent land reclamation (Anon, 1990). It is separated from Malaysia by shallow straits, 0.6 km wide at the narrowest point. The largest of the other islands are Pulau Tekong Besar (1,793 ha), Pulau Ubin (1,019 ha) and Sentosa (309 ha). The total land area, including all the islands, is 626 km?.
The topography of Singapore is predominantly low, with an average elevation of only 15.1 m (Thomas, 1991). The landscape of the main island can be roughly divided into three parts. In the centre of the island is a hilly region of granite and other igneous rocks, rising to a maximum of 162 m at Bukit Timah Hill. The western, southwestern and southern region, including most of the southern islands, consists of a variety of sharply folded sedimentary rocks with northwest-trending ridges and valleys. The eastern region is relatively flat and covered in semi-consolidated deposits of sand and gravel. Low-lying coastal plains and the lower parts of river valleys are filled with recent alluvium.
Singapore’s “rivers” are large streams with broad estuaries, which result from flood- ing of valleys incised during periods of low sea-levels in the Pleistocene. Around the coastline, cliffs and other rocky shores are of limited extent, except on some of the southern islands. Until recently, most shores consisted of mud and sand in varying proportions. Muddy shores with mangroves predominated, except along the southeast coast, which was lined with sandy beaches. Today, however, much of the coastline is entirely artificial as a result of extensive land reclamation and coastal development.
Palaeogeography
Singapore is part of the Southeast Asian extension of the great Eurasia plate and is largely unaffected by the tectonic and volcanic activity around the plate margins to the west, south and east. The regional pattern of land and sea, however, has changed dramatically many times during the last million years or so, largely as a result of changes in sea-level. During glacial periods, sea-levels were up to 200 m lower than at present, exposing most of the Sunda shelf and joining the Malay Peninsula, Sumatra, Java and Borneo into one land mass (“Sundaland”), with Singapore somewhat west of centre (Morley and Flenley, 1987). At the opposite extreme, reported Holocene sea- levels up to 5 m higher than present (Geyr and Kudrass, 1979; Pirazzoli, 1991), would have substantially reduced Singapore’s land area. Glacial periods occupied much more
3
Fig. 1 Map of Singapore showing major roads and reservoirs. Inset map shows the geographical position of Singapore.
Angiosperm Flora of Singapore 1 5
of the Pleistocene than interglacials, so the present geography of the region must be seen as the exception rather than the rule. Singapore is cut off from the Malay Penin- sula by water less than 10 m deep at the shallowest crossing, which implies a final separation about 7,000 years ago (Pirazzoli, 1991). The sea is deeper in the Singapore Straits to the south, so Singapore would have been cut off from Riau before it was separated from the Peninsula.
The repeated alternation of glacial and interglacial periods was also reflected in the region’s climate. A decrease in total rainfall and increase in rainfall seasonality during the glacial periods has been suggested for that part of the Sunda Shelf which includes Singapore (Morley and Flenley, 1987; Heaney, 1991). A pollen assemblage from middle Pleistocene Subang, 300 km northwest of Singapore, is striking for the dominance of pine and grass pollen and absence of typical rain forest taxa, suggesting a very dif- ferent climate. In Singapore itself, the Pleistocene Old Alluvium, which blankets much of the eastern part of the island, seems to have been deposited under far more seasonal climatic conditions during a period of glacial low sea-levels (Gupta ef al., 1987). The question of glacial temperatures is more controversal. Oxygen isotope and foraminiferal data suggest a lowering of sea surface temperatures by at most 2°C at the last glacial maximum, in the vicinity of the Sunda shelf (CLIMAP, 1981). However, this is very difficult to reconcile with evidence for much greater temperature changes at higher altitudes in the region (Walker and Flenley, 1979). Sea-level temperatures 5°C or more below present have been suggested for near-equatorial Amazonia (Bush et al., 1990) and this possibility must be considered for equatorial Asia (Liu, K.B., pers. comm.). In any case, it is clear that as little as 10-15,000 years ago and for most of the Pleistocene, Singapore would have been occupied by a vegetation and flora very different from today’s and probably lacking a modern analogue elsewhere in the region.
On a longer time-scale, the phytogeography of the region has been affected by its complex tectonic history. The Malay Archipelago, as it exists today, was created by a mid-Miocene collision between Australia-New Guinea and Southeast Asia, in the vicinity of Sulawesi (Audley-Charles, 1987). There has never been a dry land connec- tion between Australia and Southeast Asia, even at extreme Pleistocene low sea-levels, but the many islands between Sunda and Sahul (Australia-New Guinea) must have greatly facilitated floristic exchange.
Until recently, paleogeographic reconstructions of the region before convergence showed a huge gap between Southeast Asia and Australia, making earlier biotic inter- change between the regions unlikely (e.g. Audley-Charles, 1981). It now appears that, not only was this gap much narrower than once believed, but the whole of Southeast Asia is made up of a series of continental fragments rifted from northeastern Gond- wanaland. The dating of these events is still contentious. Even if rifting of the major fragments occurred in the Jurassic (Audley-Charles, 1987), it is unlikely that the rifted fragments carried an Angiosperm flora at the beginning of their journey north, although they may well have acted as “stepping stones” between Australia and Asia later on. If, as much of the evidence suggests, the major fragments were already welded to Eurasia by the early Mesozoic (Metcalfe, 1990), they cannot have carried angiosperms. At Gunong Belumut, 75 km north of Singapore, there is a Later Permian fossil flora of undoubted Cathaysian (i.e. tropical Eurasian) affinities, with no Gondwanic ele- ments (Hutchison, 1989). However, other blocks that make up modern Sundaland had Gondwanic floras at this time, showing they had not yet separated from that con- tinent. Moreover, smaller fragments apparently continued to be added to the margins of Southeast Asia during the Jurassic and Cretaceous. Jt thus appears that the sea
6 Gard. Bull. Sing. 44(1) (1992)
between Australia and Eurasia has never been empty, although speculation on the details of the regional palaeogeography are premature at this stage.
India, which did not rift from Gondwanaland until the early Cretaceous, and then moved very rapidly north, provides another possible route for one-way transport of Gondwanic angiosperms to Eurasia. India’s collision with Tibet occurred in the Eocene.
Human Impact
Early man arrived in southeast Asia a million or so years ago, followed by modern Homo sapiens at least 50,000 years ago. It seems likely that human population den- sities in the equatorial lowlands were low before the introduction of agriculture within the last 4,000-6,000 years but it would be a mistake to underestimate the possible impact of pre-agricultural man on the biota of the region. The arrival of Homo erectus, an adaptable and intelligent hunter, is likely to have affected populations of large, ground-dwelling herbivores and this impact would have extended into the forest canopy when, later, throwing spears or similar weapons were added to man’s arsenal. The extinction of large herbivores would have influenced forest structure both directly, through reduced grazing, browsing and trampling, and indirectly through the loss of their role in seed dispersal. Man’s use of fire — of uncertain antiquity — must have been most significant during the dryer, glacial episodes, but droughts occur even during the wet interglacials and extra sources of ignition increase the risk of fire.
The process of homogenisation of the economic and weed floras of the Old World tropics must have started early. The Malay Archipelago has been linked by a maritime trading network from prehistoric times and has had trade links with China, India and the Middle East for at least 1,500 years (Dunn, 1975). With the arrival of Europeans in the early fifteenth century and the establishment of trans-Pacific trading routes, neotropical crops and weeds also made their appearance. Maize, tobacco, chilli, peppers, papaya, pineapple and sweet potato all became established in the region before the end of the fifteenth century (Reid, 1988). In recent times, Singapore, as a port city with a large botanic gardens, may have been the point of entry to the region for many exotic plants.
Although human populations must have lived in Singapore for thousands of years, the first definite historical accounts of a settlement on the island date from the four- teenth century, when Temasek (later called Singapura) appears in Javanese, Chinese and Vietnamese records. Temasek/Singapura was probably not the great trading city described in the Malay Annals but there is archaeological evidence for a substantial settlement at the mouth of the Singapore River in the fourteenth century (Miksic, 1985). Tome Pires, who lived in Malacca from 1512-1515, says that Chinese vessels came for the “infinite quantities of the black wood that grows in Singapore” (Cortesao, 1944). Temasek went into gradual decline during the fifteenth century and the last vestiges of the settlement were burned by the Portuguese in 1613. For the next 300 years, the island disappeared from history but there is no reason to believe it was ever uninhabited.
When the British arrived in 1819, the population of Singapore island consisted of about 1,000 people. Most of these were boat dwellers: the Orang Kallang, who lived in the swamps at the mouth of the Kallang River; the Orang Seletar, who lived in mangrove areas along the north side of the island; and the Orang Gelam, in the Singapore River (Logan, 1847; Thomson, 1848). These people apparently grew no crops but may have had a significant ecological impact through their hunting and collecting activity. The remainder of the population consisted of Malays and Chinese living in
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Angiosperm Flora of Singapore 1 7
a small settlement at the mouth of the Singapore River or growing gambier in the surrounding hills.
The foundation of the British colony led to a rapid and sustained rise in population. From the beginning, Singapore was primarily a trading centre, but the cultivation of cash crops also expanded and spread into the interior of the island. Many crops were grown during the nineteenth century but, except on the sandy soils of the southeast coast, where coconuts were the major crop, most of the initial clearance of primary forest seems to have been for the cultivation of gambier (Uncaria gambir (Hunt.) Roxb.). Gambier was grown for export to China, and later Britain, where it was used for tanning leather and as a dye. It grows best on soil newly-cleared of forest and each plantation required a roughly equal area of forest to provide firewood for boiling the gambier leaves (Jackson, 1965). The Chinese gambier growers rarely had any legal title to the ground and simply moved on when the soil was exhausted and the fuelwood supply insufficient. Abandoned plantations were invaded by the grass Jmperata cylindrica (L.) P. Beauv. or by secondary scrub.
Gambier continued to be a major crop in Singapore until 1890, after which the area declined rapidly. By this time, little of the original forest cover remained and most surviving forest fragments had been heavily exploited for timber and firewood (Corlett, 199la, b). After the departure of the gambier growers, the cultivation of other crops, particularly pineapples, increased. However, it was an entirely new crop, rubber (Hevea brasiliensis (A. Juss.) M.A.), which had the major impact in the first half of this century. After the first commercial plantings in the 1900s, the area expanded rapidly, reaching a maximum in 1935, when nearly 40 per cent of Singapore’s total land area was under rubber plantations. After this, the area under cultivation declined sharply, except for a temporary increase in the production of food crops during the Japanese occupation (1942-45). The post-war era saw a decline in all crops except vegetables as agricultural land was increasingly lost to urbanisation and industrialisation. Today, less than 100 hectares are used for intensive vegetable cultivation while more than half the main island is urban in character.
Although heavily exploited for firewood, most of Singapore’s extensive mangrove forest area survived into the twentieth century. All but a few small areas have sub- sequently succumbed to conversion to brackish water ponds for agriculture, systematic reclamation for building and, more recently, the barraging of all major non-urban estuaries to create freshwater reservoirs (Corlett, 1987).
Conservation
In the early decades of the colony, exploitation and clearance of the forest ap- parently proceeded unchecked. In 1848, however, concern about possible effects on Singapore’s climate led the Governor to prohibit the further destruction of forest on the summits of hills. This prohibition seems to have been effective for Bukit Timah, at least (Corlett, 1988). By 1882, when Nathaniel Cantley was commissioned to survey the forest resources of the Straits Settlements (Singapore, Malacca, Province Wellesley and Penang), concern for the climatic effects of deforestation had largely been replaced by worries about the timber supply. Cantley, reporting that only 7 per cent of the original forest remained, proposed the creation of forest reserves (Cantley, 1884). His recommendations were accepted and, eventually, about 10 per cent of the island was protected in this way (Fig. 2). Unfortunately, most of the reserve area consisted of grassland, scrub or degraded mangrove, with little good forest. In addition to the forest reserves, an area around Singapore’s first reservoir (now called MacRitchie
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Angiosperm Flora of Singapore 1 9
Reservoir), partly covered in degraded primary forest, was protected during the 1890s as a catchment.
Most of the reserves were eventually worked for timber, handed over to squatters or otherwise developed. An area of primary forest survived at Changi until 1927, when it was cleared for construction of a military base. The decline in the forest reserves coincided with an increase in the protected catchment area as new reservoirs were constructed in the centre of the island. The expanded catchment area incorporated several fragments of disturbed primary forest, including what remained of Chan Chu Kang Forest Reserve and part of the Mandai Reserve, although this latter area was later cleared for the extension of Seletar Reservoir in 1940-41.
The Forest Reserves were finally abolished in 1936 but Bukit Timah and parts of the mangrove reserves at Pandan and Kranji were placed under the control of the Botanic Gardens. In 1951, these three areas, with the entire catchment area and 4 hectares of cliff face at Labrador, became Nature Reserves. The mangrove reserves were subsequently lost to development. Today the Nature Reserve system consists of 2,795 hectares in the centre of the island, of which 81 hectares is in Bukit Timah Nature Reserve and the rest in the Public Utilities Board Catchment Area (Fig. 2). A small area of mangrove at Sungei Buloh is protected as a bird sanctuary.
Climate
Singapore is only 137 km north of the equator and has an equatorial climate. The range of mean monthly temperatures is only 25.5-27.3 °C and of mean monthly rain- fall 160-300 mm. In Southeast Asia, similar climates are confined to the southern part of the Malay Peninsula, parts of Sumatra, much of Borneo and part of western Java. Elsewhere, only the island of New Guinea and parts of the central and western Amazon region have extensive areas of such climates. The botanical consequences of this extreme aseasonality are most obvious in urban areas, where the tree-lined streets are green all the year round but rarely show the massed flowering displays of other tropical cities. As discussed below, however, both seasonal and non-seasonal variations in the climate are of great significance for the native flora.
Despite its small size, Singapore also shows a surprising amount of spatial variation in rainfall. Mean annual rainfall exceeds 2,300 mm in the central part of the island and falls below 2,000 mm along much of the south coast (Chia and Foong, 1991). This spatial variation may have had a significant effect on plant distributions before the nineteenth century but its influence is now obscured by the effects of recent human impact.
A thorough review of Singapore’s climate with additional references can be found in Chia & Foong (1991). Here I will only discuss features of direct botanical relevance.
Seasonality
Despite its apparent constancy, Singapore’s climate is perceived by people, birds (Hails, 1987), and plants as seasonal. The time difference between the longest and shortest days of the year is only 9 minutes so photoperiod effects are unlikely, if not impossible. The long term means of air temperature, rainfall, relative humidity and solar radiation are clearly seasonal, although within a narrow range (Fig. 3). This limited seasonality is a consequence of the changes in the prevailing wind direction. The northeast monsoon prevails from November to March and the southwest (or, more accurately in Singapore, south) monsoon from May to October. During the inter- monsoon periods of April/May and October/November wind directions are variable.
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Angiosperm Flora of Singapore 1 1]
November and December are generally cooler, wetter and cloudier than other months. The solar radiation maximum is in February/March, which is also the period most prone to long dry spells. The month with the lowest mean rainfall, however, varies in different parts of the island. The hottest months are May and June. The most striking climatic change in most years is between the cloudy, wet period at the beginning of the northeast monsoon in November/December and the relatively drier and sunny period towards the end, in February/March (Fig. 3). The strength of this contrast is obscured in long-term averages because the precise timing varies from year to year but, in most years, it is the most significant interruption to the uniformity of the climate.
A single annual community-level flowering peak — around April — has been reported from several lowland forest sites in the Malay Peninsula (Ng, 1988), presumably triggered by this change. Corner (1988) suggests that there is often a second, weaker, flowering peak later in the year, triggered by a second, less predictable, dry period. There is no quantitative evidence for this but two annual peaks of leaf flushing have been described at some sites. Clear annual (e.g. Campnosperma auriculatum (Bl.) Hook. f.) or biannual (e.g. Fagraea fragrans Roxb.) flowering periodicities are fairly common in the Singapore flora.
The absence of regular seasonal climatic extremes must make non-seasonal extremes more easy for plants to detect. It is not surprising, therefore, that irregular climatic events that do not follow an annual cycle are of at least equal botanical significance in Singapore. Some of these climatic cues occur with a frequency greater than annual. The sudden drop in temperature caused by daytime thunderstorms is known to provide the trigger for already-formed flower buds to complete their development in certain orchid species (e.g. Dendrobium crumenatum Sw.) and the angsana tree (Prerocarpus indicus Willd.). Several other species which flower synchronously several times per year (e.g. Rhodamnia cinerea Jack, Timonius wallichianus (Korth.) Valeton) presumably have a climatic trigger for floral initiation.
Other climatic extremes occur with less than annual frequency. Many forest species show supra-annual reproductive cycles which may have climatic triggers. At irregular intervals of 2-10 years, the reproductive activity of many species coincides in a massive burst of flowering followed by an equally striking fruiting peak. The cue for this dramatic mass-flowering is certainly climatic but the precise event responsible is still uncertain. Dry weather (Foxworthy, 1932; Medway, 1972), increased daily sunshine hours (Ng, 1977), and a drop in minimum temperature (Ashton ef a/., 1988) have all been suggested. All three suggested triggers tend to occur together near the beginning of the year and may be associated with the El Nino-Southern Oscillation Event (Ashton et al., 1988). A mass-flowering event in Singapore in 1987 (Corlett, 1990) followed an exceptionally dry and sunny February, but there was no significant drop in minimum temperature. The consequences of this supra-annual pattern of forest phenology for the common, basically frugivorous, monkey, Macaca fascicularis, are described by Lucas & Corlett (1991).
Climatic extremes may act as constraints as well as triggers but this effect is not obvious in Singapore. The extreme minimum (19°C, in January 1934) and maximum (35.8°C, in April 1979) temperatures recorded on the island differ little from the annual extremes. Hailstorms are rare and the longest recorded dry spell was 32 con- secutive days in February and March, 1970. Singapore is well outside the typhoon belt but during the south monsoon short-lived squalls, known locally as “Sumatras’, can cause damage to isolated trees.
12 Gard. Bull. Sing. 44(1) (1992)
Microclimates
Few plants experience the “raw” climate recorded at standard meteorological stations. For the majority, climatic means and extremes are modified by the presence of other plants and non-living structures. The contrast between the microclimate near the forest floor and that above the canopy or in a large clearing is well-documented (Whitmore, 1990). Most of Singapore’s native flora must have been adapted to spending all or part of its life cycle in the cool, damp shade of the forest understorey. The only exceptions would have been canopy epiphytes, gap specialists and coastal species. Most of Singapore is now one huge, permanent clearing so the dominance of exotics and coastal species among both the planted and spontaneous flora is not surprising. Out- side the forest, there is also a marked rural-urban contrast in microclimate (Singapore Meteorological Service, 1986), attributed to heat stored in urban structures, but the botanical significance of this is unknown.
Geology and Soils
Geology
_The geology of Singapore has recently been reviewed by Thomas (1991) and a geological map of the island is available (PWD, 1975). Only a brief summary is given here.
The centre of the main island is underlain by the Bukit Timah Granite. This con- sists of acid igneous rocks, ranging from granodiorite to true granite, dated to the early-middle Triassic (230-205 Myrs B.P.). Other granites of similar or more recent age outcrop at Changi Point, Pulau Sekuda and Pulau Ubin. The Gombak Norite outcrops in a restricted area on the west of the Bukit Timah Granite, around Bukit Panjang and Bukit Gombak. It consists of basic igneous rocks ranging from norite to gabbro and is older than the Bukit Timah Granite. Most of the southern, south- western, and western part of the main island, and most of the southern islands, are underlain by a variety of sharply-folded sedimentary rocks termed the Jurong Forma- tion. These include conglomerates, sandstones, siltstones and mudstones. They are younger than the Bukit Timah Granite: probably of upper Triassic to early or middle Jurassic age (230-180 myrs B.P.).
Much of eastern Singapore and a part of the northwest is covered in semi- consolidated alluvial sands and gravels, with some silt and clay. This deposit, called the Old Alluvium, is predominantly granitic in provenance and seems to have been deposited in a braided river environment during the more seasonal climate of one or more of the Pleistocene cold stages (Gupta ef a/., 1987). Holocene deposits of various types cover low-lying coastal areas and fill the lower parts of river valleys. These include beach sands and gravels deposited during the Holocene sea-level maximum (ca. 5,000 B.P.) and a variety of other marine, estuarine and alluvial gravels, sands, muds and peats.
Soils
The soils of Singapore were classified into 24 series and mapped by Ives (1977) and have recently been reviewed by Rahman (1991). Unfortunately, soil classifications pro- duced by soil scientists, principally for agricultural purposes, often seem to have little predictive value for ecology, particularly in the tropics. This is probably because many ecologically important features of a soil are destroyed by cultivation. A soil can be cleared of its vegetation cover, suffer severe compaction, have its organic matter
Angiosperm Flora of Singapore 1 13
oxidised, its nutrients leached, and its surface layers eroded, without changing its posi- tion in the classification. Moreover, Singapore’s continuous high temperature and rain- fall, and the resulting intense weathering and leaching, have resulted in soils with similar properties on a range of different parent materials. Extreme parent materials, such as limestone and ultrabasic rocks, do not occur in Singapore.
Most soils in Singapore can be described as sandy clay loams and have a bimodal particle-size distribution. All are acidic, with a low cation exchange capacity and low to very low concentrations of available nutrients. All except those under the small areas of primary forest have undergone at least one agricultural cycle and much of Singapore is covered in soils which, whatever their taxonomic position, are character- ised by extreme soil degradation as a result of nineteenth and early twentieth century agricultural practices. Over large areas the original soil has been removed, buried or drastically altered by construction activity. A variety of different fill materials have been used for the extensive reclamation of land from the sea, including clayey sub- soil from inland construction projects and marine sand dredged from the sea bed. Sand fill resembles natural coastal deposits but clayey fill, after compaction to ensure stability, results in a soil that is completely structureless, poorly aerated and drained, and very low in fertility.
Six of the ten soil orders in the United States Department of Agriculture (USDA) Soil Taxonomy have been recognised in Singapore (Rahman, 1991). The largest area is covered in Ultisols. These are found on granite, on the fine- and mixed-grained sedimentary rocks of the Jurong Formation, and on the Old Alluvium. Soil develop- ment is greatest on well-drained, level ground over granite and least on the Old Allu- vium, where some of the less-developed soils may be better classified as Inceptisols or Entisols (Rahman 1991). Inceptisols are also found on some of the coarse-grained, resistant rocks of the Jurong Formation. Soils over granodiorite and the more basic igneous rocks typically develop into Oxisols. The soils on recent alluvium are mostly Entisols, although of widely varying properties. Other soil types of relatively minor extent occur near the coast, including highly organic Histosols formed under freshwater swamp forest, Spodosols (podzols) on old beach deposits, and the distinctive Sulfa- quents under mangrove.
Biogeography
This account of the biogeography of Singapore’s flora is based on the species list of Turner ef al. (1990), with distributional data from regional floras, principally the Flora Malesiana, Tree Flora of Malaya, Flora of Thailand, Flore du Laos, du Cam- bodge et du Vietnam, and the Flora of Australia, supplemented, where possible, by data from more recent monographs. This data has a number of obvious limitations, apart from its incomplete taxonomic coverage. Most serious is the undercollection of much of the region around Singapore, particularly Riau, Sumatra and Kalimantan. There is also the problem of non-coincidence of biogeographical and political bound- aries. In the region under consideration here, the clearest examples are the Malaysia- Thailand border, which is somewhat south of the northern limits of much of the Malesian flora, and the island of Palawan, which is biogeographically Bornean but politically part of the Philippines. Thus “Thailand” or “the Philippines”, unqualified, in a description of a species distribution, are not very helpful to the biogeographer.
Takhtajan (1986) places Singapore in the Malesian Subregion of the Malesian Region of the Paleotropical Kingdom. The Malesian Subregion is further divided into five provinces, with Singapore in the Malay Province. However, the suggested differences between the Malay Province and the adjacent Sumatran and Kalimantan Provinces
14 Gard. Bull. Sing. 44(1) (1992)
are minor. Indeed, at the species level, the most striking feature of Singapore’s flora is its broad distribution. The question of possible Singapore endemics is probably best left until after this Flora is completed, but they are undoubtedly very few, if any. Even among the inland rain forest flora — the plants most likely to have restricted distribu- tions — only 15 per cent of the 730 species for which I could obtain reasonable data are apparently confined to the Malay peninsula (including peninsular Thailand and, in a few cases, peninsular Burma or the Riau Archipelago). A further 50 per cent of forest species occur more widely in the everwet “core” of Sundaland (Sumatra, Borneo and West Java). The remaining third of the species are even more widespread, extend- ing northwards into continental Asia (21 per cent), or eastwards through Malesia towards Australia (14 per cent). The wide distribution of most Singapore forest species is no doubt the result of the unexceptional nature of the physical environment and Singapore’s position near the centre of the Sunda shelf. It does suggest, however, that whatever the effects of full-glacial aridity on the vegetation of the region, continuous rain forest was re-established on Sundaland before rising sea-levels created major barriers to dispersal.
It is important to point out that the Singapore populations of widespread species are still of conservation value because of the likelihood of ecotypic variation at the margins of the range. Moreover, deforestation to the north and south of Singapore is rapidly restricting the range of species that were widespread ten or twenty years ago.
As might be expected, the coastal flora is much more widely distributed. Caesalpinia bonduc Roxb. and Thespesia populnea (L.) Correa are effectively pantropical while a number of species range from East Africa to the western Pacific (e.g. Bruguiera gymnorrhiza (L.) Lamk., Excoecaria agallocha L.). These extremely wide ranges pre- sumably reflect both the tendency to seawater dispersal in the coastal flora and the relative uniformity of the coastal environment.
The flora of man-made open sites is essentially pantropical, consisting of species of Asian origin which have now spread around the world, and exotics from Africa and the Neotropics that have become naturalised in Singapore. As 1s true throughout the region, the recognizably exotic component of Singapore’s flora contains many more species of Neotropical origin (at least 84 species) than from Africa (14 species) (Corlett, 1988). This may reflect similarities in climate and, perhaps, agricultural systems, as well as the large number of crop and ornamental plants deliberately introduced from tropical America. Another interesting feature of Singapore’s weed flora is the rarity or absence of several pantropical exotics, abundantly-naturalised in the more seasonal climates of the region (e.g. Crassocephalum crepidiodes (Benth.) S. Moore, represented only by a single old record) and the absence of weeds of temperate origin (Corlett, 1992b). The composition of the weed flora demonstrates clearly that, far from being a “greenhouse climate” in which anything can grow, Singapore’s year-round high temperatures and rainfall exclude unadapted species as effectively as extreme cold or drought.
The biogeography of Singapore’s flora at the genus and family levels is that of the Sunda Shelf flora as a whole. The dual origin of the regional flora from both Gondwanic and Laurasian sources has long been recognised. However, the complex geological origin of the Malay Archipelago, outlined above, probably provided a multiplicity of times and routes for interchange, making the recognition of distinct “elements” in the flora difficult. For example, the family Dipterocarpaceae shows an overwhelming concentration of living species in West Malesia (Sundaland plus the Philippines) and probably entered the region from the Asian mainland, but the global distribution of living and fossil members suggests a possible ultimate origin on Gond- wanaland (Ashton, 1982). Plants of originally Gondwanic families have probably
Angiosperm Flora of Singapore 1 15
entered the region from both the south-east (after the Miocene collision between Laurasia and Gondwanaland or, earlier, via island “stepping-stones”) and from the north-west (after being “rafted” northwards from Gondwanaland on India). Some taxa (e.g. the palms, of disputed ultimate geographic origin, and the Loranthaceae) have apparently entered the region from both ends (Dransfield, 1987; Barlow, 1990).
Vegetation
Primeval Vegetation
I have been unable to find any useful description of Singapore’s vegetation before the late nineteenth century. Early maps and written accounts make insufficient distinc- tion between vegetation types or only refer to small areas. It is a reasonable assump- tion, however, that, except for sandy beaches and steep cliffs, and the immediate vicinity of the major settlement, closed canopy forest covered the whole island in 1819. To what extent this forest had been exploited, disturbed or cleared in the past is impossible now to determine. From topography, soil patterns and late nineteenth cen- tury maps, I estimate that mangrove forest would have occupied about 13 per cent of the main island, freshwater swamp forests of various types 5 per cent, and lowland evergreen rain forest the remaining 82 per cent (Corlett, 1991a). The floristic composi- tion of the rain forest must have varied considerably with soil type and topography but extensive botanical collection did not start until the 1880s, when more than 90 per cent of the forest had gone, so we have little information on this variation. The distinc- tive floras of the 10 ha forest remnant at Changi, cleared in 1927, and the 4 ha Gardens’ Jungle, now badly degraded, suggest that much of Singapore’s primeval flora may have been lost before it was collected. Even in the 1890s, Ridley could collect rain forest taxa at many sites where forest no longer exists. Known extinctions are mostly coastal, reflecting the complete destruction of the coastal forest, with the ex- ception of some small areas of mangrove.
Primary Forest Today
Today, primary rain forest, disturbed to varying extents, is confined to the 71 ha Bukit Timah Nature Reserve (which is about two-thirds primary forest) and scattered patches of various sizes, totalling about 50 ha, in the adjacent water catchment area. Most of these primary forest remnants are in the areas of overlap between the Forest Reserves established in the late nineteenth century and the current Nature Reserve system (Fig. 2). The Bukit Timah forest has apparently never been legally exploited, at least since its first protection in the 1840s (Corlett, 19885). Extensive illegal cutting of timber and firewood has, however, undoubtedly occurred at times of reduced pro- tection. At least 840 angiosperm species (excluding non-forest weeds) have been recorded from Bukit Timah in the past century (Corlett, 1990, 1991). Five families — Rubiaceae, Euphorbiaceae, Orchidaceae, Moraceae and Annonaceae — account for almost a third of the angiosperm flora. In terms of numbers of species, the Euphor- biaceae, Rubiaceae, Myrtaceae, Annonaceae and Lauraceae are the most important tree families, but the Dipterocarpaceae provides the greatest number of large tree in- dividuals (Wong, 1987). The Rubiaceae and Palmae are the biggest climber families, ferns dominate the herb layer, and ferns and orchids account for most of the epiphytes (Corlett, 1990).
The largest primary forest remnants in the catchment area are around MacRitchie Reservoir and the Nee Soon (Yishun) firing ranges. Both areas have been protected
16 Gard. Bull. Sing. 44(1) (1992)
since the late nineteenth century (as a catchment area and Chan Chu Kang forest reserve, respectively (Fig. 2)) but, before that, were probably exploited heavily. Con- tinuity of forest cover — and thus justification for considering them “primary” — is shown by the extremely rich flora, including numerous species absent from the adjacent secondary forests. The Nee Soon area also includes about 15 ha of dis- turbed freshwater swamp forest, the last remnant of the much larger area studied by Corner (1978). The floristics of these other primary forest remnants have not been investigated as thoroughly as Bukit Timah, but the floras seem to be to some extent complementary.
Secondary Forest and Scrub
The rest of the central catchment area (Fig. 2) is covered in secondary forest of various ages (Corlett, 1991c). This area was cleared of its original forest cover by the mid nineteenth century, cultivated until exhaustion and then abandoned to lalang. Protection as part of an expanded water catchment, mostly in the period 1899-1906, did not lead to an immediate regeneration of forest because of frequent grass fires, but most of the area seems to have been under woody cover by the nineteen-thirties. Some parts have been cut or burned more recently. The oldest areas of forest (?50-80 years old) are 15-20 m tall with 35-65 species >2 cm d.b.h. in 0.1 ha plots. This tall secondary forest is dominated by members of the families Guttiferae (Calophyllum spp., Garcinia spp.), Lauraceae (Lindera lucida (Bl.) Boerl., Litsea spp.), Myrtaceae (Eugenia spp., Rhodamnia cinerea Jack), and Elaeocarpaceae (Elaeocarpus spp.). It is clearly distinguished from the included primary forest remnants by its lower stature and species diversity, even canopy, poorly-developed understorey, and the complete absence of Dipterocarpaceae and other species with large, wind-dispersed seeds.
For historical reasons, tall secondary forest is confined to the central catchment area, but areas of younger secondary forest and scrub, probably all less than 40 years old, are scattered around the main island and on several offshore islands (Corlett, 1991c). This pioneer community is remarkably uniform, floristically, despite the wide range of rock types on which it occurs. Large areas contain less than 20 vascular plant species in total. The explanation for this relative floristic poverty must be severe soil degradation — chemical, physical or both. The dominant species is usually Adinandra dumosa Jack, particularly after the formation of a closed canopy has eliminated the smaller species. Transitions between the low (6-12 m), Adinandra-dominated forest and the tall secondary forest described above can be found in some parts of the central catchment area. Whether or not the outlying areas, if protected, will ever undergo this transition, in the absence of nearby seed sources, is an interesting question!
Except for small areas at the back of beaches, all herbaceous vegetation in Singapore | is secondary and results from recent or continued disturbance. Fire prevention and control have virtually eliminated the vast areas of lalang (Jmperata) grassland which dominated the Singapore landscape in the late nineteenth and early twentieth century. Spontaneous herbaceous vegetation is now most extensive on wasteland awaiting development and on land newly reclaimed from the sea. Except on the poorest soils (such as land reclaimed with sand fill), this wasteland vegetation is dominated by naturalised exotic species, particularly African grasses and tropical American legumes (Corlett, 1988a@). Exotics also dominate the weed flora of parks, gardens and other managed vegetation.
The surviving fragments of Singapore’s once extensive mangrove forests all show the effects of past exploitation and disturbance (Corlett, 1987). There are few old trees and the inland margins of most patches have been destroyed by reclamation.
Angiosperm Flora of Singapore 1 17
Moreover, land reclamation, barraging of estuaries, and developments inland have drastically changed the patterns of sediment deposition and erosion along Singapore’s coastline, so even protected mangrove areas are unstable. The rich epiphytic flora recorded earlier this century has almost entirely disappeared but most of the woody flora survives.
Managed Vegetation
For most visitors to Singapore, the lasting botanical impression is not the untamed exuberance of tropical nature but the orderly rows of matching trees and neat expanses of close-mown grass. This impression has been achieved at considerable expense in money and labour, and only after many years of careful planning and experimentation (Corlett, 1992c). Although the final image is very Singaporean, the flora of the managed vegetation is pantropical. A continued programme of plant introductions and field trials has resulted in an exceptionally diverse park and roadside flora, with the plantings often dateable from a knowledge of past changes in the favoured tree species. The few native species that are widely planted are all coastal in origin: examples include the Pong Pong (Cerbera odollam Gaertn.), the Sea Apple (Eugenia grandis Wight), the Yellow Flame (Pe/tophorum pterocarpum (DC.) Heyne), the Sea Almond (Jerminalia catappa L.), and the near-native Angsana (Pterocarpus indicus Willd.). Many early introductions were from tropical America, such as the Rain Tree (Samanea saman Merr.) and the Broad-leaved Mahogany (Swietenia macrophylla King). More recently, African Mahoganies (Khaya spp.) have been very widely planted, but no major tropical area is unrepresented.
Much of the diversity in the planted shrub flora is at the infraspecific level, with Bougainvillea (Bougainvillea X buttiana Holttum & Standley), Hibiscus (Hibiscus rosa-sinensis L.) and others represented by numerous cultivars. Managed grasslands, in contrast, are largely planted with one species, Axonopus compressus (Swartz) Beauv., propagated vegetatively and now probably the commonest plant in Singapore. Fine lawns, particularly in private gardens, are usually planted with species of Zoysia, while several cultivars of Bermuda Grass (Cynodon dactylon (L.) Pers. and hybrids) are used on golf courses.
Conclusions
Despite its small size, relatively uniform topography and recent history of massive human impact, Singapore still supports a vascular plant flora of incredible diversity. The majority of this diversity, however, is dependent on the protection of a few, small areas: most importantly, the primary forest remnants at Bukit Timah, MacRitchie and Nee Soon. Even the oldest secondary forest in the central catchment area is depauperate in comparison and its conservation significance lies more in its role as a buffer for the primary forest remnants, its importance as a habitat for vertebrates, and its potential for future floristic enrichment, by natural processes or with human inter- vention. Along the coast, the few remaining patches of mangrove forest, although of low floristic diversity, support an extremely rich and interesting fauna. Outside the forest, the flora, both spontaneous and planted, consists largely of pantropical species which, although often of considerable biological interest and aesthetic value, have no particular conservation importance.
The Flora of Singapore project is of major significance for plant conservation in Singapore. The availability of a modern Flora will greatly facilitate the detailed studies of species and habitats on which long-term conservation management of the flora will
18 Gard. Bull. Sing. 44(1) (1992)
ultimately depend. Singapore, with its stable Government, strong economy, high educa- tion standards and well-deserved reputation for long-range planning, can and should set an example for the rest of the tropical world.
Acknowledgements
Among the many people who helped in the preparation of this paper and, earlier, during my 5 year stay in Singapore, I particularly wish to thank Dr. HW. Tan, Prof. H. Keng, Mr Samsuri, Dr. PW. Lucas, Mr D.H. Murphy, Dr. Y.C. Wee and Dr. A. Gupta.
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Five New Species of Didymocarpus (Gesneriaceae) from Peninsular Malaysia
R. KIEW Department of Biology Universiti Pertanian Malaysia 43400 Serdang Selangor, Malaysia
EFFECTIVE PUBLICATION DATE: 15 MAR 1993
Abstract
Five new species of Didymocarpus are described from Peninsular Malaysia. Two, D. anthonyi Kiew and D. leiophyllus Kiew, are from the east coast and belong to sect. Boeopsis; D. Jeucanthus Kiew is from the foothills of Selangor; D. stoloniferus Kiew from Gunung Ulu Kali, Pahang; and D. salicinoides Kiew, from Trengganu and southern Kelantan, is raised to specific rank having previously been described as D. salicinus var. major Ridley. The sections Boeopsis and Salicini are defined and keys to their species provided. Didymocarpus lithophilus Kiew is validated.
Introduction
In common with several other large families of herbaceous plants, such as the Begoniaceae and Orchidaceae, the Gesneriaceae exhibits a high level of endemism (more than 90 per cent) in Peninsular Malaysia. Of the 85 odd species of Didymocarpus presently described, only D. crinitus Jack, D. platypus C.B.Cl. and D. reptans Jack (and possibly D. fasciatus Ridley) occur outside Peninsular Malaysia and southern Thailand. Within the peninsula, the majority of species is not widespread and 70 per cent are known from a single locality (Kiew, 1991).
Recent exploration of areas that are still botanically little known, such as the east coast, has led to the discovery of several new species. Two described here belong to sect. Boeopsis, a section which has a south-easterly distribution. However, even rela- tively well-known areas, such as Gunung Ulu Kali, may harbour undescribed species.
The last complete account of Malayan Didymocarpus is that in Ridley’s flora (1923). There he recognised six sections based largely on characters of habit, inflorescence and floral morphology. These sections have remained broadly useful although several contain anomalous species. Sect. Elati (sect. Eudidymocarpus of Ridley) has been redefined by Weber & Burtt (1983). Section Didymanthus contains a hotchpotch of species including the anomalous D. parviflorus and D. leucanthus (see below), D. falcatus and D. flavobrunneus, which belong to another alliance (Burtt, 1990) and a group of species with large bracts (to be excluded from sect. Didymanthus and placed in a new section based on D. venustus). Section Reptantes and sect. Heteroboea remain as Ridley circumscribed them. Section Pectinati has been redefined to exclude D. densifolius (Kiew, 1987), which is now placed in sect. Salicini (see below). Section Boeopsis is here redefined. In addition, the genus Codonoboea is now reduced to sec- tional level in Didymocarpus (Kiew, 1990). There remain several species that do not fit comfortably into any of these sections, such as D. caelestis (Ridley) Kiew, D. cordatus Jack, D. geitleri Weber, D. leucocodon (Ridley) Kiew, D. longipes C.B.Cl., D. primulinus
23
24 Gard. Bull. Sing. 44(1) (1992)
Ridley, and D. violascens Ridley. Until details of floral morphology of the majority of species are better known, not only for the peninsular species but also for species throughout the geographic range of the genus, a fundamental revision of the sections will not be possible.
Descriptions of New Species
1. Didymocarpus anthonyi Kiew sp. nov.
Differt a Didymocarpo heterophyllo Ridley statura majore, foliis longioribus, lamina in petiolum decrescente et floribus majoribus.
Typus: Trengganu, Ulu Besut R. Kiew RK 2700 (holo UPM; iso K, SING).
Erect, unbranched plant, stem woody to 37 cm by 6 mm thick, flowering at 9 cm tall. Indumentum of stem apex and petioles deep brown, densely matted with long uniseriate hairs. Leaves in a tuft at top of stem, upper internodes crowded, lower inter- nodes to 5 mm apart. Lamina oblanceolate, (12.5-) 16 (-19) by 4-4.3 cm, narrowed to apex, base narrowly cuneate, sometimes unequal, glabrous above and beneath, in life deep green above and light green beneath, drying thinly leathery. Margin in the upper half of the leaf finely and distantly serrate with a tuft of hairs in the notch, in lower half + entire, marginal strip hairy beneath. Secondary veins 11-16 pairs, midrib and secondary veins plane but conspicuous above, prominent beneath and sparsely hairy, arching close to margin, tertiary veins obscure above, prominent beneath. Petiole 1-2 cm long in youngest leaves elongating to 1.7-2.5 cm in older leaves, grooved above, geniculate abaxially.
Inflorescence 4-flowered cyme, several per axil. Peduncle 6-8.5 (-11 cm), erect with flowers held above leaves, brownish-purple, pedicel 10-17 mm. Bract pair ligulate, 6 by 1.5 mm long. Indumentum of peduncle, pedicel, bracts and calyx sparse with appressed, long, multicellular eglandular hairs with fewer short glandular hairs. Flowers nodding. Calyx divided almost to base, lobes narrowly acute, 2-3 by 1 mm. Corolla broadly campanulate, tube 5-6 by 7-8 mm, white, minutely pubescent outside, lobes broadly oblong, apex rounded, upper two 4-6 by 6 mm, pale to deep purple, reflexed, lower three 5 by 5-6 mm, white suffused purple, projecting 6-10 mm beyond upper. Stamens with thick fleshy filament, c. 2.5 mm long, joined to base of corolla tube, anthers white, broadly sagittate, 3 by 2 mm, connivent at apex. Ovary ellipsoid, 3 by 1.5 mm, purplish red, style whitish-green, 5 mm long and projecting beyond corolla tube, ovary and style minutely pubescent, stigma minute, rounded, c. 0.3 mm across, white, glistening, apex papillose. Disc none. Capsule narrowly ellipsoid, slightly curved, 3 cm long, c. 1.5 mm thick, + glabrous.
Distribution: Endemic to Peninsular Malaysia — Trengganu, Ulu Besut. Habitat: Hill slope, common on earth banks.
Specimens examined: Trengganu — Ulu Besut: Kg. Keruak 1 September 1986 S. Anthony SA 675 (SING, UPM); Kg. La 7 May 1988 R. Kiew RK 2700 (K, SING, UPM).
Notes: By virtue of its broadly campanulate, purple corolla and large, shortly stalked anthers, this species belongs to sect. Boeopsis. However, it is the most robust species in this section with stems that attain 37 cm in height. In its leaves, which are glabrous above, and in its simple cymes, it resembles D. heterophyllus Ridley, from which it is readily distinguished not only by its greater height, but also by its larger leaves, which are strongly narrowed to the base (in D. heterophyllus the leaf base is rounded), and in its larger flowers (Table 1).
jn
Five New Species of Didymocarpus 25
Table 1 Diagnostic differences between Didymocarpus anthonyi, D. heterophyllus and D. leiophyllus
Character D. anthonyi D. heterophyllus D. leiophyllus Stem height (cm) 9-37 0-7 2-19 Lamina length (cm) 12.5-19 5-13.5 7-11 Lamina width (cm) 4-43 2-4 2.5-4 Lamina base narrowed rounded narrowed Leaf margin distantly serrate crenulate + entire Petiole length (cm) 1-2.5 0.5-3 1-2
No. flowers/inflorescence 4 3-4
Calyx length (mm) 2-3 1-2 4
Corolla length (mm) 9-12 3-4 1]
This species is named for S. Anthonysamy, herbarium assistant in the Department of Biology, Universiti Pertanian Malaysia, who is an excellent field botanist and who made the first collection of this species.
Among species of Didymocarpus the indumentum of the ovary and style is variable both with regard to trichome type and their density. The difference between a finely pubescent indumentum of long-stalked trichomes and a pustulate indumentum con- sisting of short-stalked glandular ones is clearly discernible with the naked eye.
The range of stylar indumentum is illustrated in Plate 1. All species examined possess short-stalked glandular hairs, either with a single-celled rounded head (D. corneri Pl. le; D. geitleri Pl. 1g; D. yongii Pl. 1d) or with a 2-celled head (D. leucanthus Pl. la; D. leucocodon P\. 1b) or with a 4-celled head (D. anthonyi PI. lc). Except for D. yongii (Pl. 1d), they possess in addition short, 2 or 3-celled eglandular hairs on a raised base. Those of D. anthonyi are exceptionally short. D. leucanthus, D. quinquevulnerus (P|. 1h) and D. platypus possess a third type, long-stalked glandular hairs.
The density of stylar trichomes varies from extremely sparse (the style of D. anthonyi is almost glabrous), moderately sparse (D. leucanthus, D. quinquevulnerus and D. platypus) to, in most cases, dense (D. corneri, D. geitleri, D. leucanthus, D. leucocodon, and D. yongii).
Even with this small sample, trichome type is not apparently related to taxonomic affinity. Within sect. Boeopsis stylar trichome type differs among species: D. anthonyi has long eglandular and glandular hairs (the latter with a 4-celled head); D. yongii has only glandular hairs and these have a single-celled head. In addition, long-stalked glandular hairs are found in species in sect. Didymanthus (D. /eucanthus) and in sect. Heteroboea (D. quinquevulnerus) and short, glandular trichomes with a rounded head are found in sect. Codonoboea (D. corneri) and sect. Boeopsis (D. yongii).
In species where the style projects beyond the corolla tube (D. anthonyi, D. corneri and D. geitleri), the predominant trichome type is short-stalked glandular trichomes and, in contrast, eglandular hairs are extremely sparse. It is tempting to suggest that these glandular hairs function to secrete substances that attract the pollinator either by scent (none of these species has a scent perceptible to the human nose) or by sight. The indumentum is glistening in all species and in some species contrasts in colour
Plate 1. Stylar trichomes in Didymocarpus. a. D. leucanthus; b. D. leucocodon; ¢. D. anthonyi; d. D. yongii.
with the white corolla. It is pale fawn in D. corneri and magenta-purple in D. geitleri. Weber (1989) recorded the indumentum of the latter species as orange. In contrast, the white style of D. anthonyi is almost glabrous.
Long-stalked glandular hairs are found in those species which have a long corolla tube which includes the style, namely D. leucanthus and D. quinquevulnerus. It is possible, therefore, that trichome type is related to pollinator guild rather than to taxonomic affinity.
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Plate 1. Stylar trichomes in Didymocarpus (cont.). e. D. corneri (below stigma); f. D. corneri (above ovary); g. D. geitleri; h. D. quinquevulnerus.
In general, the indumentum of the ovary is more dense than that of the style (with the exception of D. leucocodon (P1. 2b) where it is more dense on the style). In some cases trichome type is the same on the style and ovary, although the hairs may be longer (e.g. D. leucanthus P|. 2a, D. anthonyi Pl. 2c) or shorter (e.g. D. leucocodon) on the ovary. In other species, (D. geitleri Pl. 2f, D. corneri Pl. 2e and D. yongii Pl. 2d) trichome type on the style and ovary is different because short-stalked glan- dular trichomes with a rounded head are absent from the ovary. In D. yongii, in addition to eglandular hairs, there are long-stalked glandular hairs on the ovary. These latter species have a transitional zone in the lower part of the style where all types of trichome are present. This is seen in D. corneri (Pl. 1f) where three trichomes types are present. The transitional zone in D. geitleri is illustrated by Weber (1989).
Plate 2. Trichomes of the ovary of Didymocarpus. a. D. leucanthus; b. D. leucocodon; ¢. D. anthonyi; d. D. yongii.
In all species examined (except for D. quinquevulnerus, Pl. 2h), there is a pre- ponderance of short eglandular hairs, which suggests they may play a protective role in the immature ovary.
2. Didymocarpus leiophyllus Kiew sp. nov. Plate 3
Differt a Didymocarpo heterophyllo Ridley inflorescentia uniflora, foliis fere integerrimis et floribus majoribus.
Typus : Trengganu, Ulu Setui R. Kiew RK 2265 (holo UPM; iso SING). Erect, unbranched plant, stem woody to 19 cm and 3-4 mm thick, flowering at
.
; Pi P a o; Ses
Plate 2. Trichomes of the ovary of Didymocarpus (cont.). e. D. corneri; f. D. geitleri; g. D. platypus; h. D. quinquevulnerus.
2 cm tall. Leaves forming a tuft at the top of the stem, lower leaves to 2.5 cm apart, spirally arranged. Lamina oblanceolate, 7-11 by 2.5-4 cm, apex acute or sometimes shortly acuminate, narrowed to base, unequal, glabrous above and beneath, in life dark green above, paler beneath, fleshy drying thinly leathery. Margin + entire, some- times shallowly dentate towards apex, marginal strip hairy beneath. Midrib prominent above and beneath, secondary veins 8-14 pairs, plane above, prominent beneath, arching close to margin, tertiary veins + obscure beneath. Petiole 1 (-2) cm. Indumentum
30 Gard. Bull. Sing. 44(1) (1992)
of stem apex, petiole, and lower surface of midrib and secondary veins of appressed uniseriate hairs.
Inflorescence 1-flowered. Peduncle and pedicel slender, purple-red, minutely pubes- cent, erect, 3.5-6 cm long with flowers held above leaves. Bract pair ligulate, 1.5-2 mm long. Calyx divided almost to base, lobes narrowly acute, 4 by 1 mm, purple-red, pubescent. Corolla broadly campanulate, tube 6 by 5 mm, white or bluish purple, minutely pubescent outside, lobes broadly oblong, apex slightly rounded, 5 by 4-5 mm, pale lilac, upper lobes reflexed. Stamens with filaments 1.5-2 mm long, broadening to the base and joined to base of corolla tube, anthers broadly sagittate, 2 by 1 mm, connivent. Ovary narrowly ovoid, 5 by 1.3 mm, style 5 mm long, yellowish, minutely hairy and projecting beyond the corolla tube, stigma slightly discoid + 1 mm across. Disc encircling base of ovary, c. 0.5 mm tall, deeply lobed. Capsule narrowly ovoid, slightly curved, 1.5-3.3 cm, purple, minutely hairy.
Distribution: Endemic to Peninsular Malaysia — Trengganu, Ulu Setui.
Habitat: Locally common in lowland forest on slopes above river banks or on steep earth banks.
Specimens examined: Trengganu — Ulu Sg. Setui 28 April 1986 R. Kiew RK 2265 (UPM, SING), 29 April 1986 RK 2272 (UPM), 30 August 1986 S. Anthony SA 662 (UPM), 31 August 1986 SA 670 (UPM), 5 November 1986 SA 7/8 (UPM).
Plate 3. | Didymocarpus leiophyllus.
Five New Species of Didymocarpus 3]
Notes: D. leiophyllus (P\. 3) is a distinctive species in its smooth leaf surface (the veins are not impressed above) and in its almost entire leaves. It belongs to sect. Boeopsis. In its flower it most resembles D. anthonyi in size (both have a larger flower than D. heterophyllus) and in colour (they both have a white or very pale purple corolla tube and lobes which are a deeper purple compared with D. heterophyllus where the corolla tube and lobes are uniformly mid-purple). Leaves of both D. leiophyilus and D. anthonyi are rather fleshy and have a pronounced pubescent marginal strip. D. leiophyllus differs from both D. anthonyi and D. heterophyllus in its single-flowered inflorescence (Table 1). Some populations of D. puncticulatus also have single-flowered inflorescences but D. /eiophyllus would not be confused with it as D. puncticulatus has hairy leaves which are punctate above and which frequently have a paler band down the centre.
3. Didymocarpus leucanthus Kiew sp. nov.
Didymocarpo parvifloro Ridley affinis sed foliis margine crenaturis praeditis supra pubescentibus et floribus albis differt.
Typus: Selangor, Ulu Ampang R. Kiew RK 2767 (holo, UPM, iso SING).
Stem prostrate with erect branching woody shoots to 1 m tall and 3 mm thick, deep purple in life. Leaves opposite, equal-sized, distant up to 7 cm apart. Lamina lanceolate, 12-13.5 by 5-5.5 cm, apex acute or sometimes acuminate, base cuneate, in dried state chartaceous, margin minutely crenate. In life dark green above and pale beneath. Lateral veins c. 9 pairs and ascending towards margin, sometimes with a minor vein parallel to lateral veins but petering out midway to margin, lateral veins and midrib plane above, prominent beneath, tertiary veins obscure above and below. Indumentum of short uniseriate hairs with c. 4 cells, on stem and petiole dense and appressed, lamina silky above and roughly pubescent beneath, with both long and short hairs. Lamina minutely pustulate beneath. Petiole terete, 1.5 to 3 cm long, deep purple in life.
Inflorescence axillary, 1-flowered, produced in a series so that axils bear buds, flowers and fruits at the same time. Peduncle and pedicel slender, 6-8 mm long, lengthening to 12 mm in fruit. Indumentum of peduncle, pedicel, bracts and calyx dense consisting of minute glandular hairs. Bracts ligulate 2 mm long. Calyx divided to base, lobes ligulate, 2 by 1 mm. Corolla narrowly tubular, white with a yellow spot at base of tube, tube 9 by 4-5 mm, minutely pubescent outside, lobes 5, oblong, apex broadly rounded, upper lobes 3 by 2.5 mm, reflexed, minutely glandular hairs on inner surface, lower lobes 3 by 4 mm, projecting 5 mm beyond upper lobes. Stamens 2, filaments slender 4 m long, anthers ellipsoid, 1.5 by 0.5 mm, connivent. Ovary narrowly cylindric 4 mm long, style enclosed within tube 3 mm long, densely pubescent, stigma rounded, 1 mm across, papillose. Disc 1 mm tall, subtending lower half of base of ovary. Capsule 3.5-4 cm long and 1 mm wide, densely pubescent.
Distribution: Endemic to Peninsular Malaysia — Selangor, Ulu Ampang.
Habitat: In lowland forest at c. 100 m, locally common and forming clumps on earth banks.
Specimens examined: Selangor, Ulu Ampang at Ampang Impounding Reservoir: 20 May 1984 R. Kiew RK 1307 (UPM); 16 August 1988 RK 2767 (UPM, SING).
Notes: Didymocarpus leucanthus most resembles D. parviflorus Ridley in its habit (it is decumbent producing erect, branching stems), in its long-petioled leaves and in its small, tubular corollas. It differs from D. parviflorus, which is a smaller, more or less
32 Gard. Bull. Sing. 44(1) (1992)
prostrate plant and which has smaller leaves (25-50 by 6-13 mm) with entire margins, yellow flowers and shorter fruits (c. 1 cm long).
Ridley (1905, 1923) included D. parviflorus in sect. Didymanthus, presumably as it has distant pairs of leaves. However, both D. parviflorus and D. leucanthus differ from other members of this section by their smaller, tubular flowers, which are not produced on long-peduncled cymes. These two species should therefore be excluded from sect. Didymanthus as it is presently circumscribed. However, until the Malayan species are better known, it is premature to erect a new section for them. For example, the little- known D. flavescens Ridley is similar to these two species in its small tubular, yellowish- white flowers on single-flowered inflorescences, but it is conspicuously different in its long peduncles, which are 2.5-7 cm long.
Among Malayan species of Didymocarpus, D. leucanthus (Pl. 4a) is unusual in possessing a nectary that does not completely surround the base of the ovary. The most common type of nectary in the genus is cylindrical and relatively large. In sect. Heteroboea, nectaries range from 0.7 mm tall (D. platypus, Pl. 4e) to 0.9 mm (D. quinquevulnerus, P|. 4f) to 1.5 mm tall in D. polyanthoides. This type of nectary is also seen in most other sections, e.g. in sect. Didymanthus (D. parvifolius), in sect. Codonoboea (D. corneri, Pl. 4d) and in sect. Boeopsis (D. yongii, Pl. 4c). That of D. yongii is unusual in being distinctly lobed.
~ Nectary morphology is not always a reliable indicator of taxonomic affinity, although all species in the D. falcatus-D. flavobrunneus-D. pyroliflorus alliance have unilateral nectaries (Weber, 1989). D. leucanthus also has a unilateral nectary but is not at all related to this alliance. Some closely related species have different nectary types, such as D. parvifolius and D. leucanthus (D. parvifolius has a cylindrical nectary c. 0.5 mm tall and D. leucanthus has a unilateral one) and D. anthonyi and D. leiophyllus (the former species does not have a nectary and in the latter the nectary is cylindrical and lobed).
Flowers of a few Didymocarpus species do not have a nectary (e.g. D. anthonyi and D. codonion) or have a very small one (D. geitleri, Weber, 1989; D. leucocodon, Pl. 4b). Weber has described the features of pollen flowers in D. geitleri and pointed out that the evolution from nectar to pollen as a floral reward has occurred in several genera of the Gesneriaceae. In the pollen flower, not only is the nectary reduced in size but the anthers are large and conspicuous in the gaping mouth of the corolla, which has a short tube. In Didymocarpus this type of flower often has a style which projects beyond the mouth of the corolla. This flower type is seen in sect. Boeopsis, in sect. Salicini and in sect. Codonoboea (D. corneri). However, in sect. Boeopsis nec- tary size ranges from large (D. yongii) to absent (D. anthonyi and D. codonion). In D. yongii the style is not exserted as it is in D. anthonyi. Some species, such as D. leiophyllus, while having an exserted style, also have a nectary.
That pollen flowers have evolved several times in the Gesneriaceae and probably also within a large genus such as Didymocarpus means that the position of D. geitleri should be reconsidered, especially as its leaf morphology and indumentum is more typical of sect. Heteroboea than sect. Boeopsis, with which Weber (1989) suggested it was allied.
There is still a great deal to be learnt about pollination in Didymocarpus. Although many Didymocarpus species produce striking flowers often in abundance (a single plant of D. quinquevulnerus Ridley can have up to 40 flowers open at any one time, although 20-30 are more usual), it is an extremely rare event to see an insect visitor. I have only observed pollination in one species, D. robinsonii Ridley, where on Gunung Tahan its flowers were visited by bumblebees. It is probable that various types of bees visit the large, trumpet-shaped flowers that have conspicuous nectar guides. For those
18 4
AP
Nectary morphology in Didymocarpus
Plate 4.
ypus
D. plat
D. leucocodon; ¢. D. yongii; d. D. corneri; e. Ss
b.
inquevulneru
. ,’
leucanthus
a. D. f. D. qu
34 Gard. Bull. Sing. 44(1) (1992)
species with narrow-tubed flowers, such as D. leucanthus, or small purple flowers of sect. Boeopsis and sect. Salicini, which do not have conspicuous nectar guides, or small white flowers with exserted styles found in D. corneri, D. geitleri and D. pyroliflorus the pollinator remains unknown. Nor is it known whether flowers of D. leucocodon, which are large, pure white and bell-like and have a relatively small nectary, are pollen or nectar flowers.
In all species of Didymocarpus that I have observed in the field, the stigma in the receptive phase is white and glistening, presumably due to secretions by the papillose cells (Pl. 5b). (Papillose cells in D. leucocodon, P|. 5c, may be undeveloped as the stigma shown is from an immature gynaecium from a flower bud.)
In general, flowers with tubular corollas and enclosed anthers and style have larger, more or less discoid, peltate stigmas, e.g. D. leucanthus (Pl. 5a), D. parviflorus and D. quinquevulnerus (P|. Sh), compared with those flowers with a projecting style, where the stigma is globose or minute and rounded (D. anthonyi, Pl. 5d; D. corneri, Pl. Sf; D. geitleri, Pl. 5g). The stigma of D. geitleri is unique among Didymocarpus species is possessing a conspicuously naked zone between the stigma and the pustular trichome layer on the style.
Plate 5. Stigma morphology in Didymocarpus. a. D. leucanthus; b. papillose surface of D. leucanthus; ¢. D. leucocodon; d. D. anthonyi.
Five New Species of Didymocarpus 35
4. Didymocarpus salicinoides Kiew stat. et nom. nov.
Synonym: Paraboea salicinia (Ridley) Ridley var. major Ridley. Flora Malay Peninsula 5 (1925) 325.
Typus: Kelantan, Kuala Aring, Yapp 193 (lecto K, isolecto CGE). Distribution: Endemic to Peninsular Malaysia — south Kelantan, Trengganu. Habitat: Lowland forest growing on earth banks.
Specimens examined: Kelantan — Kuala Aring 12 September 1899 Yapp 193 (K, CGE); Trengganu — Kemaman, Ulu Bendong 30 October 1935 Corner SFN 30027 (K), Bk.
Plate 5. Stigma morphology in Didymocarpus (cont.). e. D. yongii; f. D. corneri; g. D. geitleri; h. D. quinquevulnerus.
36 Gard. Bull. Sing. 44(1) (1992)
Kajang 2 November 1935 Corner SFN 30198 (K), Bk. Bauk 27 August 1986 S. Anthony SA 596 (UPM), Sg. Nipa 5 May 1988 R. Kiew RK 2654 (K, L, SING, UPM).
Notes: This species shares several common characters with D. salicinus Ridley. Both are plants with a wiry stem with a tuft of narrowly lanceolate leaves at the top; the leaves are flat (i.e. the veins are not impressed above), glabrous and shiny above, thinly leathery and the margin is finely serrate; the flowers are small; and the fruits are short (12-20 mm long) and narrow. In addition, both produce young leaves which are white or pale pink at the base. The species epithet is chosen to reflect the close relationship between the two species.
Ridley (1925) distinguished his var. major from the typical variety by its broader leaves and panicled cymes. Table 2 lists additional differences between them.
Table 2 Diagnostic differences between Didymocarpus salicinus and D. salicinoides
Character D. salicinoides D. salicinus Stem unbranched branched (small plants unbranched)
Leaf width (mm) 20-33 10-22 Leaf apex acute acuminate Petiole length (mm) (8-) 20 (-27) (7-) 11 (-14) Petiole & midrib with
transverse ribs + — Corolla length (mm) 3 (-4) (2-) 3 Corolla colour white with purple pale violet-pink
upper lobes
Inflorescence length (cm) (4-) 5 (-8) 3 (-4) Inflorescence type twice branched cyme simple cyme No. flowers/inflorescence 7 (-10) 2-4
In addition, the habitat of these two taxa differs. Plants of D. salicinus are rheo- phytes, while those of D. salicinoides grow on earth banks in lowland forest not neces- sarily close to streams. While differences in leaf width may be attributed to the different conditions under which they live (and indeed plants of D. salicinus that grow above the flood level do have broader leaves, Kiew 1989), even when this is taken into account there is still a difference (Table 2).
It may be a coincidence that another species pair (D. heterophyllus-D. floribundus) shows this same difference in the riverine taxon having simple cymes and the forest undergrowth species panicled cymes and that these two species pairs grow together. Thus the plants with simple cymes, D. salicinus and D. heterophyllus grow along the Sg. Tahan, and D. salicinoides and D. floribundus grow together in the Kemaman area.
Didymocarpus salicinus and D. salicinoides belong to sect. Salicini.
5. Didymocarpus stoloniferus Kiew sp. nov.
Inter congeneribus Peninsulae Malaysiae habitu, foliis et fructibus ad D. puncticulatum accedens, sed stolonibus, pedunculis brevibus et corollis majoribus et tubaeformibus differt.
Typus: R. Kiew RK 1638 (holo UPM, iso SING).
Five New Species of Didymocarpus 37
A rosette plant producing thin stolons 30 cm or longer with plantlets at intervals. Leaves in opposite pairs forming a compact rosette of c. 8 leaves. Lamina broadly elliptic 3-6 by 1.5-2.5 cm, apex acute (rarely acuminate), base rounded sometimes unequal, in life fleshy, drying membraneous, margin serrate. Indumentum on upper surface of lamina dense with 4-celled unbranched hairs with conspicuously raised hair base, dense on lower surface of lamina, midrib, veins and petiole. Midrib and secondary veins depressed above and prominent beneath, secondary veins 6-9 pairs, tertiary veins obscure above and below. Petiole slender, 6-40 mm long.
Inflorescence 1-flowered. Peduncle and pedicel 12-16 mm long. Indumentum of pedicel and calyx with glandular hairs with a multicellular stalk. Bracts linear 1.5-2 mm long. Calyx divided to base, lobes ligulate, 2-3 by 1 mm. Corolla trumpet-shaped, tube 17 mm long by 1.5 mm wide at base dilating to 9 mm across at mouth, glabrous, whitish tinged purplish-pink, throat white with 2 lemon-yellow nectar guides, lobes pale purple-pink, equal-sized, broadly oblong, apex rounded, 4 by 4-5 mm. Stamens 2, filaments slender 4 mm long, anthers ellipsoid, 2 by 1 mm, connivent. Ovary ovoid, 2.5 by 1 mm, style slender 19 mm long, ovary and style densely pubescent, stigma discoid c. 0.5 mm across, not projecting beyond the corolla tube. Disc cylin- drical, c. 0.5 mm tall. Capsule 8-10 by 2.5 mm, spreading pubescent (Burtt & Stone B 11690).
Distribution: Endemic to Peninsular Malaysia — Pahang, Gunung Ulu Kali.
Habitat: Growing in moss on large granite rocks or rockfaces in upper montane forest at c. 1,600 m a.s.l.
Specimens examined: Pahang, G. Ulu Kali — 12 October 1978 B.L. Burtt & B.C. Stone B 11690 (E); 25 March 1985 Ruth Kiew RK 1638 (UPM, SING).
Notes: This species has been found at just two sites: one population was growing on the drier side of large rocks in forest (B 1/690), the other on a sheer rock face some 15 m high, which forms one side of a damp, dark gully (RK /638). Although the latter population has been visited at all times of year over a period of more than ten years, it has only been found in flower on two occasions (March 1985 and October 1989), when many plants were in flower. It is interesting that the other collection was also made in October (1978) when “the plants were flowering freely” (Burtt, pers. comm.).
The assignment of this species to one of the presently circumscribed sections is problematic. On the one hand its small rosette habit and short fruits ally it with sect. Boeopsis but it has neither the long peduncles nor the short campanulate corolla of this section; on the other hand its trumpet-shaped flower with distinct yellow nectar guides resembles species in sect. Heteroboea but it is not a robust plant with a woody stem and large leaves. Although it superficially resembles D. puncticulatus in sect. Boeopsis in its one-flowered inflorescence, its short fruit and small crenate leaves, which are hairy above; it differs from this species as D. puncticulatus does not have stolons, its leaves are punctate above and have a broad pale green band down the midrib, its inflorescence has a long peduncle and its flowers are shortly campanulate. In possessing stolons D. stoloniferus is unique among Malayan Didymocarpus.
Didymocarpus lithophilus Kiew Validated
Mr B.L. Burtt has pointed out to me that the application of Didymocarpus komp- soboea C.B.Cl. to a Malayan taxon being simply a misidentification, the new name D. lithophilus (Kiew, 1989) therefore requires a latin diagnosis to be valid, which is here supplied,
38 Gard. Bull. Sing. 44(1) (1992)
Didymocarpus lithophilus Kiew spec. nov. Gardens’ Bull. Singapore 42 (1989): 54.
Synonym: D. kompsoboea auct.; Ridley Trans. Linn. Soc. 2nd Ser. 3 (1893) 328; FI. Mal. Pen. 2 (1923): 518 — non C.B.Cl. in DC Mon. Phan. 5 (1883) 92.
Typus: Ridley 2152 Kuala Tahan, Pahang (holo K; iso SING).
Didymocarpo platypodi affinis sed venatione (in D. lithophilo areolis oblongis; in D. platypode areolis polygonalibus) et fructibus brevioribus differt. Differt a Didymocarpo rugoso foliis tenuibus non bullatis et pedunculis duplo longioribus.
Section Boeopsis
Section Boeopsis includes species that are smallish rosette plants with broadly cam- panulate flowers. The earliest Malayan species in this group, D. heterophyllus, was described by Ridley in 1893. In 1896, he grouped it with D. puncticulatus Ridley in sect. Kompsoboea. The latter includes species with a rosette habit but their flowers are larger and trumpet-shaped compared with the smaller campanulate ones of either D. heterophyllus or D. puncticulatus. (Sect. Kompsoboea is not represented in Penin- sular Malaysia.) In 1905 Ridley described a new section, Acaules, which he defined as comprising plants that are ‘Stemless or nearly so. Leaves crowded’ and in which he included D. violaceus Ridley, D. lacunosus Hook. f., D. pumilus Ridley as well as D. heterophyllus, D. perditus Ridley and D. puncticulatus. (Acaules is not available as a sectional name in Didymocarpus, as its lectotype, D. lacunosus has been transferred to Chirita.)
In 1907 Ridley described a new section, Boeopsis, to accommodate D. perditus, D. puncticulatus, D. heterophyllus and D. battamensis Ridley, the latter a species from Pulau Batam, an island south of Singapore. He described the species in this section as being small plants with short corolla tubes and two short stamens with thick sigmoid filaments and subglobose or elliptic anthers.
In his 1923 account of the genus, Ridley defined sect. Boeopsis as comprising plants with ‘leaves crowded in a tuft at the top of a woody root stock, peduncles slender, flowers usually small’ and in this he included D. longipes C.B.Cl., D. primulinus Ridley, D. soldanella Ridley, D. pumilus and D. grandiflorus Ridley, as well as those included in his 1907 account. D. grandiflorus (now renamed D. ridleyanus Burtt), on account of its distant pairs of leaves and large tubular flowers, obviously does not belong to this section and is now placed in sect. Didymanthus (Kiew, 1989). D. longipes and D. primulinus are both anomalous within this section in possessing yellow flowers with narrow corolla tubes. All other species in sect. Boeopsis have purple, campanulate corollas.
D. longipes is quite unlike any other Malayan species in its leaves and its condensed cymes borne on long peduncles. Its narrow tubular flowers recall those of D. flavo- brunneus Ridley and D. falcatus Kiew. In 1896 Ridley had noted that ‘it is difficult to find any species really nearly allied to this’ and he did not place it in sect. Komp- soboea. In his 1905 account he included D. /ongipes in sect. Didymanthus (i.e. not in sect. Acaules with D. heterophyllus and D. puncticulatus). Burtt (1954) selected D. longipes as the lectotype of sect. Boeopsis based on Ridley’s flora account of 1923 as he did not realize that this section had already been published in 1907 where it did not include D. longipes (Burtt, pers. comm.)
It is proposed here that sect. Boeopsis be redefined in its original 1907 sense and that D. longipes, D. primulinus and D. ridleyanus be excluded from the section. Burtt (1971) returned most species that Ridley had included in Paraboea sect. Campanulati
Five New Species of Didymocarpus 39
to Didymocarpus sect. Salicini. However, sect. Salicini is best kept in Ridley’s original sense for narrow-leaved species (see below). Only two of the remaining Paraboea species conform to the circumscription of sect. Boeopsis, viz. D. floribundus (Hender- son) Burtt and D. rubiginosus (Ridley) Burtt, both of which have a rosette habit and purple, campanulate flowers. Apart from the two new species described above, another three recently described species, D. codonion Kiew, D. n.sp (proposed to be named D. oreophilus Kiew) and D. yongii Kiew all belong to this section. D. perditus Ridley is a synonym of D. puncticulatus (Kiew, 1987). Therefore at present, this section in- cludes eleven species. D. heterophyllus is here chosen as the lectotype of the section as it 1s typical in its small rosette habit, its purple campanulate flowers and its short capsule.
Section Boeopsis Ridley J. Str. Br. Roy. Asiatic Soc. 49 (1907) 22; Ridley Fl. Mal. Pen. 2 (1923) 508.
Lectotype: D. heterophyllus Ridley Trans. Linn. Soc. 2nd Ser. 3 (1893) 329.
Small to medium-sized herbs, stemless or not, with a rosette of usually oblanceolate leaves, flowers held above the leaf rosette in a lax cyme of 3-18 (rarely 1-2) flowers, corollas short and broadly campanulate, purple (sometimes pink) without yellow nectar guides, stamens with short, thick filaments, anthers large, broadly oblong and prominently positioned in the mouth of corolla tube, style either contained within corolla tube or projecting well beyond it, ovary short, capsule short (up to 2.5 cm long).
Key to Malayan Species of Section Boeopsis
eer ewes Ce OUP... . tc oo ns Lee ee SEU LORS Es he nee soe cele eee ee 2 Perea velvety above, veins and lamina concolorous }ia.6 2.54. ch wee cee ee D. rubiginosus
2 eae manrous apove,. veins Outlined in white «22005 .6 ge ek wl le ere owe eee ee D. yongii TR ACEO TATRA ONIN eS oe gs gh A wei eee Cs elgg Fe hw ee eee Nees we 3 Drm eCiiee ett ONC. OF SONICHINIES 2 TOWGIS 25 os ecie Sv cle anc was tle cee ee ee cece eee 4 eat taDMIMS “ARONG, MIGUSIE ENTIRE eh Gee. eae se wie ela ees ob es D. leiophyllus ee gg Tee a ST 21 a eg ae eee a ear 5
5. Leaf sparsely hairy and minutely punctate above, fruit to 15 mm long ..................
aaa! elatit aiplibe a ee tee el ee le oF bee ene te ©, wales 6 6's she wei e ai © S's, 9 0.0 Sm 8 «a © 8 2, 0e@ © © © ©
5. Leaf densely velvety and not punctate above, fruit to 25 mm long ............. D. n. sp.
a OrCeCCtiCe CyMitse Will S7OF TMOTe TIQWETS | oi 6 cw icine re wl cee ee eee tesens 6 ene een TED OE NTIGIC LIOWEES sau. Ca, al. sce nS ws A eine Se nw sw ee alee eee em enes 7 7. Leaf petiole 2.5-7 cm, calyx 1-1.5 mm, corolla to 4 mm long ............... D. codonion
7. Leaf petiole 0.5-2 cm, calyx 3-5 mm, corolla to 7-9 mm long ............ D. floribundus
OO BSS a a ne Se SG ee a ee ee ii 8 RI PTEN GIOVE | 6 Fetes ' uidias 5 os Vanna s oy pile Regia Wa nied ve cmv ew ees D. pumilus RS ae of Sa ee ee ee ee 9 obese 12 -19-en long, Dase. NATTOWED 6.42). 55 ois eed Few eee eee eens D. anthonyi ama 14590 ION, DAase TOUNGER f. ) hw oa .2ic tnd ee eee eee eee wee eees 10
10. Leaf oblanceolate, apex acute, fruit c. 25 mm long .............. D. heterophyllus
10. Leaf obovate, apex rounded, fruit c. 1S mm long .................. D. soldanella
The geographic range of species in sect. Boeopsis is centred on the east and south of Peninsular Malaysia (Fig. 1). The majority are confined to the lowlands with the exception of D. puncticulatus, which has an altitudinal range from near sea level
Fig... 1.
Distribution of Didymocarpus species belonging to Section Boeopsis. 1. D. anthonyi; 2. D. codonion; 3. D. floribundus; 4. D. heterophyllus; 5. D. leiophyllus;
6. Didymocarpus sp. nov.; 7. D. soldanella; 8. D. puncticulatus; 9. D. rubiginosus; 10. D. pumilus
and 11. D. yongii.
Five New Species of Didymocarpus 4]
to 1,000 m. The few that are confined to the mountains include D. rubiginosus on Gunung Tahan (1,300-2,000 m), and D. n. sp. on G. Setong (1,000 m), D. pumilus on Fraser’s Hill (1,000 m) and D. so/ldanella on G. Kerbau, the latter three are from the Main Range but notably not collected from the western side. Apart from D. soldanella (unfortunately poorly known from a single scrap of a fruiting specimen), the other montane species all have leaves that are densely hairy on the upper surface, as does D. puncticulatus. The grey, silky indumentum of D. pumilus recalls the appearance of species of Loxocarpus and indeed it is this section that includes the Didymocarpus species with the shortest fruits (Kiew, 1987). However, all species of Loxocarpus have much smaller flowers and the cymes are more compressed.
Section Salicini
Ridley (1896) described species in section Salicini as being ‘small, short flowered species with narrow willow leaves crowded at the top of a short woody stem’. He included in this section D. pectinatus, D. salicinus and D. densiflorus, (the last an undescribed species based on a specimen collected by H.J. Kelsall from G. Janing, Johore). In 1905 he added D. serratifolius.
Ridley did not include this section in his 1923 account in which he transferred D. salicinus to Paraboea sect. Campanulati. The species that remained in Didymocarpus, D. densifolius, D. pectinatus and D. serratifolius, he placed in a new section Pectinati. (Section Pectinati is a distinct group of species with deeply serrate or pectinate leaf margins and small white tubular flowers. For this reason D. densifolius was excluded from this section, Kiew 1987).
In returning species from Paraboea Sect. Campanulati to Didymocarpus sect. Salicini, Burtt (1971) only excluded D. cordatus and D. tahanicus from sect. Salicini. However, apart from D. caeruleus, D. filicifolius and D. salicinus, the other species do not conform to Ridley’s concept of the section comprising species with willow-shaped leaves. Kiew (1987) has suggested that sect. Salicini be used in its original restricted sense. It presently includes the following species: D. densifolius Ridley (syn. Paraboea caerulea Ridley and D. azureus Burtt, Kiew 1987), D. salicinoides, D. salicinus Ridley (syn. D. filicifolius Ridley, Kiew, 1989) and D. tiumanicus (Ridley) Burtt. (Although D. holttumii (Henderson) Burtt has narrowly lanceolate leaves, 8-11 by 1.5-2 cm, it does not belong to this section as its leaves are arranged in distant pairs. In addition, its flower buds are reported as yellow. Flower colour in sect. Salicini ranges from pink to purple to bluish-purple.)
Key to Species in Section Salicini
ESS STAG GR IOe Te Yea ae SO: en an re re ee D. tiumanicus aE IRIN ONES IVE 8 inc an koe ated Hh we Se wees as eee eee ee ee 2
2. Leaves 10-19 cm long, decurrent, leaf margin entire, flowers 12-17 mm long .................. D. densifolius
a wine Sine e a aale as Aen 6 a em we eS ew @ © 2 eee Se 6 ee ee Eee eS he Bee eee 8 8 8
2. Leaves 5-7 cm long, petiolate, leaf margin minutely serrate, flowers 2-6 mm long ............ 3 3. Cymes simple, petiole not transversely ribbed ............. 0. cece eee eee ee eees D. salicinus 3. Cymes panicled, petiole and midrib transversely ribbed .................--. D. salicinoides
As mentioned above, D. salicinoides and D. salicinus are closely similar. They differ from D. densifolius and D. tiumanicus in their smaller flowers and slender peduncles. (D. tiumanicus has corollas c. 12 mm long). This raises the suspicion that this section may include rheophytes, and that the tufted habit and willow-shaped leaves reflect
42 Gard. Bull. Sing. 44(1) (1992)
ecological adaptation rather than relatedness of the species. The purple campanulate corolla with a wide mouth and conspicuous white anthers is also seen in sect. Boeopsis. However, until the range of floral structure in Didymocarpus is more fully understood, it is premature to split this section further.
Acknowledgements
I should like to thank Universiti Pertanian Malaysia for funding this project; S. Anthonysamy for assistance in the field; the Curators of the Herbaria at Cambridge Botany School, Royal Botanic Gardens Kew, and Singapore Botanic Gardens; Brian Chapman and A.J. Burgess for technical assistance in preparing material for and in using Joel JSM-35CF scanning electron microscope in Dept. Anatomy, Cambridge University and B.L. Burtt for his wise advice and helpful criticism and for supplying me with a fruiting specimen of Didymocarpus stoloniferus.
References
Burtt, B.L. (1954). Studies in the Gesneriaceae for the Old World. Sections proposed by Ridley. Notes Roy. Bot. Gard. Edinb. 21: 203-225.
____. (1971). Studies in the Gesneriaceae of the Old World. Didymocarpus sect. Salicini. Notes Roy. Bot. Gard. Edinb. 31: 43-44.
____. (1990). Gesneriaceae of the Old World. II. A new Didymocarpus from Sumatra. Edin. J. Bot. 47: 235-237.
Kiew, R. (1987). The herbaceous flora of Ulu Endau, Johore-Pahang, Malaysia, in- cluding taxonomic notes and descriptions of new species. Mal. Nat. J. 41: 201-234.
____. (1989). Didymocarpus (Gesneriaceae) on Gunung Tahan, Malaysia. Gard. Bull. Sing. 42: 47-64.
____. (1990). Reassessment of the generic status of Codonoboea (Gesneriaceae) and its species. Blumea 35: 167-176.
. (1991). Herbaceous Plants. In R. Kiew (ed.). The State of Nature Conservation in Malaysia. Malayan Nature Society, Kuala Lumpur, pp. 71-77.
Ridley, H.N. (1893). On the flora of the eastern coast of the Malay Peninsula. Trans. Linn. Soc. 2nd Ser. 3: 329.
___. (1896). Cyrtandraceae Malayenses — Didymocarpus sect. Salicini and sect. Kompsoboea. J. Linn Soc. Bot. 32: 514.
_____. (1905). The Gesneriaceae of the Malay Peninsula. J; Roy. As. Soc. Str. Br. 43: 27-57.
_____. (1907). New or rare Malayan Plants. Section Boeopsis. J. Roy. As. Soc. Str. Br. 49: 22.
____. (1923). Didymocarpus. Fl. Mal. Pen. 2: 506-523. _____. (1925). Didymocarpus. Fl. Mal. Pen. 5: 325.
Weber, A. (1989). Didymocarpus geitleri, a remarkable new species of Gesneriaceae with deceptive pollen flowers. Pl. Syst. Evol. 165: 95-100.
Weber, A. & Burtt, B.L. (1983). Didymocarpus corchorifolius and its allies (Gesneriaceae). Blumea 28: 291-309.
Notes on the Development of the Fruit-bodies of Four Malayan Species of Amanita (Basidiomycetes)
E.J.H. CORNER 91 Hinton Way Great Shelford
Cambridge CB2 5AH England
EFFECTIVE PUBLICATION DATE: 15 MAR 1993
Abstract
The development of the fruit-bodies was observed under natural conditions in the forest. Those of A. elata, A. princeps and A. virginea took 12-14 days to reach maturity when they persisted for merely 1-3 days. Expanded fruit-bodies soon became fly-blown and this hastened their decay. A. elata and A. princeps fructify early in the fungus season, as do most Malayan species, but A. virginea appears towards the end of the season. The presence of these species is revealed only for a few days twice each year.
Introduction
These observations were made in 1929-1931. The species used to fructify in roughly the same places every year in the Singapore Botanic Gardens Jungle and at Bukit Timah Forest Reserve, as if they were mycorrhizal, though I could not associate them with particular trees. However, this fact enabled me to disturb the humus gently in the likely spots and discover the young primordia. When to look for them was a few days after heavy rain had soaked the ground after the drier months of January-February and July-August (Corner, 1935). These species were described by Corner and Bas (1962).
Amanita elata (Mass.) Corner et Bas
From 22 March and 22 September 1930 I watched the successful development of 6 fruit-bodies in the Singapore Botanic Gardens Jungle. They reached, eventually, overall heights of 70-95 mm with pilei 25-80 mm wide. Several other fruit-bodies which I began to measure rotted off before the stem emerged from the volva. Measurements were made at about 8 a.m. daily. On day 1, the unopened volva was 8-10 mm high, 4-5 mm wide. By day 3, it had grown to 16-25 mm high, 9-13 mm wide. At 8 a.m. on day 4, the volva had ruptured, evidently during the night, and the stem had begun to project the pileus; the overall height was 26-63 mm but the convex pileus was merely 14-20 mm wide; the volva had ruptured into flat pieces on the unopended and pale umber pileus. On day 5, four fruit-bodies A-D were fully expanded, 70-78 mm high with plane pilei 30-60 mm wide. Two fruit-bodies, E and F, were c. 80 mm high with _ half-open pilei 30-46 mm wide. On day 6, fruit-bodies A-D were the same but E and F had fully expanded, 80-95 mm high with plane pilei 36 and 80 mm wide respectively. On day 7, A-D were dead. On day 8, E and F had collapsed by 4 p.m.
Full expansion from the volva had taken 48-72 hours and seemed to occur mainly during the night. The plane pileus persisted sporing for some 50-60 hours. From the incidence of heavy rain at that time, I judged that the mycelium had taken c. 10 days
43
44 Gard. Bull. Sing. 44(1) (1992)
to develop the primordia to their state on day 1. The full life of the fruit-body, there- fore, would be 14-15 days with a sporing period of c. 2 days or 50-60 hours.
The largest fruit-bodies that I recorded for this species had stems 13 cm long and pilei 9 cm wide. Such fruit-bodies might require an extra day for development and en- joy another day of sporing.
Amanita sp. aff. A. fritillaria (Berk.) Sacc.
On 15 March 1931 I marked two young fruit-bodies of this species in the Singapore Botanic Gardens Jungle. They were expanding with overall height 30 mm and pilei 11 mm wide. They expanded fully overnight and next morning were 73 mm high with plane pilei 38 mm wide. They lasted, evidently sporing, in this state for c. 36 hours before collapsing.
Amanita princeps Corner et Bas
In March and September 1930, I watched the development of 18 specimens of this lofty species. It grew in the deep shade of Fern Valley in Bukit Timah Forest Reserve. My observations were made at 3-4 p.m. The youngest specimens found were enclosed in the volva 15-21 mm wide. In 2 days the volva had enlarged to 32-48 mm wide. The next day, which was day 4 in the sequence, the volva had ruptured, evidently at night, and the stem had reached its full height 15-25 cm but the pilei were only one quarter to half open with the intact veil still covering the gills. On day 5, the pileus was fully expanded, plane or concave, 10-19 cm wide. The fruit-bodies then persisted for some 36-48 hours before becoming rotten. Many flies and small beetles had crawled over the expanding pilei to lay their eggs, and larvae together with the heavy rain hastened the demise of the fruit-bodies. Early development up to the rupture of the volva pro- bably took some 12 days. In my experience this conspicuous fungus could be seen merely on 3-4 days, twice a year in March and September.
Amanita virginea Mass.
This fungus is unlike other species of the genus in the Malay Peninsula because it fruits towards the end of the fungus season after 2-3 months of rainy weather. The fruit-bodies are not to be found in the usual run of fungus about March and September but in May or November-December. In 1929 I watched the development of 10 fruit-bodies which came up in the Singapore Botanic Gardens Jungle in the second half of November and in the first half of December. Four of these failed to ‘grow beyond an early stage when the primordia were merely a few days old. The others conformed to the sequence shown in Table 1.
The primordia took 8-10 days to develop from 10-15 mm high to the fully expanded state. The sporing period from the rupture of the veil to the collapse of the fruit-body varied from 30-70 hours. The expanded fruit-bodies were soon swarmed over by little flies, and how long they would last clearly depended on the extent to which they were fly-blown.
In 3 fruit-bodies the veil began to rupture about noon but was not fully broken and detached until 4 hours later. In one case the veil ruptured during the night. The veil split irregularly and fell to the ground in fairly large pieces.
It seemed likely that the primordia 10-15 mm high were not more than 3-4 days old. All the primordia and the freshly expanded fruit-bodies had very firm, turgid and compact texture. On section, a pale amber fluid issued from the cut surface, especially of the pileus and stem-apex.
Notes on Fruit-bodies of Malayan Amanita 45
Table 1 Fruit-body development of Amanita virginea
Day Height overall Pileus width 8 a.m. mm mm 1 10-15 9-10 pileus a small hump 2 20 14 3 25 18 4 38 23 3 45 30-35 6 60 40-45 7 75-85 50-55 8 Soo ioe } veil rupture 9 105-140 135-145 10 110-150 150-190 fully grown References
Corner, E.J.H. (1935). The seasonal fruiting of Agarics in Malaya. Gard. Bull. Str. Settl., 1X: 79-88.
and Bas, C. (1962). The genus Amanita in Singapore and Malaya. Persoonia 2: 241-304.
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Notes on the Rare Fern, Pteris holttumii C. Chr.
AZIZ BIDIN and RAZALI JAMAN Botany Department, Faculty of Life Sciences Universiti Kebangsaan Malaysia, 43600 Bangi
Selangor, Malaysia
EFFECTIVE PUBLICATION DATE: 15 MAR 1993
Abstract
Pteris holttumii C. Chr. was found in the vicinity of the lowland dipterocarp forest of Dent Peninsula, of Lahad Datu. It is the second record for Malaysia, and the only known record from the lowlands for the species.
Introduction
There is no comprehensive study on the Sabah ferns reported to date. The only detailed reference available is the work of Christensen and Holttum (1934) on the Mount Kinabalu ferns. The more recent treatments on the subject are rather general or restricted to specific taxa only (Price, 1987; Bidin & R. Jaman, 1989; Bidin, R. Jaman & K.M. Salleh, 1988). The richness of ferns in Sabah as exemplified by Mount Kinabalu, which habours about 500 species, will only be known once thorough studies have been conducted on the Crocker and Trus Madi Ranges as well as the lowlands.
In one of the many collecting trips to Sabah in search of ferns, the authors came across a handsome fern of the genus Preris in the lowlands of the Tabin Wildlife Reserve of Dent Peninsula (near Lahad Datu, East Sabah, alt. 50 m). The fern, Pteris holttumii C. Chr. was found on a steep river bank near a waterfall. Extensive search in the area failed to find the species in other localities. Specimens collected are de- posited at the Universiti Kebangsaan Malaysia Herbarium (UKMB) and a live plant brought back is grown in the Fernery of the same University (Fig. 1).
The find constitutes the second record for the species in Sabah. In describing the species in 1934, Christensen wrote: “This splendid new species, which I dedicate to its collector, is the finest novelty discovered in recent years....” The species collected by R.E. Holttum near Dallas Mt. Kinabalu (alt. +850 m) in 1931 was never recorded again in Sabah until the Tabin specimen surfaced. As for the region, the only find- ing for the species was by Hovenkamp & De Joncheere in Palu, Sulawesi at 500 m (Hovenkamp & De Joncheere, 1988).
Observations
External Features of P. holttumii
The gross morphological characters of the species resembles Acrostichum aureum in terms of size and divisions of frond. These characters prompted Christensen and Holttum to suggest that Acrostichum is derived from the Pterideae.
Rhizomes creep horizontally, slightly beneath ground, bearing solitary fronds at short intervals; thickly covered (especially at stipe base) by long wiry roots. Stipes are
47
Fig. 1. Pteris holttumii growing at the Universiti Kebangsaan Fernery.
grooved on the inner side, scattered, pale to yellowish, hard conicle prickles present. Fronds are simply pinnate; pinnae uniform, basal pinnae not branched on the basi- scopic sides near the base, basiscopic side of the base fused to the rachis, each pinna about 40 cm long. Rachises and upperside of costae are grooved, lower costae promi- nent. Veins are reticulate, forming up to 10 series of aerioles. Sori are marginal, elongated, without inner indusium (Fig. 2 A-E).
Endomorphic Characteristics of the Stipe
In transectional view, the stipe is subsulcate in outline enclosing a single vascular bundle which runs throughout the stipe. The bundle is a modified U-shaped strand with a wide base.
Ogura (1972) stressed the importance of the shape of the xylem strand in segregating families and genera of ferns, including relationships among taxa. In Preris holttumii the xylem strand follows the outline of the bundle with both ends curved inside but without hooks, which in Ogura’s classification is termed as the non-hippocampus type (Fig. 2B).
Summary
Pteris holttumii was first collected in Sabah in 1931 and described in 1934. Exten- sive botanical surveys by later workers in other parts of Borneo (Iwatsuki ef al. 1980; Iwatsuki & Kato, 1980a, & b, 1981 & 1983a & b) did not include the fern in their lists. With the present finding and that of Hovenkamp and De Joncheere in Sulawesi, it is established here that P holttumii is found in the lowlands as well as at high elevations.
i)
ana !
My
MW
(AY
V4
=
eT UM
=<" a —_, a
a
—s ——
Had i \ de | WS UY ¥ oN \\ HMA Ly), a INS Ani Pteris holttumii. A. Part of rhizome, showing roots and stipe bases. B. Xylem configuration middle of stipe. C. Part of frond, showing pinnae bases and sori outline. D. Vennation of pinna
nO) Hae
Fig. 2. E. Sporangium.
50 Gard. Bull. Sing. 44(1) (1992)
The species differs from the rest of the genus in having hard conicle prickles throughout the stipe as well as in the basal pinnae not being branched on the basiscopic sides near the base. It is the only species in Preris with reticulate venation. It is hoped that with the availability of a live specimen in our collection, the cytology of the species would be determined in due time in order to give some indications on the phylogenetic relationship within the genus.
Acknowledgements
We would like to thank Mr. Lamri Ali of Sabah Parks for allowing the authors to carry out a number of field surveys in the various localities in Sabah. The work was funded by research grant No. IRPA 4-07-03-007.
References Bidin, A., Jaman, R. and Salleh, K.M. (1988). A new species of Adiantum from Trus Madi Range, Sabah. Gard. Bull. Sing. 41(2): 45-48.
Bidin, A. and Jaman, R. (1989). The Pteriodophytes of Tabin Wildlife Reserve. Sabah Museum Journal. Special Issue. (In press).
Christensen, C. and Holttum, R.E. (1934). The ferns of Mount Kinabalu, Gard. Bull. Str. Settl. 7(3): 191-316.
Hovenkamp, P. and De Joncheere, G.J. (1988). Additions to the fern flora of Sulawesi. Blumea 33: 395-409.
Iwatsuki, K., Mogea, J.P., Murata, G. and Kartawinata, K. (1980). A botanical survey in Kalimantan during 1978-1979. Acta Phytotax. Geobot. 31(1-3): 1-23.
Iwatsuki, K. and Kato, M. (1980q@). Enumeration of Kalimantan Pteridophytes collected during 1978-1979(1). Acta Phytotax. Geobot. 31(1-3): 24-43.
. (19805). Enumeration of Kalimantan Pteridophytes collected during 1978- 1979(2). Acta Phytotax. Geobot. 31(4-6): 164-181.
_____. (1981). Enumeration of Kalimantan Pteridophytes collected during 1978-1979(3): Acta Phytotax. Geobot. 32(1-4): 121-132.
____. (1983a). Additions to the enumeration of East Kalimantan Pteridophytes 1. Acta Phytotax. Geobot. 34(1-3): 55-65.
____. (1983b). Additions to the enumeration of East Kalimantan Pteridophytes 2. Acta Phytotax. Geobot. 34(4-6): 130-141.
Ogura, Y. (1972). Comparative Anatomy of Vegetative Organs of the Pteridophytes. 2nd ed. Gebruder Borntraeger. Berlin, Stuttgart.
Price, M.G. (1987). Sabah ferns collected by John Beaman. Contr. Univ. Mich. Herb. 16: 189-200.
A Botanical Survey of Pulau Ubin
I.M. TURNER, H-TW. TAN, K.S. CHUA, HAJI SAMSURI BIN HAJI AHMAD and Y.C. WEE
Department of Botany National University of Singapore Lower Kent Ridge Road Singapore 0511
EFFECTIVE PUBLICATION DATE: 15 MAR 1993
Abstract
An intense botanical survey of Pulau Ubin, a 1019.2 ha island within the Republic of Singapore, found at least 332 native and naturalized vascular plant species. These are listed in the paper together with 40 species found to have escaped from cultivation. Previous botanical records for Pulau Ubin are also collated. The contemporary flora is dominated by early successional and ruderal species in addition to mangrove and beach forest elements. The low diversity and relatively high frequency of aliens (71 out of 332 spp.; 21 per cent of the flora) reflects the high degree of human disturbance on the island.
Introduction
Pulau Ubin is an island within the Republic of Singapore. It was chosen as a site for an intensive botanical survey because as one of the least urbanised and under botanized areas of the city state it was believed likely to possess an interesting flora. Additionally, development of Pulau Ubin is currently a topic of public interest in Singapore. Basic biological information such as a plant species list is prerequisite to the development of nature conservation programmes. This survey will therefore be of value to those involved in future decisions concerning changes in land use on Pulau Ubin.
Site
Pulau Ubin (N 1° 25’, E 103° 57’) is a granite island of 1019.2 ha (Ng, 1988) lying off the north-east coast of Singapore Island (Fig. 1). The main land uses at present are granite quarrying, agriculture, horticulture, aquaculture and recreational activities, largely of an athletic nature. Pulau Ketam and P. Sekudu, islets to the south of Ubin, were also included in the survey.
Methods
Most of the specimens were gathered by a team of 19 collectors during the period 17-22 June 1990. Some collections have been made subsequently on occasional visits to the island up to May 1991. The collections have been identified, largely by matching to named specimens in the Herbarium, Singapore Botanic Gardens (SING). The Pulau Ubin specimens have been deposited in the Herbarium, Department of Botany, National University of Singapore (SINU).
5]
PULAU UBIN
Sekudu 4
1km
103° 59° E
Fig. 1. | Map showing locality of study site.
Results and Discussion
The names of the species collected are given in Table 1, the few sterile collections that could not be identified have been omitted. A total of 332 native and naturalized species were collected plus 40 species of cultivated plants that had escaped or were relics of cultivation in now abandoned areas. The distinction between alien and escaped species is somewhat arbitrary and is based largely on Turner, Chua and Tan (1990). The Ubin flora included 24 pteridophytes and one gymnosperm; the rest (including all the escapes) being angiosperms.
The Ubin flora represents about one eighth of the total flora recorded for Singapore. The species list is made up largely by common weed, secondary forest and mangrove species. Primary forest species are nearly completely lacking; a Knema species sapling being the only real rain forest tree found. The orchids Spathoglottis plicata and Thrix- spermum amplexicaule were the most notable collections, though the former may possibly be a relic of cultivation. Eight species, Adiantum latifolium, .Hemigraphis primulaefolia, Pennisetum polystachyon, Scurrula parasitica, Sesamum radiatum,
A Botanical Survey of Pulau Ubin 53
Talinum paniculatum, Thysanolaena latifolia and Typha angustifolia, were not pre- viously recorded for Singapore. All but Scurrula parasitica can be called weeds, being able to colonise disturbed sites readily.
Pulau Ubin was presumably covered in lowland rain forest and mature mangrove forest until at least the middle of the Nineteenth Century. A list of previous botanical records from the island (Table 2) does include some species one would associate more strongly with primary vegetation such as Chisocheton macrophyllus, Cyathostemma viridiflorum, Dipterocarpus c.f. sublamellatus, Forrestia gracilis, Lithocarpus elegans, Phoebe grandis and several celastraceous climbers, but the general impression is that even by the 1880s and 1890s when Ridley and Hullett were collecting on Pulau Ubin much of the vegetation must have been secondary. One has to conclude that all of the terrestrial vegetation has been cleared at some time, much of this by the turn of the century. The state of the mangrove is little better. There are earlier records for 156 species from Pulau Ubin, of which 111 were not re-collected in the survey. This clearly reflects the rapid changes that have occurred on the island.
Two late Nineteenth Century collections of Ridley from Pulau Ubin have been des- cribed as isosyntypes. Neither of these species, Chisocheton macrophyllus and Ardisia singaporensis, were re-collected and are thus very probably extinct in one of their type localities. Hullett’s collection of Claoxylon longifolium from Ubin (Hullett s.n., March 1885) was described as a syntype of Claoxylon longifolium var. brachystachys by J.D. Hooker (Hooker 1887, p. 411) but this variety is now generally not recognized and the name reduced to synonymy with the type variety (e.g. Airy Shaw, 1972, Whitmore, 1973). Luerssen (1882) described Phegopteris subdecurrens from a fern collection made on Ubin (Kehding 2960) but this taxon is now generally synonymized to Tectaria semipinnata (Roxb.) Morton (e.g. Holttum, 1981).
The broad floristic changes brought about by human activities can be examined by comparing the relative abundances of the commonest angiosperm families in the current flora of Ubin and that recorded for Singapore as a whole (Table 3). The Orchidaceae are the commonest family in the flora of Singapore. Only five of the 194 species were found in the current survey on Pulau Ubin. The Leguminosae and the Euphorbiaceae are among the top three families with most species on contemporary Ubin. Both of these families have many weedy, alien species that have added to their diversity.
The flora of Pulau Ubin is clearly depauperate. Bukit Timah Nature Reserve has 854 species recorded (Corlett, 1990), nearly three times the number in one-fifteenth the area. Human interference on Pulau Ubin has lead to a semi-natural vegetation dominated by relatively few early successional species with a fairly high representation, 21 per cent of the flora, of alien species. It would be difficult to make a case for the conservation of the present day vegetation of Pulau Ubin on solely botanical grounds from an international perspective. However, from a Singapore standpoint, the patches of mangroves and the larger areas of belukar are of value in a country with so little natural vegetation remaining. Any plans for their destruction merit in-depth considera- tion of all the possible alternatives before being allowed to proceed.
Acknowledgements
The National University of Singapore Botany Honours Class (1990-1991) did most of the collecting on Pulau Ubin. We are most grateful for their efforts and enthusiasm. Our thanks also go to Soong Beng Ching, Shawn Lum and David Burslem for their assistance. We are grateful to the Director of the Singapore Botanic Gardens for allowing us access to the Herbarium. The survey was supported by the National Univer- sity of Singapore.
Table 1 List of species collected on Pulau Ubin since June 1990. Representative collection given in square brackets. Marginal note: a = alien species naturalized in Singapore, e = escape from cultivation.
PTERIDOPHYTES
Acrostichum aureum L. [M.F. Choong 22]
Acrostichum speciosum Willd. [H.H. Neo 11]
Adiantum latifolium Lamk. [MLY. Kok 32]
Asplenium nidus L. [A. Ho & J. Lee 11]
Blechnum orientale L. [K.S. Tan 47]
Ceratopteris thalictroides (L.) Brongn. [K.S. Chua & HW. Tan 427] Davallia denticulata (Burm.) Mett. [A. Ho & J. Lee 13] Dicranopteris linearis (Burm. f.) Underw. [A. Ho & J. Lee 1] Drynaria quercifolia (L.) J. Sm. [A. Ho & E.M. Sim 21] Lindsaea ensifolia Sw. [A. Ho & E.M. Sim 8]
Lycopodium cernuum L. [A. Ho & J. Lee 7]
Lygodium circinnatum (Burm. f.) Sw. [H.H. Neo 103] Lygodium microphyllum Cav.) R. Br. [A. Ho & J. Lee 5]
Nephrolepis biserrata (Sw.) Schott [A. Ho & J. Lee 3]
Phymatosorus scolopendria (Burm.) Pichi Serm. [I.M. Turner 42] Pityrogramma calomelanos (L.) Link [M. Chan & Roslina 31]
Platycerium coronarium (Koenig) Desv. [T.S. Teo 59]
Pteridium caudatum (L.) Maxon ssp. yarrabense (Domin) Parris [K.S. Tan 56] Pteris vittata L. [H.H. Neo 203]
Pyrrosia longifolia (Burm.) Morton [Latifah 6] Pyrrosia piloselloides (L.) Price [A. Ho & J. Lee 12]
Schizaea digitata (L.) Sw. [H.H. Neo 206] Stenochlaena palustris (Burm.) Bedd. [E.M. Sim & L.P. Ng 32] Taenitis blechnoides (Willd.) Sw. [H.H. Neo 5]
SPERMATOPHYTES
Acanthaceae
Acanthus ebracteatus Vahl [E.M. Sim & L.P. Ng 18] Acanthus volubilis Wall. [H.H. Neo 117]
Andrographis paniculata (Burm. f.) Nees [J. Lee 104] Asystasia nemorum Nees [E.M. Sim & L.P. Ng 14] Hemigraphis primulaefolia (Nees) Vill. [E.M. Sim & L.P. Ng 39]
Thunbergia fragrans Roxb. [M. Chan & Roslina 32] Thunbergia grandiflora Roxb. [T.S. Teo 124]
Agavaceae Sansevieria trifasciata Prain [L.P. Ng 21]
Aizoaceae Sesuvium portulacastrum L. [L.P. Law 4]
Table 1 (Continued)
Amaranthaceae
Celosia argentea L. [K.S. Tan 37] Cyathula prostrata (L.) Bl. [K.S. Chua 101] Gomphrena globosa L. [J. Sim 31]
Anacardiaceae
Anacardium occidentale L. [K.S. Chua & H:TW. Tan 440] Bouea macrophylla Griff. [M.F. Choong 58]
Buchanania arborescens (Bl.) Bl. [T.S. Teo 25] Campnosperma auriculatum (Bl.) Hook. f. [I.M. Turner 20] Mangifera indica L. [J. Sim 33]
Annonaceae Desmos dasymaschala (Bl.) Safford [M.F. Choong 38]
Apocynaceae
Allamanda cathartica L. [J. Sim 40]
Alstonia angustiloba Miq. [H.H. Neo 205] Catharanthus roseus (L.) G. Don [A. Ho & E.M. Sim 7] Cerbera odollam Gaertn. [I.M. Turner 111]
Araceae
Alocasia macrorrhizos (L.) G. Don [M.F. Choong 42] Colocasia esculenta (L.) Schott [H.H. Neo 10] Dieffenbachia seguine (Jacq.) Schott [T.S. Teo 70]
Araliaceae
Arthrophyllum diversifolium Bl. [L.P. Law 10] Polyscias fruticosa (L.) Harms [T.S. Teo 60] Schefflera elliptica (Bl.) Harms [H.H. Neo 111]
Asclepiadaceae
Dischidia major (Vahl) Merr. [M-Y. Kok 36] Dischidia nummularia R. Br. [J. Lee 1]
Hoya parasitica (Roxb.) Wall. ex. Wight [Latifah 117]
Avicenniaceae
Avicennia alba Bl. [J. Sim 23]
Avicennia officinalis Bl. [T.S. Teo 16] Avicennia rumphiana Hall. f. [T.S. Teo 20] Bignoniaceae
Spathodea campanulata Beauv. [M.Y. Kok 27]
Bombacaceae
Durio zibethinus J. Murray [K.S. Tan 17]
Boraginaceae Cordia cylindristachya R. & S. [E.M. Sim & L.P. Ng 43] Heliotropium indicum L. [{Roslina 45]
Table 1 (Continued)
Bromeliaceae
Ananas comosus (L.) Merr. cv. Mauritius [J. Sim 27] Cannaceae
Canna indica L. [M. Chan 30]
Capparaceae
Cleome rutidosperma DC. [Roslina 37]
Caricaceae
Carica papaya L. [J. Sim 21]
Casuarinaceae
Casuarina equisetifolia J. R. & G. Forst. [J. Sim 7] Combretaceae
Lumnitzera littorea (Jack) Voigt [M.Y. Kok 38] Lumnitzera racemosa Willd. [J. Sim 26]
Terminalia catappa L. [J. Sim 1]
Commelinaceae
Commelina diffusa Burm. f. [L.P. Law 18]
Compositae
Ageratum conyzoides L. [L.P. Ng & L.P. Law 2]
Bidens pilosa L. [K.S. Chua 100]
Blumea balsamifera (L.) DC. [K.S. Chua & HTW. Tan 443] Eclipta prostrata (L.) L. [M.F. Choong 43]
Elephantopus scaber L. [A. Ho & E.M. Sim 14]
Emilia sonchifolia (L.) DC. ex Wight [A. Ho & E.M. Sim 15] Erigeron sumatrensis Retz. [M.Y. Kok 25]
Gynura procumbens (Lour.) Merr. [L.P. Ng 8]
Mikania cordata (Burm. f.) B.L. Robinson [M.Y. Kok 21]
Pluchea indica (L.) Less. [Roslina 19]
Sparganophora sparganophorus (L.) C. Jeffrey [K.S. Chua & H.TW. Tan 428] Spilanthes iabadicensis A.H. Moore [K.S. Chua & H-T'W. Tan 100] Synedrella nodiflora (L.) Gaertn. [J. Lee 3]
Tridax procumbens L. [M. Chan 33]
Vernonia cinerea (L.) Less. [M.Y. Kok 33]
Wedelia biflora (L.) DC. [MLY. Kok 11] Wedelia trilobata (L.) Hitch. [E.M. Sim & L.P. Ng 42]
Youngia japonica (L.) DC. [T.S. Teo 125] Connaraceae
Cnestis palala (Lour.) Merr. [T.S. Teo 57] Convolvulaceae
Erycibe tomentosa BI. [T.S. Teo 54]
Ipomoea aquatica Forsk. [A. Ho & E.M. Sim 34] Ipomoea batatus (L.) Lamk. [L.P. Law & L.P. Ng 20] Ipomoea cairica (L.) Sweet [J. Sim 41]
Merremia tridentata (L.) Hallier f. [M. Chan 53]
e
Table 1 (Continued)
Cucurbitaceae Cucumis sativa L. [M.F. Choong 50]
Cyperaceae Bulbostylis barbata (Rottb.) Clarke [Roslina 21]
Cyperus aromaticus (Ridl.) Mattf. & Kuk [Roslina 6] Cyperus compactus Retz. [Latifah & H.H. Neo 2] Cyperus compressus L. [Roslina 1]
Cyperus cyperinus (Retz.) Valck. Sur. [Latifah 2] Cyperus halpan L. [M. Chan 13]
Cyperus javanicus Houtt. [A. Ho & E.M. Sim 66] Cyperus kyllingia Endl. [Latifah 18]
Cyperus pilosus Vahl [Latifah & H.H. Neo 3] Cyperus trialatus (Boeck.) Kern [Latifah 1]
Fimbristylis cymosa R. Br. [Latifah 12]
Fimbristylis dichotoma (L.) Vahl [Roslina 4] Fimbristylis griffithii Boeck. [M. Chan 10] Fimbristylis polytrichoides (Retz.) R. Br. [Latifah 11] Fimbristylis schoenoides (Retz.) Vahl [H.H. Neo 204]
Hypolytrum nemorum (Vahl) Spreng. [H.H. Neo & Latifah 1]
Scleria corymbosa Roxb. [M. Chan & Roslina 27] Scleria levis Retz. [Latifah 10]
Thoracostachyum bancanum (Miq.) Kurz [Latifah 13]
Dilleniaceae Dillenia suffruticosa (Griff.) Mart. [E.M. Sim & L.P. Ng 8] Tetracera indica (Christm. & Panz.) Merr. [E.M. Sim & L.P. Ng 27]
Dioscoreaceae
Dioscorea glabra Roxb. [T.S. Teo 29] Dioscorea laurifolia Wall. ex Hook. f. [K.S. Chua & H‘TW. Tan 437]
Elaeocarpaceae
Elaeocarpus ferrugineus (Jack) Steud. [I.M. Turner 38] Elaeocarpus pedunculatus Wall. ex Mast. [J. Sim 5]
Muntingia calabura L. [T.S. Teo 43]
Eriocaulaceae Eriocaulon longifolium Nees [M. Chan 12]
Erythroxylaceae Erythroxylum cuneatum (Miq.) Kurz [K.S. Tan 30]
Euphorbiaceae
Antidesma velutinosum Bl. [L.P. Law 8]
Baccaurea motleyana (M.A.) M.A. [A. Ho & E.M. Sim 13]
Breynia coronata Hook. f. [T.S. Teo 10]
Bridelia stipularis (L.) Bl. [M. Chan & Roslina 29]
Claoxylon indicum (Reinw. ex Bl.) Endl. ex Hassk. [M.F. Choong 19] Codiaeum variegatum (L.) Bl. [T.S. Teo 64]
og yo HY O HY DY
Table 1 (Continued)
Croton hirtus L’Héritier [K.S. Tan 28]
Euphorbia hirta L. [M.F. Choong 40]
Excoecaria agallocha L. [K.S. Tan 3]
Glochidion superbum Baill. [I.M. Turner 113]
Hevea brasiliensis (Willd. ex A. Juss.) M.A. [MLY. Kok 7]
Macaranga conifera M.A. [MLY. Kok 54]
Macaranga gigantea (Rchb. f. & Zoll.) M.A. [L.P. Law & L.P. Ng 25] Macaranga griffithiana M.A. [I.M. Turner 116]
Macaranga heynei 1.M. Johnston [K.S. Tan 19]
Macaranga hypoleuca (Rchb. f. & Zoll.) M.A. [M.Y. Kok 26] Macaranga triloba (Bl.) M.A. [L.P. Law & L.P. Ng 7]
Mallotus paniculatus (Lamk.) M.A. [M. Chan & Roslina 25]
Manihot esculenta Crantz [MLY. Kok 28] Manihot glaziovii M.A. [H.H. Neo 104]
Phyllanthus acidus (L.) Skeels [T.S. Teo 50]
Phyllanthus amarus Schum. & Thonn. [A. Ho & E.M. Sim 35] Phyllanthus debilis Klein ex Willd. [E.M. Sim & L.P. Ng 7] Phyllanthus urinaria L. [E.M. Sim & L.P. Ng 35]
Ricinus communis L. [K.S. Tan 48] Sapium discolor (Champ. ex Benth.) M.A. [T.S. Teo 9] Suregada multiflora (Juss.) Baill. [I1.M. Turner 30]
Flagellariaceae Flagellaria indica L. [M. Chan 11]
Gnetaceae Gnetum macrostachyum Hook. [J. Sim 13]
Goodeniaceae Scaevola sericea Vahl [MLY. Kok 39]
Gramineae
Axonopus compressus (Swartz) Beauv. [M. Chan 15] Bambusa glaucescens (Willd.) Sieb. [T.S. Teo 28] Centotheca lappacea (L.) Desv. [Latifah 3]
Chloris barbata Swartz [A. Ho & E.M. Sim 27] Chrysopogon aciculatus (Retz.) Trin. [Latifah 15]
Coix lacryma-jobi L. [T.S. Teo 39]
Cynodon dactylon (L.) Pers. [K.S. Chua & HW. Tan 447] Cyrtococcum accrescens (Trin.) Stapf [Latifah & H.H. Neo 5] Dactyloctenium aegyptium (L.) P. Beauv. [Roslina 17] Digitaria ciliaris (Retz.) Koel. [K.S. Chua & H-:TW. Tan 446] Echinochloa colona (L.) Link [M. Chan & Roslina 34] Eleusine indica (L.) Gaertn. [M. Chan 18]
Eragrostis pilosa (L.) P. Beauv. [Latifah 9]
Imperata cylindrica (L.) P. Beauv. [M. Chan 8]
pe)
Table 1 (Continued)
Ischaemum indicum (Houtt.) Merr. [Roslina 3] Ischaemum muticum L. [M. Chan 1]
Mnesithea glandulosa (Trin.) Koning & Sosef [T.S. Teo 121] Panicum maximum Jacq. [L.P. Law 23]
Paspalum conjugatum Berg. [Latifah 6] Paspalum orbiculare Forst. f. [Latifah 7]
Pennisetum polystachyon (L.) Schult. [Roslina 16]
Pogonatherum paniceum (Lamk.) Hack. [K.S. Chua & H:TW Tan 445] Rhynchelytrum repens (Willd.) C.E. Hubb. [Roslina 7]
Saccharum arundinaceum Retz. [L.P. Law 17]
Sporobolus indicus (L.) R. Br. [M. Chan 12]
Thysanolaena latifolia (Roxb. ex Hornem.) Honda [A. Ho & E.M. Sim 19] Zoysia matrella (L.) Merr. [M. Chan 14]
Guttifera
Calophyllum inophyllum L. [MLY. Kok 51] Calophyllum pulcherrimum Wall. ex Choisy [H.H. Neo 207]
Garcinia mangostana L. [K.S. Tan 15] Garcinia nigrolineata Planch. ex T. Anders. [E.M. Sim & L.P. Ng 5]]
Hypoxidaceae Curculigo orchioides Gaertn. [K.S. Tan 27]
Ixonanthaceae Ixonanthes reticulata Jack [M.F. Choong 17]
Labiatae
Hyptis brevipes Poit. [A. Ho & E.M. Sim 20] Hyptis capitata Jacq. [K.S. Chua 99] Hyptis suaveolens (L.) Poit. [A. Ho & E.M. Sim 45]
Leucas zeylanica (L.) R. Br. [E.M. Sim & L.P. Ng 36] Ocimum basilicum L. [A. Ho & E.M. Sim 33]
Lauraceae
Cassytha filiformis L. [L.P. Law 21] Cinnamomum iners Reinw. ex BI. [K.S. Tan 49] Neolitsea zeylanica Merr. [M.Y. Kok 29]
Leeaceae Leea indica (Burm. f.) Merr. [M-Y. Kok 31]
Leguminosae Abrus precatorius L. [J. Lee 4]
Acacia auriculiformis A. Cunn. ex Benth. [J. Sim 15] Acacia mangium Willd. [I.M. Turner 44]
Alysicarpus vaginalis (L.) DC. [E.M. Sim & L.P. Ng 48] Andira inermis (W. Wright) H.B.K. ex DC. [I.M. Turner 115] Archidendron clypearia (Jack) I. Nielsen [M.F. Choong 16]
bab)
og © Y &
Table 1 (Continued)
Caesalpinia crista L. [K.S. Chua & H-TW. Tan 426] Calopogonium mucunoides Desv. [L.P. Ng 10] Canavalia cathartica Thou. [A. Ho & J. Lee 15]
Cassia alata L. [L.P. Law 19] Cassia lechenaultiana DC. [K.S. Chua 80] Cassia mimosoides L. [L.P. Ng & L.P. Law 19]
Centrosema pubescens Benth. [A. Ho & E.M. Sim 44] Clitoria laurifolia Poir. [J. Sim 18]
Crotalaria mucronata Desv. [K.S. Tan 8]
Dalbergia candenatensis (Dennst.) Prain [T.S. Teo 17]
Derris elliptica (Roxb.) Benth. [M-Y. Kok 3] Derris trifoliata Lour. [A. Ho & J. Lee 14]
Desmodium heterocarpon (L.) DC. [L.P. Ng & L.P. Law 14] Desmodium heterophyllum (Willd.) DC. [M. Chan & Roslina 31] Desmodium umbellatum (L.) DC. [M.F. Choong 11]
_ Entada spiralis Ridl. [K.S. Tan 12]
Intsia bijuga (Colebr.) O. Ktze. [K.S Chua & H.TW. Tan 441]
Mimosa invisa Mart. ex Colla [T.S. Teo 42] Mimosa pigra L. [M.F. Choong 28] Mimosa pudica L. [M. Chan & Roslina 30]
Neptunia plena (L.) Benth. [T.S. Teo 71] Peltophorum pterocarpum (DC.) Backer ex Heyne [K.S. Tan 25] Pterocarpus indicus Willd. [A. Ho & E.M. Sim 40]
Lemnaceae Lemna perpusilla Torrey [K.S. Chua & H-TW. Tan 449]
Liliaceae
Cordyline fruticosa (L.) A. Chev. [L.P. Law 15] Dianella ensifolia (L.) DC. [M. Chan 19] Gloriosa superba L. [J. Lee 102]
Linaceae Indorouchera griffithiana (Planch.) Hallier f. [H.H. Neo 208]
Loganiaceae
Fagraea acuminatissima Merr. [I1.M. Turner 105]
Fagraea fragrans Roxb. [MLY. Kok 12]
Loranthaceae
Dendrophthoe pentandra (L.) Mig. [A. Ho & J. Lee 10] Macrosolen cochinchinensis (Lour.) Tiegh. [H.H. Neo 114] Scurrula parasitica L. [P-T. Chew & A.S. Chew 46]
Magnoliaceae Michelia champaca L. [J. Lee 5]
Malpighiaceae Tristellateia australasiae A. Rich. [E.M. Sim & L.P. Ng 20]
Table 1 (Continued)
Malvaceae
Hibiscus rosa-sinensis L. [L.P. Ng 14] Hibiscus tiliaceus L. [E.M. Sim & L.P. Ng 4]
Sida rhombifolia L. [E.M. Sim & L.P. Ng 41] Thespesia populnea (L.) Soland. ex Correa [J. Lee 8] Urena lobata L. [E.M. Sim & L.P. Ng 34]
Melastomataceae
Clidemia hirta D. Don [E.M. Sim & L.P. Ng 6] Dissochaeta gracilis (Jack) Bl. [K.S. Chua 104] Melastoma malabathricum L. [E.M. Sim & L.P. Ng 2] Memecylon edule Roxb. [K.S. Tan 20]
Meliaceae Xylocarpus granatum Koen. [M.F. Choong 4]
Menispermaceae Fibraurea tinctoria Lour. [L.P. Ng & L.P. Law 4] Limacia scandens Lour. [M.F. Choong 20]
Moraceae Artocarpus integer (Thunb.) Merr. [J. Lee 6] Artocarpus heterophyllus Lamk. [A. Ho & E.M. Sim 26]
Ficus aurata Miq. [K.S. Tan 63]
Ficus fistulosa Reinw. ex Bl. [M.F. Choong 49]
Ficus grossularioides Burm. f. [M.F. Choong 6]
Ficus heteropleura Bl. [M.Y. Kok 19]
Ficus microcarpa L. f. [T.S. Teo 34]
Ficus variegata Bl. [K.S. Tan 36]
Ficus virens Ait. var. glabella (Bl.) Corner [M.F. Choong 36]
Streblus elongatus (Miq.) Corner [MLY. Kok 47]
Musaceae Musa acuminata Colla cultivar [J. Sim 32]
Myricaceae Mpyrica esculenta Buch.-Ham. [L.P. Law & L.P. Ng 8]
Mpyristicaceae Knema sp. [K.S. Tan 51]
Mpyrsinaceae Ardisia crenata Sims [T.S. Teo 72] Ardisia elliptica Thunb. [T.S. Teo 14]
Embelia ribes Burm. [M.Y. Kok 16]
Myrtaceae
Eugenia jambos L. [K.S. Tan 54]
Eugenia longiflora (Pres) F.-Vill. [M.Y. Kok 14] Eugenia palembanica (Miq.) Merr. [K.S. Tan 18] Eugenia spicata Lamk. [T.S. Teo 1]
pe)
Table 1 (Continued)
Psidium guajava L. [MLY. Kok 30]
Rhodamnia cinerea Jack [J. Sim 19]
Rhodompyrtus tomentosa (Ait.) Hassk. [Roslina 15]
Tristaniopsis whitiana (Griff.) Wilson & Waterhouse [M-Y. Kok 40]
Nepenthaceae Nepenthes gracilis Korth. [E.M. Sim & L.P. Ng 54]
Olacaceae Ximenia americana L. [K.S. Tan 1]
Onagraceae Ludwigia hyssopifolia (G. Don) Exell [M. Chan & Roslina 30]
Opiliaceae Champereia manillana (Bl.) Merr. [E.M. Sim & L.P. Ng 13]
Orchidaceae
Bromheadia finlaysoniana (Lindl.) Rchb. f. [M. Chan 61] Dendrobium crumenatum Sw. [A. Ho & J. Lee 9] Spathoglottis plicata Bl. [A. Ho & E.M. Sim 42] Thrixspermum amplexicaule (Bl.) Rchb. f. [T.S. Teo 32] Vanilla griffithii Rchb. f. [T.S. Teo 26]
Oxalidaceae
Oxalis barrelieri L. [E.M. Sim & L.P. Ng 9]
Oxalis corniculata L. [J. Lee 2]
Palmae
Arenga pinnata (Wurmb) Merr. [H.H. Neo 112] Caryota mitis Lour. [K.S. Tan 10]
Cocos nucifera L. [K.S. Tan 5]
Licuala spinosa Wurmb [J. Sim 12]
Nypa fruticans Wurmb [J. Sim 29]
Oncosperma tigillarium (Jack) Ridl. [I.M. Turner 37]
Pandanaceae
Pandanus amaryllifolius Roxb. [K.S. Tan 46] Pandanus odoratissimus L. f. [J. Sim 2] Pandanus yvanii Solms. [L.P. Law & L.P. Ng 3]
Passifloraceae
Passiflora foetida L. [K.S. Tan 13]
Passiflora laurifolia L. [E.M. Sim & L.P. Ng 44] Passiflora suberosa L. [A. Ho & J. Lee 45]
Pedaliaceae
Sesamum radiatum Schum. [K.S. Chua & H:TW. Tan 82]
Piperaceae Peperomia pellucida (L.) H.B.K. [M.F. Choong 39]
Table 1 (Continued)
e Piper betle L. [I.M. Turner 53] Piper sarmentosum Roxb. ex Hunter [M. Chan & Roslina 29]
Plantaginaceae a Plantago major L. [M.F. Choong 52]
Portulacaceae a Talinum paniculatum (Jacq.) Gaertn. [M.F. Choong 57]
Rhamnaceae Colubrina asiatica L. ex Brongn. [T.S. Teo 38]
Rhizophoraceae
Bruguiera cylindrica (L.) Bl. [T.S. Teo 19] Bruguiera gymnorrhiza (L-.) Lamk. [T.S. Teo 23]
Ceriops tagal (Perr.) C.B. Robinson [M.F. Choong 29] Gynotroches axillaris Bl. [A. Ho & E.M. Sim 56]
Rhizophora apiculata Bl. [M.F. Choong 31] Rhizophora mucronata Poir. [M.F. Choong 47]
Rubiaceae
a Borreria alata (Aubl.) DC. [M.F. Choong 15] Borreria articularis (L. f.) FN. Will. [E.M. Sim & L.P. Ng 29] Borreria laevicaulis (Miq.) Ridl. [A. Ho & E.M. Sim 9] Borreria setidens (Miq.) Bold. [K.S. Chua & HW. Tan 434]
a Diodia ocymifolia (Willd. ex R. & S.) Bremek. [A. Ho & E.M. Sim 17] Guettarda speciosa L. [J. Sim 9] Hedyotis corymbosa (L.) Lamk. [L.P. Law & L.P. Ng 61] Ixora congesta Roxb. [T.S. Teo 24]
a Morinda citrifolia L. [A. Ho & E.M. Sim 3] Morinda umbellata L. [E.M. Sim & L.P. Ng 47]
Oxyceros longiflora (Lamk.) Yamazaki [E.M. Sim & L.P. Ng 31] Scyphiphora hydrophyllacea Gaertn. f. [T.S. Teo 15]
Tarenna costata (Mig.) Merr. [T.S. Teo 4] Tarenna fragrans (Nees) K. & V. [T.S. Teo 3]
Timonius wallichinus (Korth.) Valeton [J. Sim 2]
Rutaceae Euodia roxburghiana (Cham.) Benth. ex Hook. f. [K.S. Chua & H-‘TW. Tan 108] e Murraya koenigii (L.) Spreng. [M. Chan 27]
Santalaceae Dendrotrophe varians (Bl.) Miq. [I.M. Turner 102]
Sapindaceae Allophyllus cobbe (L.) Raeusch. [M.Y. Kok 22] Guioa pleuropteris (Bl.) Radlk. [A. Ho & E.M. Sim 57] Mischocarpus sundaicus Bl. [1.M. Turner 36] e Nephelium lappaceum L. [MLY. Kok 45]
Table 1 (Continued)
Sapotaceae Planchonella obovata (R. Br.) Pierre [T.S. Teo 12]
Scrophulariaceae Limnophila sessiliflora Bl. [K.S. Chua & H:TW. Tan 431]
Lindernia antipoda (L.) Alston [E.M. Sim & L.P. Ng 30] Lindernia crustacea (L.) Fyv.M. [K.S. Chua & H:TW. Tan 432] Lindernia sessiliflora (Benth.) Wettst. [K.S. Chua & HITW. Tan 433]
Scoparia dulcis L. [A. Ho & E.M. Sim 18]
Simaroubaceae Brucea javanica (L.) Merr. [T.S. Teo 45]
Smilacaceae Smilax megacarpa DC. [K.S. Chua & H.TW. Tan 425]
Solanaceae
Physalis minima L. [K.S. Chua 71]
Solanum melongena L. [K.S. Chua 72]
Solanum torvum Sw. [MLY. Kok 17]
Sonneratiaceae
Sonneratia alba J.J. Smith [K.S. Chua & H.TW. Tan 442] Sonneratia ovata Backer [J. Lee 7]
Sterculiaceae
Commersonia bartramia (L.) Merr. [M.Y. Kok 52] Heritiera littoralis Dryand. ex W. Ait. [I.M. Turner 45] Pterospermum diversifolium Bl. [T.S. Teo 40]
Symplocaceae Symplocos fasciculata Zoll. [E.M. Sim & L.P. Ng 52]
Theaceae Adinandra dumosa Jack [E.M. Sim & L.P. Ng 26] Eurya acuminata DC. [MLY. Kok 56]
Thymelaeaceae Linostoma pauciflorum Griff. [Latifah & H.H. Neo 4]
Tiliaceae Triumfetta rhomboidea Jacq. [M.F. Choong 55]
Turneraceae Turnera ulmifolia L. [Roslina 12]
Typhaceae Typha angustifolia L. [1.M. Turner 117]
Ulmaceae
Trema cannabina Lour. [K.S. Tan 34] Trema tomentosa (Roxb.) Hara [A. Ho & E.M. Sim 40]
e Clerodendrum paniculatum L. [T.S. Teo 61] e Clerodendrum philippinum Schauer [A. Ho & E.M. Sim 22] Clerodendrum villosum Bl. [K.S. Chua 69] Gmelina asiatica L. [J. Lee 103] a Lantana camara L. [T.S. Teo 31] Premna corymbosa (Burm. f.) Rottl. & Willd. [K.S. Tan 35. Stachytarpheta indica (L.) Vahl [K.S. Chua 95] Vitex pinnata L. [J. Sim 14] Vitex trifolia L. [K.S. Tan 52] Vitaceae Ampelocissus elegans (Kurz) Gegnep. [M-Y. Kok 7] Cissus hastata (Miq.) Planch. [A. Ho & E.M. Sim 6] Zingiberaceae e Languas galanga (L.) Stuntz. [L.P. Law 9] Table 2 Species previously recorded from Pulau Ubin; representative collections are indicated in square brackets. Species PTERIDOPHYTES
Table 1 (Continued)
Umbelliferae Centella asiatica (L.) Urb. [Roslina 28]
Urticaceae Laportea interrupta (L.) Chew [H.H. Neo 105]
Verbenaceae
Clerodendrum inerme (L.) Gaertn. [E.M. Tim & L.P. Ng 49] Clerodendrum laevifolium Bl. [E.M. Sim & L.P. Ng 44]
Adiantum flabellulatum L. Asplenium macrophyllum Sw. Cheilanthes tenuifolia (Burm.) Sw. Drynaria sparsisora (Desv.) Moore
Humata heterophylla (Sm.) Desv.
Lindsaea divergens Hk. & Grev.
Phymatosorus scolopendria (Burm.) Pic. Ser.
Pityrogramma calomelanos (L.) Link
Pteris ensiformis L. [Ridley 3040]
Pyrrosia lanceolata (L.) Farwell [Ridley 9510] Tectaria semipinnata (Roxb.) Morton [Ridley 6027]
Reference
Ridley 1900 Ridley 1900 Ridley 1900 Johnson 1977 Johnson 1977 Johnson 1977 Johnson 1977 Johnson 1977
Ridley 1900 Johnson 1977
Table 2 (Continued)
Species Reference SPERMATOPHYTES
Agavaceae
Dracaena elliptica Thunb. Ridley 1900 Dracaena porteri Bak. Ridley 1900 Amaranthaceae
Amaranthus lividus L. [Ridley 4690] Keng 1990 Cyathula prostrata (L.) BI. Ridley 1900 Amaryllidaceae
Crinum asiaticum L. Ridley 1900 Annonaceae
Cyathostemma viridiflorum Griff. [Ridley s.n. Feb 1894] Keng 1990 Apocynaceae
Urceola lucida (DC.) Hook. f. [Ridley s.n. 1894] Ridley 1900 Urnularia flavescens (Hook. f.) Stapf Ridley 1900 Willughbeia coriacea Wall. [Ridley 9501] Ridley 1900 Willughbeia grandiflora Dyer ex Hook. f. [Ridley s.n. 1893] Markgraf 1972 Araceae
Aglaonema simplex Bl. [Ridley s.n. 1890] Ridley 1900 Araliaceae
Schefflera cephalotes (C.B. Clarke) Harms. Ridley 1900 Schefflera lanceolata Ridl. Ridley 1900 Asclepiadaceae
Calotropis procera (Ait.) Ait. f. — esc. cult. Ridley 1900 Avicenniaceae
Avicennia officinalis Bl. Ridley 1900 Bignoniaceae
Dolichandrone spathacea (L. f.) K. Schum. Ridley 1900 Celastraceae
Reissantia indica (Willd.) Hallé Ridley 1900 Salacia chinensis L. Ridley 1900 Salacia grandiflora Kurz [Ridley 4784] Ridley 1900 Commelinaceae
Aclisia secundiflora (Bl.) Bakh. f. [Ridley 4759] Ridley 1900 Forrestia gracilis Ridl. [Ridley 4810] Ridley 1900 Compositae
Adenostemma lavenia (L.) O. Kuntze [Hullett 78] Ridley 1900
Blumea riparia (Bl.) DC. [Ridley s.n. 1894] oes,
Table 2 (Continued) Species
Eleuthanthera ruderalis (Sw.) Sch.-Bip. [Furtado 18629] Erigeron sumatrensis Retz. [Furtado 18342]
Synedrella nodiflora (L.) Gaertn. [Furtado 18624] Tridax procumbens L. [Furtado 18347]
Wedelia biflora (L.) DC. [Hullett 387]
Connaraceae Connarus planchonius Schellenb.
Cyperaceae
Fimbristylis ferruginea (L.) Vahl Scleria oblata S17. Blake [Furtado 18630]
Dipterocarpaceae Dipterocarpus c.f. sublamellatus Foxw. [Ridley s.n. 1890]
Euphorbiaceae
Acalypha indica L.
Antidesma velutinosum Bl. [Hullett 629]
Claoxylon longifolium Endl. ex Hassk. [Hullett s.n. 1885] Galearia fulva (Tul.) Miq.
Glochidion microbotrys Hook. f.
Macaranga triloba (Bl.) M.A.
Suregada multiflora (Juss.) Baill.
Fagaceae
Lithocarpus elegans (Bl.) Hatus. ex Soepadmo Lithocarpus wallichianus (Lindl. ex Hance) Rehd. [Ridley 7479]
Flagellariaceae Flagellaria indica L.
Goodeniaceae Scaevola sericea Vahl
Gramineae
Chloris barbata Sw. [Furtado 18348]
Digitaria ciliaris (Retz.) Koel. [Furtado 18637]
Digitaria violascens Link [Ridley s.n. 1894]
Eragrostis tenella (L.) P. Beauv. ex R. & S. [Ridley s.n. 1892] Ischaemum muticum L. [Furtado 18625]
Leptaspis urceolata (Roxb.) R. Br. [Ridley 369]
Mnesithea glandulosa (Trin.) Koning & Sosef [Ridley s.n. 1892]
Guttiferae
Calophyllum inophyllum L.
Garcinia eugeniaefolia Wall. ex T. Anders. [Ridley 9488] Garcinia hombroniana Pierre [Ridley 4791]
Garcinia nervosa Miq.
Reference
Ridley 1900 Ridley 1923
Keng 1990 Ridley 1900
Ridley 1900 Blake 1961
Keng 1990 Ridley 1900 Ridley 1900 Ridley 1900 Ridley 1900 Ridley 1900 Ridley 1900
Ridley 1900
Soepadmo 1970
Ridley 1900
Ridley 1900
Ridley 1900
Ridley 1900 Ridley 1900 Ridley 1900 Ridley 1900
Table 2 (Continued)
Species
Labiatae
Hyptis suaveolens (L.) Poit. [Furtado 18344] Ocimum tenuiflorum L. [Furtado 18622]
Lauraceae
Actinodaphne macrophylla (Bl.) Nees [Ridley 9489] Litsea umbellata (Lour.) Merr. [Ridley s.n. 2 Mar 1893] Neolitsea zeylanica Mert.
Phoebe grandis Merr.
Leguminosae
Albizia retusa Benth. [Ridley 4752] Alysicarpus vaginalis (L.) DC.
Canavalia cathartica Thou. [Hullett 463] Crotalaria mucronata Desv. [Furtado 18634] Dalbergia candenatensis (Dennst.) Prain [Ridley 4678] Dalbergia junghuhnii Benth.
Derris heptaphylla (L.) Merr. [Hullett 6194] Derris trifoliata Lour. [Furtado 18346] Pithecellobium ellipticum (Bl.) Hassk. Pongamia pinnata (L.) Pierre [Ridley s.n. 1891]
Loganiaceae
Fagraea auriculata Jack
Fagraea racemosa Jack ex Wall. Malpighiaceae
Tristellateia australasiae A. Rich.
Malvaceae
Abutilon indicum (L.) Sweet
Hibiscus surattensis L. [Furtado 18623]
Thespesia populnea (L.) Sol. ex Correa [Ridley 4624] Melastomataceae
Diplectria viminalis (Jack) O. Ktze.
Pogonanthera pulverulenta Bl. [Hullett 391] Meliaceae
Chisocheton erythrocarpus Hiern
Chisocheton macrophyllus King [Ridley 4767, isosyntype]
Dysoxylum cauliflorum Hiern [Hullett 392] Sandoricum koetjape (Burm. f.) Merr. Menispermaceae
Cyclea laxiflora Miers
Moraceae
Artocarpus dadah Miq. [Ridley 4721] Ficus fistulosa Reinw. ex BI.
Reference
Keng 1990 Ridley 1900 Ridley 1900 Keng 1990
Ridley 1900 Ridley 1900 Ridley 1922
Keng 1990 Keng 1990 Ridley 1900 Ridley 1900 Ridley 1900 Ridley 1900
Ridley 1900 Ridley 1900
Ridley 1900
Ridley 1900 Ridley 1900 Ridley 1900
Keng 1990 Keng 1990
Ridley 1900 Ridley 1900 Ridley 1900 Ridley 1900
Ridley 1900
Ridley 1900 Ridley 1900
Table 2 (Continued) Species
Ficus kerkhovenii Val.
Ficus laevis BI.
Ficus obscura Bl. var. borneensis (Miq.) Corner Ficus pellucido-punctata Griff.
Myristicaceae
Knema glaucescens Jack
Knema globularia (Lamk.) Warb. [Ridley 4817] Myrsinaceae
Aegiceras corniculatum (L.) Blanco
Ardisia singaporensis Ridl. [Ridley 2816, isosyntype] Ardisia villosa Roxb. [Ridley 2809]
Myrtaceae
Eugenia leucoxylon (Korth.) Mig. [Ridley 9486] Tristaniopsis whitiana (Griff.) Wilson & Waterhouse [Ridley 4970] Oleaceae
Olea brachiata (Lour.) Merr.
Ochnaceae Gomphia serrata (Gaertn.) Kanis
Orchidaceae
Bulbophyllum medusae (Lindl.) Rchb. f.
Corymborkis veratrifolia (Reinw.) Bl. [Ridley 2037]
Cymbidium finlaysonianum Lindl. [Goodenough s.n. 1894] Grammatophyllum speciosum BI.
Renanthera elongata Lindl.
Spathoglottis plicata Bl. [Furtado 18621]
Thrixspermum calceolus (Lindl.) Rchb. f. [Goodenough s.n. 23/5/1896] Vanilla griffithii Rchb. f.
Palmae
Caryota mitis Lour. [Goodenough 3148]
Licuala spinosa Wurmb [Ridley 3166]
Nenga pumila (Mart.) Wendl. [Goodenough s.n. 1890] Orania sylvicola (Griff.) H.E. Moore [Ridley 3146] Pandanaceae
Pandanus odoratissimus L. f.
Pandanus parvus Ridley
Passifloraceae
Passiflora foetida L. [Furtado 18626]
Piperaceae Piper caninum Bl.
Rhamnaceae Ventilago malaccensis Ridl.
Reference
Keng 1990 Ridley 1900
Ridley
1900
Ridley 1900
Ridley 1900 Keng 1990
Ridley 1900 Ridley 1900 Ridley 1900
Keng 1990
Ridley Ridley Ridley Ridley
Ridley
Ridley Ridley
Ridley Ridley
Ridley
Ridley
Ridley Ridley
Ridley
1900
1900
1900 1900
1900
Keng 1990
Table 2 (Continued) Species
Rhizophoraceae Bruguiera cylindria (L.) Bl. [Ridley 366]
Rubiaceae
Borreria articularis (L. f.) FN. Will. [Furtado 18339] Borreria laevicaulis (Miq.) Ridl. [Furtado 18343] Gaertnera viminea Hook. f. ex Clarke [Ridley 9500] Ixora congesta Roxb.
Lasianthus cyanocarpus Jack [Ridley 9499]
Morinda umbellata L.
Ophiorrhiza singaporensis Ridl.
Oxyceros longiflora (Lamk.) Yamazaki [Ridley 9487] Psychotria griffithii Hook. f.
Psychotria malayana Jack
Psychotria rostrata Bl.
Uncaria glabrata (Bl.) DC.
Urophyllum streptopodium Wall. ex Hook. f. [Hullett 393]
Rutaceae Clausena excavata Burm. f.
Sapindaceae
Cardiospermum halicacabum L. Lepisanthes rubiginosa (Roxb.) Leenh. [Hullett 386] Mischocarpus sundaicus Bl. [Ridley 9495]
Solanaceae
Datura candida (Pers.) Pasq. — esc. cult. Physalis minima L. [Ridley 367] Solanum torvum Sw. [Furtado 18628]
Sterculiaceae
Pterospermum diversifolium Bl. [Ridley 387] Sterculia coccinea Jack
Turneraceae Turnera ulmifolia L. [Furtado 18633]
Urticaceae
Poikilospermum cordifolium (Borg.-Petr.) Merr. Poikilospermum suaveolens (Bl.) Merr. Verbenaceae
Clerodendrum inerme (L.) Gaertn. Stachytarpheta indica (L.) Vahl [Furtado 18627] Vitex trifolia L.
Zingiberaceae Hornstedtia leonurus (Koenig) Retz. [Ridley 9494]
Reference
Ridley 1900
Keng 1990
Ridley 1900 Ridley 1900 Ridley 1900 Ridley 1900 Ridley 1923
Ridley 1900 Ridley 1900 Keng 1990
Ridley 1900 Ridley 1900
Ridley 1900
Ridley 1900 Ridley 1900
Ridley 1900 Keng 1990
Ridley 1900 Ridley 1900
Ridley 1900 Ridley 1900
Keng 1990
Keng 1990
Holttum 1950
A Botanical Survey of Pulau Ubin 71
Table 3 The five angiosperm families with the most species for the flora of Singapore (after Turner et a/. 1990) and the contemporary flora of Pulau Ubin. Total number of species with number of alien species in brackets.
Singapore Pulau Ubin
Orchidaceae 194 (0) Gramineae 26 (8)
Rubiaceae 147 (7) Leguminosae 25 (10)
Euphorbiaceae 127 (13) Euphorbiaceae 24 (8)
Gramineae 119 (18) Cyperaceae 19 (1)
Cyperaceae 95 (3) Compositae 18 (5) References
Airy Shaw, H.K. (1972). The Euphorbiaceae of Siam. Kew Bull. 26, 191-363. Blake, ST. (1961). Two new species of Scleria. Blumea 11, 219-223.
Corlett, RT. (1990). Flora and reproductive phenology of the rain forest at Bukit Timah, Singapore. J. Trop. Ecol. 6, 55-65.
Holttum, R.E. (1950). The Zingiberaceae of the Malay Peninsula. Gard. Bull. Sing. 13, 1-249.
____. (1981). The fern genus TJectaria Cav. in Malaya. Gard. Bull. Sing. 13, 132-147.
Johnson, A. (1977). A student’s guide to the ferns of Singapore Island, Second Edi- tion, Singapore University Press, Singapore.
Keng, H. (1990). The concise flora of Singapore: gymnosperms and dicotyledons, Singapore University Press, Singapore.
Luerssen, C. (1882). Pteridologische Notizen. Botanisches Centralblatt 11, 26-31.
Markgraf, F. (1972). Florae Malesianae Praecursores LIII. Apocynaceae II. 6. Urnu/aria, 7. Willughbeia, 8. Kopsia. Blumea 20, 407-425.
Ng, PS. (1988). Singapore facts and pictures 1988. Information Division, Ministry of Commuications & Information, Singapore.
Ridley, H.N. (1900). The flora of Singapore. J. Straits’ Branch Roy. As. Soc. 33, 27-196.
____. (1922). The flora of the Malay Peninsula, Volume I, L. Reeve & Co., London. ____. (1923). The flora of the Malay Peninsula, Volume II, L. Reeve & Co., London.
Soepadmo, E. (1970). Florae Malesianae Praecursores XLIX Malesian species of Lithocarpus Bl. (Fagaceae). Reinwardtia 8, 197-308.
Turner, I.M., Chua, K.S. and Tan, H.TW. (1990). A checklist of the native and natural- ized vascular plant species of the Republic of Singapore. J. Sing. Nat. Acad. Sc. 18 & 19, 58-88.
Whitmore, T.C. (1973). Euphorbiaceae. Tree Flora of Malaya, Volume 2, 34-136.
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j 4 THE GARDENS’ BULLETIN SINGAPORE
, VOL. 44 (Part 2) December 1992 4 SS ESTEE ea a ’ 6 " CONTENTS \
' WAH, TT, WEE, Y.C. and PHANG, S.M.: Diatoms from Marine Environments of Peninsular Malaysia 8s 2a; aed va nna Ss stgecptivaers Wied nicinalve Hipw FBSak arias ne has be ne dere ? 73-125
TAN, HTW., IBRAHIM, Ali bin and CHUA, K.S:: f MME FH F107 Of Sitigapore, [vvicic-sensnedes consesrcccodevsdpacsstecsescecesetecencceeness 127-133
' _OGINUMA, Kazuo, LUM, Shawn KY., LEE, Y.H. and TOBE, Hiroshi: Karyomorphology of Some Myrtaceae from Singapore .............::::::eseeeeeeeees 135-139
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THE GARDENS’ BULLETIN SINGAPORE
VOL. 44 (Part 2) December 1992
CONTENTS
WAH, T.I., WEE, Y.C. and PHANG, S.M.: Diatoms from Marine Environments of Peninsular Malaysia Ee tbe ea 2 aeRO SIA ar eee pet pe ae Cone Shi s PEO Pe ne 73-125
TAN, H.-TW., IBRAHIM, Ali bin and CHUA, K:S.: Pannen tC PIGTA Gl SINPADOTE, [ .....c556.fecccas Roe Seeckea Meson del adaaeden esse oeencceees 127-133
OGINUMA, Kazuo, LUM, Shawn K.Y., LEE, Y.H. and TOBE, Hiroshi: Karyomorphology of Some Myrtaceae from Singapore .............:.: cee 135-139
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Diatoms from Marine Environments of Peninsular Malaysia and Singapore
T.T. WAH, Y.C. WEE and S.M. PHANG! Department of Botany, National University of Singapore, Singapore 0511, Republic of Singapore ‘Institute of Advanced Studies, University of Malaya, 59100 Kuala Lumpur, Malaysia
EFFECTIVE PUBLICATION DATE: 15 MAR 1993
Abstract
A total of 230 taxa of diatoms belonging to 58 genera were recorded from 12 locations in Peninsular Malaysia and 14 locations in Singapore. The most common genera are Navicula and Nitzschia.
Introduction
For a long time now the only detailed studies on the marine diatoms of the South- east Asian region have been those by Mann (1925) and Allen & Cupp (1935) on the Philippines Islands and the Java Sea, respectively. More recent studies include those by Takano (1960) who collected from the Arafura and eastern Timor Seas; Wood (1963) who listed many species from the Indonesian waters; Podzorski & Hakansson (1987) on the freshwater and marine diatoms of Palawan, Philippines; and Wah & Wee (1988) on the diatoms of mangrove environments of Singapore and southern Peninsular Malaysia. This paper gives an account on the diatoms of marine environments of Peninsular Malaysia and Singapore.
Materials and Methods
Materials collected for diatoms were from seaweeds obtained from the littoral and sublittoral zones of coastal areas. In addition, sand, molluscs, stones, debris and sediments were also collected. A total of 12 locations in Peninsular Malaysia and 14 locations in Singapore were visited during 1986-87 (Fig. 1). Samples of planktonic diatoms came from the Zoological Raffles Collection of the National University of Singapore. These were collected from the vicinity of Sisters Island in 1968-69. Perma- nent slides were prepared after the method of Gerloff & Natour (1982), details of which are described in an earlier publication (Wah & Wee, 1988). All taxa were iden- tified from the prepared slides using the classical criteria of size, shape and ornamen- tation. Relative abundance of the taxa refers to the abundance of the diatoms in the slides referred to. Details of slide numbers giving collection locations and substrata are tabulated in Table I. Slides were deposited in the Cryptogamic Herbarium of the Botany Department, National University of Singapore.
Results
A total of 230 taxa of diatoms belonging to 58 genera were recorded from 12 loca- tions in Peninsular Malaysia and 14 locations in Singapore. Only five genera were
73
100°E 102° 104°
ss WFO 6° - aoPP oPRa oPK 4 5 oPTe PPa P. Malaysia 4° 3° r d\pt °0 = 9 9 9° O Legs Po Singag bre N
Fig. la. | Map of Peninsular Malaysia showing collecting sites (CR: Cape Rachado; La: Pulau Langkawi; Po: Pontian; PB: Pulau Besar; PD: Port Dickson; PK: Pulau Kapas; PP: Pulau Perhentian; PPa: Pulau Pangkor; PRa: Pulau Redang; PT: Pulau Tioman; PTe: Pulau Tenggul; Sm: Sementa).
‘(YTB ISVOD ISOM ‘AA ‘SENE iL ‘PURIST SIOISIS “1S ‘SUBMEQUIDS 19S fBSOJUIS :G ‘UIQA, Neng ‘Ad ‘INeyT ueJoY Ne[Nd Ad ‘sty JIseg wig ‘suvho7] :o7 *y41ed IOPRIGL’] “T {Ye ISkOD Ise_ -q ‘WUeyeas BURYD :SD ‘IsuRYD :D) Sous ZuNdaTJOO BuIMOYsS WOdeBuIs Jode = “QI “BI
w>Ol O M aye re) S
=
O71 9 dd
Table 1
List of slides giving collection locations and materials collected.
Locations
East Coast Park Changi
Kranji
Loyang
Pulau Ubin Pulau Ubin Pulau Ubin Pulau Ubin Pulau Ubin
East Coast Park Changi
Labrador Park Labrador Park Labrador Park Port Dickson Port Dickson Pasir Ris
Pasir Ris
Pulau Retan Laut Pulau Tioman Pulau Tioman Pulau Tioman Pulau Tioman Pulau Tioman Pulau Tioman Pulau Tioman Changi Sembawang Sembawang
West Coast Park West Coast Park Sentosa
Sentosa
Pontian
Around Sisters Island Around Sisters Island Around Sisters Island Around Sisters Island Around Sisters Island Pulau Kapas Pulau Redang Pulau Pangkor Sementa
Pulau Langkawi Pulau Besar Pulau Tenggul Cape Rachado Pulau Perhentian Tuas
Tuas
Tuas
Tuas
Materials
seaweeds
seaweeds
mollusc
stones
roots and algae mud
sand
reddish sand
rock. 7
seaweeds
seaweeds
red algae
sediment
seaweeds
sediment and sand seaweeds
sand
seaweeds
seaweeds
molluscs
debris
seaweeds
water
sand
black sediment pole
seafarm debris foam
seaweeds
seaweeds
sand
sand
submerged grasses sandy mud surface pumping, 16.4.68 bottom pumping, 16.4.68 surface towing, 21.5.68 surface pumping, 14.1.69 bottom pumping, 14.1.69 seaweeds
seaweeds
seaweeds
red algae
water sample seaweeds
coral
seaweeds
seaweeds
sand
seaweeds filamentous algae debris, stone
4: re
‘ ; '
Marine Diatoms of Peninsular Malaysia and Singapore 77
represented by ten or more taxa, and these were Amphora (14), Diploneis (21), Navicula (29), Nitzschia (20) and Pleurosigma (11). The most common genera were Navicula and Nitzschia, the former represented by 27 species and the latter by 19 species.
Systematics
For the sake of convenience, the list of taxa presented here is arranged alphabet- ically by genus with species and their varieties listed alphabetically within each genus. Figures are given for all taxa.
Achnanthes Bory 1822
A. brevipes var. intermedia (Kutz.) Cl. Figs. 2-3 References: Cleve 1894-95, 27(3), p. 193; Hustedt 1959, 7(2), p. 425, figs. 877d, e. Description: Length 25-42 yum, breadth 9-10 um, 8-10 striae in 10 pm. United into short filament.
Distribution: Very common. Found on slides 1, 135, 257, 339, 347, 353. Comments: A cosmopolitan species.
A. hauckiana Grun. Figs. 4-5 References: Cleve 1894-95, 27(3), p. 190; Patrick & Reimer 1966, p. 267, pl. 17, figs. 25-32.
Description: Length 16-25 wm, breadth 8-10 ym, 7-10 striae in 10 pm. Distribution: Common. Found on slides 125, 353.
Comments: A cosmopolitan species. Freshwater to brackish.
A. lewisiana Patr. Figs. 6-7 References: Patrick & Reimer 1966, p. 266, pl. 17, figs 19, 20.
Description: Length 14 pm, breadth 6 um, 15 striae in 10 pm.
Distribution: Common. Found on slides 273, 343, 353, 355.
Comments: Freshwater species.
A. longipes Ag. Figs. 8-9 References: Cleve 1894-95, 27(3), p. 195; Hendey 1964, p. 174, pl. 28, figs. 1-6; pl. 42, fig. 2.
Description: Length 62-69 ym, breadth 11-14 um, 6-7 costae and 9-10 striae in 10 um. Solitary or joined into filament; attached by mucous stipe.
Distribution: Very common. Found on slides 12, 116.
Comments: A cosmopolitan species.
Actinocyclus Ehrenb. 1838
A. ehrenbergii var. sparsa (Greg.) Hust. Fig. 10 References: Hendey 1964, p. 84; Foged 1984, p. 15, pl. 19, fig. 3. wae Description: Diameter 37-66 ym, 5-6 areolae and 9-11 marginal striae in 10 um. Distribution: Common. Found on slides 48, 119, 124, 138.
Comments: A cosmopolitan species.
A. octonarius Ehrenb Fig. 11
References: Hendey 1964, p. 83, pl. 24, fig. 3; Priddle & Fryxell 1985, p. 108. Description: Diameter 60 pm, 7 areolae and 16 marginal striae in 10 pm.
78 Gard. Bull. Sing. 44(2) (1992)
Distribution: Not common. Found on slides 95, 339. Comments: A cosmopolitan species.
A. platensis Mill. Melchers. Fig. 12 References: Hendey 1958, p. 43, pl. 5, figs. 1, 2.
Description: Diameter 56-77 um, 20 marginal striae in 10 pm.
Distribution: Common. Found on slide 3.
Actinoptychus Ehrenb. 1839
A. senarius (Ehrenb.) Ehrenb. Fig. 13 References: Hendey 1964, p. 95, pl. 23, figs 1 and 2; Priddle & Fryxell 1985, p. 110, fig. A.
Description: Diameter 39 pm.
Distribution: Not common. Found on slides 4, 79, 116, 323, 343.
Comments: A cosmopolitan species.
Amphora Ehrenb. 1840
A. acutiuscula Kitz. Figs. 14-15 References: Prowse 1962, p. 55, pl. 17, figs. e-f, n, q, v-w; pl. 18, fig. b; Patrick & Reimer 1975, p. 77, pl. 14, figs. 9, 10.
Description: Length 35-55 ym, breadth 6-13 wm, 11-12 dorsal, 18-20 ventral striae in 10 pm.
Distribution: Common. Found on slides 12, 17, 43, 297.
Comments: A freshwater species.
A. angusta var. eulensteinii Grun. Fig. 16 References: Cleve 1894-95, 27(3), p. 135.
Description: Length 81 yum, breadth 15 ym, 11 dorsal and 14 ventral striae in 10 pm. Distribution: Not common. Found on slides 100, 357.
Comments: A cosmopolitan species.
A. angusta var. oblongella Grun. Fig. 17 References: Cleve 1894-95, 27(3), p. 135.
Description: Length 52-57 ym, breadth 8-10 ym, 12-15 striae in 10 um. Distribution: Rare. Found on slides 183, 353.
A. angusta var. ventricosa (Greg.) Cl. Fig. 18 References: Cleve 1894-95, 27(3), p. 135; Navarro 1982, p. 31, pl. 20, figs. 1-2. Description: Length 49-58 ym, breadth 10-13 ym, 14-15 striae in 10 pm. Distribution: Common. Found on slides 335, 355.
A. coffeiformis (Ag.) Kitz. Fig. 19 References: Prowse 1962, p. 56, pl. 17, figs. h, o.
Description: Length 30-33 ym, breadth 6-7 wm, 18-20 striae in 10 pm. Distribution: Not common. Found on slides 1, 257, 335. Freshwater to brackish. Comments: A cosmopolitan species.
ese: 't /
Marine Diatoms of Peninsular Malaysia and Singapore 79
A. crassa Greg. Fig. 20
References: Hendey 1964, p. 262. Foged 1975, p. 10, pl. 36, fig. 10.
Description: Length 132 um, breadth 18 um, 5 dorsal, 6 ventra costae in 10 ym. Distribution: Rare. Found on slide 93.
Comments: A cosmopolitan species.
A. decipiens Cl. Fig. 21 References: Cleve 1894-95, 27(3), p. 108, pl. V, figs. 16-18.
Description: Length 41 pm, breadth 17 wm, 12 striae in 10 um. Distribution: Rare. Found on slide 12.
A. graeffi var. minor Perag. Fig. 22
References: Hendey 1964, p. 263, pl, 37, fig. 8. Description: Length 38 um, breadth 9 um, 11-12 striae in 10 pm. Distribution: Rare. Found on slides 2, 12.
A. holsatica Hust. Figs. 23-26
References: Caljon 1983, p. 119, pl. 21, figs. 2-8.
Description: Length 18-42 um, breadth at girdle view 8-14 um and at valve view 5 um, 12-20 striae in 10 pm.
Distribution: Very common. Found on slides 1, 12, 122, 257, 259, 273, 335, 337, 339, 341, 343, 347, 350, 353.
Comments: A cosmopolitan species. Freshwater to brackish.
A. ostrearia Bréb Fig. 27
References: Cleve 1894-95, 27(3), p. 129.
Description: Length 58 ym, breadth 15 in valve view, 38 um in girdle view, 11-12 striae in 10 pm.
Distribution: Rare. Found on slide 38.
Comments: A cosmopolitan species. Freshwater to marine.
A. proteus Greg. , Fig. 28 References: Prowse 1962, p. 58, pl. 17, fig. u; pl. 18, fig. a; Cleve 1894-95, 27(3), p. 103. Description: Length 56 um, breadth 8 in valve view, 20 um in girdle view, 9-12 striae in 10 pm.
Distribution: Rare. Found on slide 3.
Comments: A cosmopolitan species.
A. turgida Greg. Fig. 29 References: Cleve 1894-95, 27(3), p. 123; Hendey 1964, p. 264.
Description: Length 32 um, breadth 9-10 um, 15 striae in 10 um.
Distribution: Rare. Found on slides 273, 333, 341, 351.
Comments: A cosmopolitan species. Marine to brackish.
A. valida Perag. Fig. 30 References: Cleve 1894-95, 27(3), p. 102. <a . . Description: Length 82 um, breadth 16-18 «wm in valve, 50 «wm in girdle view, 7 striae in 10 pm.
Distribution: Rare. Found on slide 124.
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Figs. 2-29
hauckiana, 9. A. longipes,
raphe valve and girdle view. Fig. 10. Actinocyclus ehrenbergii var. sparsa. Fig. 11. A. octonarius. Fig. 12. A.
5. A
Figs. 2-3. Achnanthes brevipes var. intermedia, raphe valve and rapheless valve. Figs. 4-
raphe valve and rapheless valve. Figs. 8-
7 A. lewisiana
raphe valve and rapheless valve. Figs. 6-
16. A. angusta vat. var. minor. Figs. 23-26. A. holsatica, Fig, 274mm
15. Amphora acutiuscula. Fig.
eulensteinii. Fig. 17. A. angusta var. oblongella. Fig. 18. A. angusta var. ventricosa. Fig. 19. A. coffeiformis.
14-
13 Actinoptychus senarius. Figs. 20. A. crassa. Fig. 21. A. decipiens. Fig. 22. A. graeffi
platensis. Fig.
Fig.
/
i
ostrearia. Fig. 28. A. proteus. Fig. 29. A. turgida.
Marine Diatoms of Peninsular Malaysia and Singapore 81
A. wisei (Salah) Simonsen. Fig. 31
References: Foged 1975, p. 12, pl. 26, fig. 11.
Description: Length 14 um, breadth 5 um, 16 striae in 10 pm. Distribution: Rare. Found on slide 273.
Comments: A cosmopolitan (?) species.
Asterionella Hassall 1855
A. japonica Cl. & MOIll. Fig. 32 References: Cupp 1943, p. 188, fig. 138; Hendey 1964, p. 158, pl. 21, fig. 1.
Description: Length 92-104 ym, inflated length 22-24 um, inflated breadth 11-12 um. Distribution: Not common. Found on slide 321.
Bacillaria Gmelin 1788
B. paradoxa var. tumidula Grun. Fig. 33
References: Navarro 1982, p. 51.
Description: Length 106-149 um, breadth 10-13 ym, 15-20 striae, 7-9 keel puncta in 10 pm.
Distribution: Common. Found on slide 2.
Comments: A cosmopolitan species. Freshwater to marine.
Bacteriastrum Schadbolt 1853
B. delicatulum Cl. Fig. 34 References: Gran & Angst 1931, p. 463, fig. 46a-—b; Hendey 1964, p. 139, pl. 6, fig. 2. Description: Diameter 16 ym, 9 bristles. United to form chains.
Distribution: Not common. Found on slide 317.
Comments: Common in temperate waters.
B. elongatum Cl. Fig. 35 References: Cupp 1943, p. 99, fig. 57; Hendey 1964, p. 139, pl. 6, fig. 3. Description: Diameter 14 um, 9 bristles.
Distribution: Not common. Found on slide 339.
Comments: Common in tropical seas.
B. hyalinum Lauder. Fig. 36 References: Cupp 1943, p. 96, fig. 56(A); Hendey 1964, p. 139, pl. 6, fig. 1. Description: Diameter 24-39 ym, 14-27 bristles. United to form chains. Distribution: Common. Found on slide 317.
Biddulphia Gray 1821
B. biddulphiana (Smith) Boyer (B. pulchella Gray.). Figs. 40-42 References: Hendey 1964, p. 101, pl. 25, fig. 1.
Description: Length 76-101 um, breadth 69-82 ym in pervalvar axis; length 77-90 «um, breadth 38-58 pm in top view. Colonial, united with their processes to form short chains. Distribution: Common. Found on slides 12, 355.
Comments: A cosmopolitan species.
82 Gard. Bull. Sing. 44(2) (1992) 7
B. mobiliensis (Bailey) Grun. Fig. 37 References: Gran & Angst 1931, p. 490, fig. 74; Cupp 1943, p. 153, fig. 110; Hendey 1964, p. 104, pl. 22, fig. 3.
Description: Length 69 um in apical axis, breadth 44 wm, 11 striae in 10 pm. Solitary or in chains.
Distribution: Rare. Found on slide 323.
B. petitiana (Leud.-Fortm.) Mann. Figs. 38-39
References: Mann 1925, p. 43, pl. 10, figs. 4, 5. Description: Length 70 um in pervalvar axis; length 66 »m, breadth 38 pm in top view. Distribution: Rare. Found on slide 75.
B. recticulata Roper. Fig. 43
References: Boyer 1926-27, p. 128, Hendey 1958, p. 48.
Description: Length 88-96 wm, breadth 68-96 um in pervalvar axis (girdle view). Distribution: Rare. Found on slide 333.
Comments: A cosmopolitan species (?).
B. vesiculosa (Ag.) Boyer. Fig. 44
References: Lebour 1930, p. 181, pl. 4, fig. 1. Description: Length 91-104 y~m, breadth 52 um in pervalvar axis (girdle view). Distribution: Rare. Found on slides 4, 278.
Caloneis Cleve 1891
C. alpestris (Grun.) Cl. Fig. 45
References: Patrick & Reimer 1966, p. 587, pl. 54, fig. 9. Description: Length 73 ym, breadth 15 yum, 20 striae in 10 pm. Distribution: Rare. Found on slide 12.
C. bacillum (Grun.) Cl. Fig. 46 References: Cleve-Euler 1955, 5(4), p. 102, fig. 1147 a-c; Patrick & Reimer 1966, p. 586, pl. 54, fig. 8.
Description: Length 21-25 ym, breadth 10-12 ym, 21-23 striae in 10 pm. Distribution: Not common. Found on slides 341, 350.
Comments: A cosmopolitan species. Freshwater.
C. egena (A. Sch.) Cl. Fig. 47
References: Cleve 1894-95, 26(2) p. 66; Foged 1984, p. 24, pl. 44, fig. 10. Description: Length 25 um, breadth 6 pm. Distribution: Rare. Found on slide 345.
C. liber (W. Sm.) Cl. Fig. 48
References: Cleve 1894-95, 26(2) p. 54; Hendey 1964, p. 229, pl. 29, fig. 2. Description: Length 39 ym, breadth 11 pm, 18 striae in 10 pm. Distribution: Rare. Slide 31.
C. linearis (Grun.) Boyer. Fig. 49
References: Hendey 1964, p. 230, pl. 29, fig. 3; Foged 1984, p. 26, pl. 44, figs. 4,5.
Figs. 30-40 (horizontal common scale bar and those of Figs. 33, 36 and 40 = 10 um)
Fig. 30. Amphora valida. Fig. 31. A. wisei. Fig. 32. Asterionella japonica. Fig. 33. Bacillaria paradoxa var. tumidula. Fig. 34. Bacteriastrum delicatulum. Fig. 35. B. elongatum. Fig. 36. B. hyalinum. Fig. 37. Bid- dulphia mobiliensis, girdle view. Figs. 38-39. B. petitiana, valve view and girdle view. Fig. 40. B. biddulphiana, valve view.
84 Gard. Bull. Sing. 44(2) (1992)
Description: Length 48 um, breadth 8 pm. Distribution: Not common. Found on slide 345. Comments: A cosmopolitan species.
Campylodiscus Ehrenb. 1841
C. fastuosus Ehrenb. Fig.
References: Hendey 1964, p. 290, pl. 40, fig. 13. Description: Diameter 54-68 pm. Distribution: Not common. Found on slides 141, 339, 343, 345, 357.
Comments: Common on sandy beaches of all North Sea coasts (Hendey, 1964).
C. hypodromus Brun & Tempeére. Fig.
References: Hendey 1964, p. 291. Description: Diameter 62-113 um, 2-3 costae in 10 pm. Distribution: Rare. Found on slide 345.
C. ralfsii W. Sm. Fig.
References: Hendey 1970, p. 161, pl. 5, fig. 53; 1964, p. 291. Description: Diameter 28-56 wm, 2-3 costae in 10 wm. Distribution: Not common. Found on slides 79, 345, 347.
Chaetoceros Ehrenb. 1844
C. danicum Cl. Fig.
References: Hendey 1964, p. 122, pl. 10, fig. 5. Description: Diameter 25-31 ym. Solitary or in chains. Distribution: Common. Found on slide 317.
C. lorenzianum Grun. Fig.
References: Gran 1905, p. 76, fig. 90; Hendey 1964, p. 124, pl. 26, fig. 1. Description: Diameter 17 ym. Solitary or in chains.
Distribution: Not common. Found on slide 323.
Comments: A cosmopolitan species.
C. peruvianum Brightw. Fig.
References: Hendey 1964, p. 123, pl. 9, fig. 3; Priddle & Fryxell 1985, p. 40. Description: Diameter 30 pm. Solitary or in chains.
Distribution: Not common. Found on slide 323.
Comments: A cosmopolitan species.
C. tetrastichon Cl. Fig.
References: Hendey 1964, p. 123, pl. 11, fig. 1. Description: Diameter 16 ~m. United to form chains. Distribution: Rare. Found on slide 323.
Climacosphenia Ehrenb. 1841
C. moniligera Ehrenb. Figs. 57-
References: Cupp 1943, p. 178, fig. 128.
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