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—(. 94 102/ ISSN 0374-7859
DEL \ Wo! ;
THE GARDENS’ BULLETIN
SINGAPORE
Volume 39
(1986)
A periodical reflecting the interests and
activities of the Botanic Gardens
Singapore
Published by the Botanic Gardens
Parks and Recreation Department
Ministry of National Development
Cluny Road, Singapore 1025.
CONTENTS
Volume 39
PART 1 — Ist June 1986
WILDE. W.J.J.O. DE:
A New Account of the Genus Horsfieldia (Myristicaceae). Pt 4 ......... cece e eee es
KENG, Hsuan:
Peden fis! OF SCC F Ianis Or SINGAPOTe (A)... 5... cf... o. 2. ete ee ene vagy nnetdecsesececeerees
FOONG, T.W. and C.N. YANG:
The Evaluation of the Effectiveness of Water-absorbent Polymers in Reducing the
USO Pe USS Be SO Be ee tb iy cl: en end foe ee en ee
PART 2 — list December 1986
CORNER: E/3°H.:
The Agaric Genus Panellus Karst. (including Dictyopanus Pat.) in Malaysia
BIDIN, Aziz & JAMAN, RAZzatt:
A New Species of Platycerium from Peninsular Malaysia ......................s.eceeeeeeeeeeee ees
HOLTTUM, R.E.:
STUDIES IN THE FERN-GENERA ALLIED TO Tectaria Cav. VI: A conspectus of genera in the
Old World regarded as related to Tectaria, with descriptions of two genera .................
TINDALE, Mary D.:
A New Genus and Three New Species of Pteridophytes from North Eastern
Queensland
KOSTERMANS, A.J.G.H.:
Notes on Asiatic Cassine L. (Celastraceae)
STONE, BENJAMIN C.:
The Genus Pandanus (Pandanaceae) on Christmas Island, Indian Ocean ...................
WONG, Yew Kwan:
The Use of Tifgreen and Tifdwarf Bermuda Grasses in two Singapore Golf Courses
ec
Pages
67-95
97-101
103-147
149-151
169-175
177-191
193-202
203-214
215-219
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INDEX
Volume 39
Basionyms and synonyms appear in italics. Page numbers in italics indicate the presence of illustra-
tions.
Acanthus ebracteatus 82
volubilis 82
Adenosma capitatum 75
indicum 75
inopinatum 75
javanicum 75
ovatum 75
Aenigmopteris 158, 162
Aeschynanthus parviflora 78
purpurascens 78
radicans 78
wallichii 78
Aleurodiscaceae 104
Amanita 143
Amauroderma 109, 119, 112
Ambullia sessiliflora 75
Amphiblestra 164
Andrographis paniculata 82
Angelonia salicariaefolia 75
Aniseia martinencis 67
Anisomeles indica 93
ovata 93
Arenga 133
Argyreia ridleyi 68
Arrabidaea magnifica 80
Aspidiaceae
Aspidium lanuginosum 161
Asystasia coromandeliana 83
gangetica 83
intrusia 83
nemorum 83
Atalopteris 164, 165
Ataxipteris 160, 161, 162, 164
Avicennia alba 88
intermedia 88
lanata 88
marina 88
officinalis 88
Bacopa monnieri 75
Baeospora 143
Barleria cristata 83
lupulina 83
Beloperone guttata 83
Boletus 109, 110
Bonamia semidigyna 68
Bonnaya brachiata 76
reptans 76
veronicaefolia 76
Browallia speciosa 72
Brunsfelsia americana 72
calycina 72
Calamus
Calliaspidia guttata 83
Callicarpa longifolia 88
Calophanoides quadrifaria 83
Campanella 129
Camptodium 164
Capsicum annuum 72
var. acuminatum 72
var. cerasiforme 72
var. grossum 72
Carmona retusa 71
Cassine L. 178, 181, 185
albens comb. nov. 178, /79; 181, 183, 185
australe 188
balae sp. nov. 180, 181, 183, 185, 187
congylos sp. nov. 185, 187
elliptica 188, 189
glauca (Rottb.) O.K. 177, 178; 180, 181, 782,
185, 187
grossa comb. nov. 177, 178, 181
koordersii sp. nov. 186, 188, 191
obiensis 188, 191
paniculata 178, 181, 183
roxburghii 181, 183
viburnifolia 185
Cassine, Vernacular Names: Nerija 180,
Helmedija (Waelmedija) 183; Kurativa 186;
Kurubu vachchi 186, Nareloo, Neraloo 186,
Perun 186, Piyaree, 186
Celastrus glaucus Vahl 181, 183
glaucus Auct. (non Vahl) 181
Centranthera humifusa 75
tranquebarica 75
Cestrum nocturnum 72
Chlamydogramme gen. nov. 157, 164
hollrungii comb. nov. /56, 157
Cionidium 157
Citharexylum quadrangulare 88
spinosum 88
Clerodendrum deflexum 89
disparifolium 89
fragrans 89
indicum 89
inerme 89
laevifolium 89
myrmecophilum 89
nutans 89
paniculatum 89
penduliflorum 89
philippinum 89
var. pleniflorum 89
phyllomega var. myrmecophyllum 89
siphonanthus 89
thomsonae 89
villosum 89
Clitocybula 143
Coleus atropurpureus 95
blumei 95
parviflorus 95
tuberosus 95
Collybia 104, 110
Congea velutina 90
Cordia cylindristachys 71
215
216
dichotoma 71
obliqua 7\
subcordata 71
Coveniella gen. nov. 159, 160, 169
poecilophlebia (Hook.) Tind. comb. nov. 169
Crossandra infundibuliformis 83
undulaefolia 83
Ctenitis 153, 158, 159, 160, 162, 164, 165, 170
canacae comb. nov. 161
submarginalis 165
Ctenitop. submarginalis group 159
Cuscuta australis 68
hygrophilae 68
Cyclopeltis 160, 162
Cyphomandra betacea 73
Cyrtandra pendula 78
Cyrtanthera carnea 85
Datura candida 72
metel 73
Deplanchea bancana 79
Dictyopanus 104-107, 113, 124
copelandii 131
foliicolus 112
gloeocystidiatus 104, 131
luminescens 132
orientalis 110, 135
pusillus 104, 136
var. pseudorhipidium 104
var. sublamellatus 104
rhipidium 136
Dictyoxiphium 157, 164
Didymocarpus perdita 78
platypus 78
Diplazium assimile 174
queenslandicum sp. nov. 174, /74
Dipteracanthus repens 86
Dolichandrone spathacea 80
Dryopsis 153
Dryopteris 158
Dryopteris group 153
Dryopteris subg. Ctenitis 158, 162
Dryopteris protensa 158
subcincisa 158
Duranta repens 90
Dysophylla auricularia 95
Ehretia buxifolia 71
microphylla 71
Elaeodendron Jacq.f. 178
Elaeodendron (non O.K.) 178
glaucum (non O.K.)) 181
glaucum (Rottb.) Pers. 178, 183, 185
var. macrocarpum 186
var. montanum 185
var. roxburghii 183
glaucum sensu Roxb. 180, 183
paniculatum 180, 183
roxburghii 177, 180, 183
Elaeodendrum Murray 178
Endocomia 60
Endopogon ridleyi 83
Entoloma 109, 110
Episcia cupreata 79
fulgida 79
Gard. Bull. Sing. 39 (1986)
Eranthemum nervosum 84
pulchellum 84
Erycibe festiva 68
griffithi 68
leucoxyloides 68
maingayi 68
malaccensis 68
Euonymus grossa 178
Evolvulus alsinoides 68
Favolaschia 104, 105, 110, 111
minima 131
Favolaschiaceae 104
Fadyenia
Fittonia verschaffelti 84
var. argyroneura 84
Ganoderma 112
Gendarussa vulgaris 84
Gmelina asiatica 90
elliptica 90
hystrix 90
villosa 90
Golf course grasses:
Axonopus compressus 209, 211
Bermuda grass, Santa Ana 203
Bermuda grass, Tifgreen 203, 204, 205, 206,
209, 214
Bermuda grass, Tifdwarf 203, 204, 205, 206,
209, 214
Borreria setidens 209, 2/3
Bracharia distachya 207, 209, 210
Curvularia, pathogen 214
Cynodon crosses:
dactylon 203
transvaalensis 203
Cynodon x magennisii 203
incompletus var. hirsutus 203
(See also under Bermuda grass)
Cyperus kyllingia 207, 209, 210
radians 208, 209, 214
Desmodium triflorum 209
Digitaria didactyla 209, 2/0
setigera (D. marginata) 209, 2/2
Eleusine indica 209, 2/2, 214
Eragrostis 209
Euphorbia hirta 209, 2/3
thymifolia 209, 2/3
Ischaemum spp. 209
Panicum spp. 209
Graptophyllum pictum 84
Gymnopteris decurrens 154
holrungii 157
Harmsiopanax aculeatus 60
Heliotropium indicum 71
Heimiella 112
Hemigramma 154, 157
latifolia 154, 157
x Tectaria crenata 154
decurrens 154
holrungii 157
siifolia 156
Hemigraphis alternata 84
colorata 84
Herpestis monnieri 75
Heterogonium pinnatum 166
sagenioides 166
Index to Volume 39
Heterogonium Pres] emend. Holttum 157. 158
pinnatum x Tectaria aurita 158
Hippocrates indica 183
Holmskioldia sanguinea 90
Horsfieldia, Index of Names 62-65
excluded species 60-61
Hydrolea zeylanica 70
Hygrophila erecta 84
quadrivalvis 84
meianthos 84
phlomoides 84
quadrivalvis 68
spinosa 85
Hygrophorus 110
Hyptis brevipes 93
capitata 93
suaveolens 94
Ipomoea alba 68
aquatica 68
batatas 69
bona-nox 68
digitata 69
gracilis 69
horsfalliae 69
illustris 69
littoralis 69
pes-caprae ssp. braziliensis 69
pinnata 69
pulchella auct. 69
quamoclit 69
reptans 68
triloba 70
tuba 70
Jacaranda filicifolia 80
Jacobinia carnea 85
coccinea 85
Justicia betonica 85
gendarussa 84
quadrifaria 83
vasculosa 85
Lantana aculeata 90
camara 90
var. aculeata 90
Lastrea 161
Lastreopsis 158, 159, 162, 170
grayi 170
tenera 172
tinarooensis sp. nov. 170, / 72
walleri sp. nov. 172, /73
Lentinus 146
Leonitis nepetaefolia 94
Leonurus sibiricus 94
Leptochilus pentagonalis 154
stifolius 157
Leucas lavandulifolia 94
linifolia 94
zeylanica 94
Lettsomia ridleyi 68
Limnophila laxa 75
sessiliflora 75
Lindernia anagallis 76
antipoda 76
crustacea 76
ruellioides 76
sessiflora 76
viscosa 76
Lippia nodiflora 91
Lycium chinensis 73
Lycopersicum esculentum 73
var. cerasiforme 73
Mangifera glauca Rottb. 180, 181, 183, /84
Marasmiellus 143
Marasmius 104, 110, 117, 118, 139
Mazus rugosus 76
Megalastrum gen. nov. 158, 159. 161*
canacae comb. nov. 161
exaggeratum comb. nov. 161
lanatum comb. nov. 161
lanuginosum comb. nov. 161, /63
magnum comb. nov. 161
villosum (L.) Holttum comb. nov. 161, /63
Mentha arvensis 94
spicata 94
Merremia caespitosa 70
convolvulacea 70
hederacea 70
Microcarpaea minima 76
muscosa 76
Mycena 139, 143
Myristica brachiata 3
mayjuscula 49
Neerija 177
dichotoma 180; 181, 183
Nephrodium crinitum var. exaggeratum 161
magnum 161
Nicoteba betonica 85
Nicotiana tabacum 73
Nothoperanema 153
Ocimum americanum 94
basilicum 94
canum 94
tenuiflorum 94
Omphalina 143
Orthosiphon aristatus 95
stamineus 95
Oudemansiella 112
Pachystachys coccinea 85
Pajanelia longifolia 80
multijuga 80
Pandanus,
subg. Lophostigma sect. Brongniarta 199
subg. Loph sect. Barrotia 199
subg. Pandanus, sect. Pandanus 199, 200
subg. Rykia 201
subg. Rykia sect. Roussinia 199, 202
subg. Ryk sect. Asterodontia 199
christmatensis 195, 196, 196, 198, 199, 199,
201, 201
elatus 195, 195, 196, 197, 20], 201, 202
intraconicus 200, 201
kirkii 202
leram 190, 190, 20/, 201
var. andamanensium 199
navitatis 195
odoratissimus 198, 202
platycarpus 200
tectorius 198, 199, 200, 202
Pandorea pandorana 80
Panellus, Index to Taxa 147
Panellus 104-16, 110-112, 113, 117, 118, 124,
218
Panellus
127, 139, 143
key to subgenera 114
subgen. Megalopanellus subgen. nov. 104,
109, 110, 143
subgen. Megalopanus 110
key to species 139
subgen. Mesopanellus subgen. nov. 104, 109,
110, 139
subgen. Mitellus 110, 111, 139
subgen. Panellus 104, 109, 110, 111, 112, 114
key to lamellate species 114-115
key to poroid species 128
albifavolus sp. nov. 108, 111, 112, 128, 729
alutaceous sp. nov. 108, //5, 115
ambiguus sp. nov. 106, 111, 116
aureofactus 104, 106
bambusarum sp. nov. 16, 110, //7, 117
bambusifavolus sp. nov. 106, 108, 129, /30,
132-133
brunneifavolus sp. nov. 106, 133, 130
brunneomaculatus sp. nov. 106, 111, 118
copelandii 106, 131
cystidiosus 111
dichotomus sp. nov. 104, 106, 110, 118, 179,
120
var. dichotomus 120
var. pinnatus var. nov. 120
?diversipes 124
dumonti 110
exiguus sp. nov. 106, 108, 120, /2/
fuscatus sp. nov. 108, 121, 722, 139
gloeocystidiatus comb. nov. 106, /07, 109,
12; 173,130; 131
glutinosus sp. nov. 135, 139, 140
hispidifavolus sp. nov. 108, /32, 132
intermedius sp. nov. 106, 110, 123
var. Stenoporus var. nov. 124
longinquus 108, 124
luminescens comb. nov. 106, /07, 112, 1/3,
1245132
magnus sp. nov. 110, 123, 1/42, 143, 144, 145
megalosporus sp. nov. 106, 111, 112, 7/3, 133,
155
microsporus sp. nov. 106, 108, 110, 111, 112,
113, 134, 134
mirabilis 127
nubigenus 126
orientalis comb. nov. 106, 111, 772, 112, 1/3,
134,435,135
parvulus sp. nov. 106, 110, 117, 125, 125
pauciporus sp. nov. 106, 136
pendens sp. nov. 108, 126
pusillus 106, 107, 132, 136
pyruliferus sp. nov. /40, 141
serotinus 110
stenocystis 118
stypticus 104, 106, 112, 120
sublamellatus 100
sublamelliformis sp. nov. 106, 137, 137
sublevatus sp. nov. 106, 110, 127, 127
Peronema 153
Perilepta dyeriana 86
Peristrophe acuminata 85
roxburghiana 85
tinctoria 85
Gard. Bull. Sing. 39 (1986)
Peronema canescens 90
Petraea volubilis 91
Petunia axillaris x P. interifolia 73
hybrida 73
Phegopteris lanata 161
Phyla nodiflora 91
Pinanga 133
Platycerium coronarium 149, /50, 151
holttumii 149
platylobum sp. nov. 149
ridleyi 149, 151
Plectranthus scutellarioides 95
Pleocnemia 159, 160, 162, 164
Pleuroderris 164
Pleurotus 104, 113.
decipiens 146
Pluteus 109, 110, 143
Pogostemon auricularius 95
Polypodium grande 161, 164
poecilophlebium 160, 169, 171
villosum L 161
Polyporus 111
Porana volubilis 70
Porpholoma 143
Premna angustior 91
corymbosa 91
foetida 91
integrifolia 91
parasitica 91
punctulata 91
ridleyi 91
trichostoma 91
Paeuderanthemum kingii 85
reticulatum 86
Pseudotectaria 160, 162
crinigera 162
decaryana 162
Psomiocarpa 158, 162
Pteridrys 160, 162
Quamoclit pinnata 69
Quercifilix 154, 157
Radermachera gigantea 80
lobbii 80
Rhapis 133
Rhododendron 146
Roussinia indica 20/, 201
Ruellia amoena 86
repens 86
tuberosa 86
Russelia equisetiformis 77
Juncea 77
Sagenia 157
venation 154
Saintpaulia ionantha 79
Salvia coccinea 95
Sanchezia nobilis 86
Saritaea magnifica 80
Schematiza cordiae (a beetle) 71
Scoparia dulcis 77
Sesamum orientale 81
indicum 81
Sinningia speciosa 79
Solanum ferox 73
mammosum 74
melongena 74
nigrum 74
Index to Volume 39
torvum 74
tuberosum 74
wrightii 74
Spathodea campanulata 80
Sphenodesma pentandra 91
Stachytarpheta indica 91
jamaicensis 92
mutabilis 92
Staurogyne griffithiana 86
setigera 86
Stenolobium stans 81
Stenostemia 157
Stephanophysum longifolium 86
Stereospermum chelonoides 80
fimbriatum 80
personatum 80
Striga asiatica 77
hirsuta 77
Strobilanthes dyerianus 86
Tabebuia chrysantha 81
pallida 81
rosea 81
stans 81
Tecomaria capensis 81
Tectaria 153, 154, 157, 158, 159, 160, 162, 164,
165
Tectaria, allies 164
Tectaria group 153, 154
Tectaria & closely allied genera 154
key to the paleaotropic genera 165-166
Sect. Tectaria 154, 157, 158, 162, 164
Sect. Sagenia stat. nov. 157, 158, 159, 162, 164
aurita 158
brauniana 157, 165
decaryana 162
decurrens 154
dicksonioides 160
fuscipes 154
incisa 164
hilocarpa 154, 155
ingens 157
pedata 157
Tectaridium 157
Tectona grandis
Teijsmanniodendrom coriaceum 92
pteropodum 92
Thelypteridaceae 170
Thunbergia affinis 86
alata 87
erecta 87
fragrans 87
grandiflora 87
laurifolia 87
Torenia polygonoides 77
Tournefortia tetrandra 71
wallichii 71
Tricholoma 104
Triplophyllum 158, 160, 162
dicksonioides 165
Tristania 132
Trogia 143, 146
Utricularia albina 77
bifida 77
caerula 77
exoleta 77
fluitans 77
griffithii 77
punctata 77
Vandellia crustacea 76
hirsuta 76
pedunculata 76
sessiliflora 76
Vitex coriacea 92
negundo 92
pinnata 92
pteropoda 92
pubescens 92
tenera 92
trifolia 93
vestita 93
Water absorbent polymers 97-1()1
as soil conditioners 97
as water-holding additives 97
composition of planting mix 98, 99
index for plant-available water 98
on Hibiscus rosa-sinensis 98, 99, 100
on Mussaenda Dona Luz 98, 100
on Tecomaria capensis 98, 99, 100
plants susceptible to water stress 98
soil mixes used 98
their structure 97
total water-holding capacity and plant mix 99
DTW 98, 99, 100
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-
THE GARDENS’ BULLETIN
fi SINGAPORE
VOL. 39 (Part 1) Ist June 1986 |
CONTENTS
re
PAGES
WILDE, W. J.J.O. bE:
A New Account of the Genus Horsfieldia (Myristicaceae), Pt 4 ..............c ccc eceeeeeeeeeee 1-65
) KENG, Hsuan:
N Peneesice List of Sced Plants of Singapore.(X) 5.62) ani nde bos oot ovecsnececseceeeee 67-95
x FOONG, T.W. and C.N. YANG:
X The Evaluation of the Effectiveness of Water-absorbent Polymers in Reducing the
SSE RSS Shot oC Re ole OA a SR 97-101
Published by the Botanic Gardens
Parks and Recreation Department
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FY CHANGE
THE GARDENS’ BULLETIN
SINGAPORE
VOL. 39 (Part 1) Ist June 1986
CONTENTS
PAGES
WILDE, W.J.J.O. DE:
A New Account of the Genus Horsfieldia (Myristicaceae), Pt 4 ........ 0... ceeccceccceeccee eee 1-65
KENG, Hsuan:
Annotated List of Seed Plants of Singapore (X) ............cccccceeececueecceueeseuseceereceesuees 67-95
FOONG, T.W. and C.N. YANG:
The Evaluation of the Effectiveness of Water-absorbent Polymers in Reducing the
SS Fit LoS fae ag 0 Rg a 97-101
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Singh, H. (1967). Sclereids in Fagraea. Gard. Bull. Sing. 22, 193-212.
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Medinilla alternifolia B\., Mus. Bot. Lugd.-Bat. I:2 (1849) 19.
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A New Account of the Genus Horsfieldia (Myristicaceae), Pt 4*
W.J.J.O. de WILDE
Rijksherbarium, Leiden, The Netherlands
EFFECTIVE PUBLICATION DATE: 26 JULY 1986
Contents
page
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71. Horsfieldia hirtiflora de Wilde, sp. nov. Fig. AC (71); 27
Ramuli validi, cristati, primum pilis 0.3-0.4 mm longis obtecti. Folia disticha vel sparsa, membranacea,
elliptico-oblonga usque ad oblonga, 17-30 cm longa, nervis supra prominentibus. Perianthia mascula
pubescentia, breviter obovoidea, c. 2.5 mm diam., 4-valvata. Androecium indistincte triquetrum an
non, antheris 7, suberectis, in parte superiore liberis. Pedicellus pubescens, basi articulatus. —
Type: Rahmat si Boeea 9257 (L; iso: MICH, n.v.).
Tree 10 m. Twigs subterete, in the apical portion + angular and ridged, lower
down distinctly lined or ridged, 4-6(-15) mm diam., dark brown, glabrescent, hairs
0.3-0.4 mm; bark lower down finely to coarsely striate, lenticels not very conspi-
cuous, older bark not flaking. Leaves in 2 or sometimes in 3 rows, membranous,
elliptic-oblong to oblong, broadest above the middle, 17-30 x 6-11 cm, base +
long-cuneate, top rather shortly acute-acuminate; upper surface glabrous, drying
brown-olivaceous, lower surface drying brown, glabrous but midrib + late glabres-
cent, without scattered larger blackish dots; midrib above raised to rather flattish,
glabrescent; nerves 15-20 pairs, above slender, raised, glabrous, the lateral arches
usually + sunken, distinct; tertiary venation forming a lax network indistinct
above; petioles 15-25 x 2.5-3.5 mm, late glabrescent; leaf bud 8-10 x 4-5 mm, with
hairs c. 0.3-0.4 mm. Inflorescences behind the leaves, densely rusty pubescent with
hairs c. 0.3-0.6 mm; in GC: 3 or 4 times ramified, rather many-flowered, 10-12 x 5-6
cm, common peduncle 35-40 mm, the flowers in clusters of 3-5 each; bracts elliptic-
oblong, acute, 5-10 mm long, densely pubescent, caducous; flowers 3- or 4-valved,
perianth pubescent with hairs 0.1-0.3 mm, pedicel pubescent with hairs 0.2-0.3 mm,
at base articulate; 9 inflorescence rather few-flowered, c. 4 cm long. Male perianth
subglobose-obovoid, in transverse section rounded, collapsing slightly on drying,
2.2-2.5 X 2.0-2.5 mm, top broadly rounded, base + rounded; pedicel 1.5-2.5 mm
long; perianth at anthesis cleft from c. 2 to nearly %, valves 0.3-0.4 mm thick.
Androecium globose to obovoid, (1.2-)1.5 mm diam., subcircular to blunt-angular
in cross-section; anthers 6 or 7, acutish, 0.7-0.8 mm long, free for c. 2-way,
somewhat curved towards the centre, column largely hollowed out, the basal part
of the androecium consisting of a tapering androphore 0.5-0.7 mm long. Female
flowers not seen. Fruits 1-3 per infructescence, broadly ellipsoid-obovoid, top
broadly rounded, base narrowly rounded and short-attenuate, 5.0-5.7 x 3.8-4.5
cm, glabrous, drying dark brown, sparsely coarsely tubercled, pericarp 10(-15) mm
thick; stalk 2-3 mm long, pubescent; perianth not persistent.
*Continued from Gdns’ Bull. Sing. 38 (2): 225
Received on 17 July 1984
Fiz. 27 Horsfieldia hirtiflora de Wilde. .
a. icafy twig with immature male inflorescence, note lined twig, dispersed leaves and bracts
resent in inflorescence, x 2; b, submature male flower, lateral view, x l2ne, ditto, opened,
“howing androecium, x 12; d, immature androecium, longitudinal section, schematic, x 12; e,
~ortion of twig with infructescence, fruit mature, x /%.— a-d, from Rahmat si Boeea 9257; e,
trom Kostermans 22048.
New account of Horsfieldia 4 3
Distribution. Sumatra (Tapanuli, E. Coast).
SUMATRA. Tapanuli: Kostermans 22048 — E. Coast: Rahmat si Boeea 9257, 9362.
Ecology. Forest, 100-500 m alt; on sandstone. Flowers June-July, fruits in
December.
NOTES
1. Fieldnotes. Tree 10 m, diam. 12 cm. Bark rough, hard, black. Fruit yellow,
subglobose, 8 cm diam.
2. Surely closely related to H. brachiata with which it has the distinctly ridged
twigs in common, but differing by the coarser habit, larger and hairy flowers, and
large fruits.
3. The leaves towards the apex of the twigs in Kostermans 22048 and Rahmat Si
Boeea 9257 are not distichous but, rather, arranged in 3 rows.
4. Sinclair identified the specimen Rahmat Si Boeea 9257 as H. brachiata var.
sumatrana, Rahmat Si Boeea 9632 as H. brachiata var. brachiata.
72. Horsfieldia brachiata (King) Warb. Fig. 1C(72).
Myristica brachiata King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 311, pl. 144 — Horsfieldia brachiata
(King) Warb., Mon. Myrist. (1897) 325; Gamble, Mat. Fl. Mal. Penins. 5, 23 (1912) 218; Ridley, FI.
Mal. Penins. 3 (1924) 59 — H. subglobosa (Miq.) Warb. var. brachiata (King) Sinclair, Gard. Bull.
Sing. 16 (1958) 431, fig. 51 E. — H. brachiata (King) Warb. var. brachiata: Sinclair, Gard. Bull. Sing.
28 (1975) 9 — Type: Malaya, Scortechini 1649 (CAL. n.v.; iso: K, L), King 4704 (CAL, n.v.; iso: K,
L; BO, PDA, n.v.), 6771 (CAL, n.v.; iso: BM, K; FI, G, n.v.), Griffith 4351 (CAL, n.v.; iso: K, P,
U; A, G, M, S, n.v.).
Tree 10-35 m. Twigs + angular or subterete, or more or less flattened towards
the apex, almost always distinctly lined or ridged in between the petioles, or
sometimes lines evident only in a part of the material, twigs towards the apex
2-7(-18) mm diam., grey-brown to blackish, generally early glabrescent from dark
brown or rusty hairs (0.1-)0.2-0.4 mm long; bark lower down finely to coarsely
striate, in colour not very contrasting with lenticels; bark when older not flaking.
Leaves in two rows, membranous, elliptic-oblong to oblong(-lanceolate), broadest
at or slightly above the middle, 12-26(-30) x 4-9(-11) cm, base cuneate, top
acute-acuminate; upper surface glabrous, drying olivaceous to brown or sometimes
blackish, with the midrib glabrescent, lower surface drying light brown, early
glabrescent but midrib sometimes later glabrescent; not dotted; midrib raised
above, glabrous; nerves 12-20 pairs, above slender, raised, glabrous, the lateral
arches usually not distinct above; tertiary venation forming a lax network usually
not distinct above; petioles 8-13(-20) x 2-3 mm, glabrescent; leaf bud 8-15 x 3-4
mm, with hairs 0.2-0.4 mm long. Inflorescences sparsely to densely pubescent with
dendroid hairs 0.2-0.5 mm, sometimes partly glabrescent; in CO’: 3-4 times ramified,
many-flowered, 7-18(-22) x 5-16(-18) cm, common peduncle 6-18 mm long, the
flowers in loose clusters of 3-6; Q-inflorescences rather many-flowered, 3-8 Xx 2-6
cm; bracts oblong-lanceolate, acutish, 3-5 mm long, pubescent, caduous; perianths
3 (or 4)-valved, glabrous; pedicels in CO pubescent in various degree with hairs
0.1-0.2 mm, in Q glabrescent, at base articulate. Male perianths subglobose or
4 Gard. Bull. Sing. 39(1) (1986)
slightly depressed-globose to broadly obovoid, in transverse section usually slightly
angular, sometimes rounded, 1.0-1.5 x 1.2-1.8 mm, top (broadly) rounded, base
rounded to short-cuneate, glabrous; pedicel (1.0-)1.5-2.5 mm long, slightly taper-
ing or not; perianth at anthesis cleft to 2-74, not or but little collapsing on drying,
valves 0.2-0.4(-0.5) mm thick. Androecium depressed-globose to obovoid in out-
line, (0.5-)0.7-1.0 x 0.8-1.2 mm, + rounded or usually + 3-angular in transverse
section; anthers 6-10 (12-20 thecae), 0.5-0.7 mm long, mutually free for c. /2-way,
usually curved towards the centre, column largely hollowed out, at base continuing
into the 0.2-0.3-mm long androphore, slightly tapering or not. Female perianth
broadly ellipsoid, 2.2-2.5 x 1.8-2.0 mm, glabrous, cleft at anthesis to c. ¥%3, valves
c. 0.3-0.4 mm thick, pedicels 1-1.5 mm long, glabrous, ovary ovoid, 1-1.4 x 0.8-1.2
mm, glabrous, stigma 2-lobed, c. 0.2 x 0.4 mm. Fruits 4-12(-20) per infructescence,
broadly ellipsoid, top narrowly rounded, base (broadly) rounded, 2.0-2.8(-4.0, see
notes) X 1.8-2.2(-3.0) cm, glabrous, drying brown to dark brown, not warted nor
lenticellate, pericarp 1.5-4(-7) mm thick; stalk 1.5-3 mm long; perianth not per-
sisting.
Distribution. Peninsular Thailand, Malaya (Kedah, Kelantan, Perak, Treng-
ganu, Pahang, Malacca, Johore), Sumatra, Borneo (Sarawak, incl. one deviating
coll., see notes; Sabah; E. & NE. Kalimantan); not found in Singapore and most of
Kalimantan.
THAILAND. Peninsular: (Larsen c.s.) Fl. of Thailand 31184, (Phusomsaeng 424) 36470, (Sang-
khachand 1132) 36570, 52357; A.F.G. Kerr 7426, 7690, 12426, 18185; Put 1184; Shimizu et al. (Kyoto)
8181.
MALAYA. FRI 0417, 4941, 5606, 10387, 11424, 11872, 13310, 13743, 15307, 15807, 17562: Griffith
(d.d. 1845; Kew Distr. 4351); Kep 94517, 94688, 98501, 100129, 104313, 104996; (Kunstler in) King 4704,
6771; KL (of Malaya Univ.) 3095; Ridley 4459; Shah MS. 1541, (& Noor) 1751; Scortechini 1649; SFN
23869, 25991, 28707, 28963, 29466, 31617, 33753, 34569, 34708, 36791, 40772.
SUMATRA. E. Coast: Bartlett 7110; Rahmat si Toroes 4124, 4657; Yates 1601 — Djambi: Roos &
Franken 1653 — Palembang: Lambach 1341.
BORNEO. Sarawak: S 34908 (large fr., see notes), 39/62 — Sabah: Amdjah 1088; San 21509, 48435,
69290, 74417, 74544 NE. & E. Kalimantan (incl. Tarakan, E. Kutei): Kostermans 4988, 5078, 9018,
9171; Meyer 2259, 2380, 2385.
Ecology. Primary and secondary lowland rain forest; often in flatland forest,
marshy forest, riverside forest, peaty forest, forest on alluvial plains, poor forest on
soil with stagnant water, but also on hill sides; found on alluvial soils, brown soil,
sandy soil (in Tristania forest, Sabah), sandstone, peaty soils, loam soil with lime;
often near streams; 0-400 m alt. Flowers and fruits throughout the year.
Vernacular names. Pérédah boeroeng (Palembang), Kajoe darodong (Tapa-
noeli).
NOTES
1. Fieldnotes. Usually a slender tree, with straight bole once recorded as with
buttresses to 50 cm high. Bark + smooth, pale brown to dark brown, generally
shallowly vertically fissured c. 1 cm apart, sometimes recorded as + laminated or
scaly, or cracked. Living bark 8-10 mm thick, pinkish to reddish-brown, exuding
reddish sap. Wood whitish to pale brown; no heartwood. Twigs with raised lines.
Flowers greenish-yellow to dark yellow, scented. Fruits yellow-green, yellow or
yellow-orange.
New account of Horsfieldia 4 5
2. The present species is largely the same as H. subglobosa var. brachiata or H.
brachiata var. brachiata as delimited by Sinclair in 1958 and 1975 respectively. I also
agree that it is very close to H. polyspherula; in fact, it stands in several respects
more or less between the vars. polyspherula and sumatrana of H. polyspherula. H.
brachiata is in most cases easily recognized by its weak to strong raised lines on the
twigs on both sides from petiole to petiole. Its fruits are rather uniform in shape and
size, 20-28 mm long, and thus + intermediate between H. polyspherula var.
polyspherula and var. sumatrana (see there). H. brachiata has the leaves
rather like those of H. polyspherula var. sumatrana, viz. generally membranous,
and drying to pale, dull olivacous above, and pale cinnamon below. Its flowers are
rather uniform, mature male buds 1.2-1.8 mm diam., with a usually + triquetrous
androecium (but see note 3) consisting of 6-10 stamens, and they do not differ from
those of H. polyspherula s.1. Sterile and flowering specimens, in which the apical
and lower twig portions are not sufficiently represented may be difficult to place.
Note that H. brachiata generally has stouter inflorescences in contrast with H.
polyspherula. The former occurs quite commonly in evergreen forests in Peninsular
Thailand, where H. polyspherula is not found yet.
3. Variation. The number of anthers in the androecium is usually 6-8, only in the
material from Peninsular Thailand did I find 9 or 10.
The shape of the mature male perianths in bud is generally subglobose or
short-obovoid, with the transverse section faintly 3-angular, corresponding with the
generally angular shape of the androecium. Only in a few specimens from Malaya
(FRI 0417, Kep 94517, 104313) are the mature male buds rather depressed-globose
and circular in section; also the androecium in these specimens is + rounded in
section, but otherwise there is no reason to exclude them from the present species.
The specimen SFN 40772, from Trengganu, Malaya, is a relatively glabrous
plant. The twigs are almost glabrous, including the apex, and the tomentum of the
leaf bud is composed of hairs only c. 0.1 mm long.
4. Some deviating and doubtful specimens. FRI 17562, from Perak, Malaya, has
twig apices rather sharply 2-angled, and hence would belong to H. brachiata. The
older wood sample, however, is not lined, the fruit is also relatively large, c. 30 x
26 mm, and the colour of the dry leaves rather blackish. Probably this specimen
belongs to H. polyspherula var. sumatrana.
The Malayan specimens King’s Coll. 6771 (syntype) and KEP 100129 are also
somewhat doubtful because of the rather indistinct lines on the twigs.
S 34908 from Sarawak (Kapit, 5th Div.) is a stout specimen and in bad condition;
it has 2 flowers and at L is a single fruit measuring c. 40 x 30 mm; the pericarp is +
woody, c. 5-7 mm thick. H. brachiata is not common in Sarawak and this large-
fruited specimen probably represents a separate taxon. It was collected in a
kerangas-mossy forest at c. 800 m, the highest altitude ever collected of the species.
73. Horsfieldia pachyrachis de Wilde, sp. nov. Fig. 1C(73)
Ramuli validi, primum pilis c. 0.2 mm longis obtecti, deinde glabrescentes. Folia chartacea, 16-26 x
6.5-9 cm, nervis supra prominentibus. Inflorescentiae masculae validae, costa 5-8 mm diam. Perian-
6 Gard. Bull. Sing. 39(1) (1986)
thia mascula subglobosa, 1.5-2 mm diam., 3-valvata. Androecium triquetrum, antheris 5-7, erectis.
Pedicellus basi articulatus. — Type: b.b. 28/28 (L; iso: K; BO, SING, n.v.).
Tree. Twigs terete, towards the apex 5-7 mm diam., blackish-brown, glabres-
cent, tomentum rusty to grey-brown with hairs c. 0.2 mm, bark of older twigs not
seen; lenticels conspicuous. Leaves in 2 rows, thinly chartaceous, obovate-oblong
to oblong, broadest at or somewhat above the middle, 16-26 x 6.5-9 cm, base
(long-)attenuate, top broadly acutish; upper surface drying olivaceous brown to
blackish brown, glabrous, lower surface drying dark brown, without larger blackish
dots, early glabrescent but on the midrib towards the base tomentum vestigial with
hairs 0.2-0.3 mm; midrib rather slender above, near the transition of the petiole
2.5-3 mm wide, raised; nerves 14-17 pairs, raised above, the marginal arches not
very distinct; tertiary venation forming a lax network, flat or + sunken, indistinct;
petioles 7-11 x 3.5-4.5 mm, glabrescent; leaf bud c. 15 X 4 mm, with hairs c. 0.2
mm. Inflorescences apparently behind the leaves (see notes), densely to sparsely
pubescent with dendroid hairs 0.2-0.3 mm, glabrescent, in C’: very stout and rather
compact, 4 or 5 times ramified, many-flowered, c. 14 x 10 cm (not fully expanded),
the main axis stout, towards the base 5-8 mm diam., length of common peduncle
not known (see notes); flowers in clusters of 4-10, perianth 3-valved, glabrous,
pedicel thinly pubescent towards base with hairs 0.1-0.2 mm long, at base distinctly
articulate; bracts caducous, those of uppermost ramifications + elliptic, subacute,
densely pubescent, 3-5 x 2-4 mm. Male perianth (possibly somewhat submature)
broadly globose to broadly obovoid, apically slightly depressed, c. 1.5 X 2.0 mm,
top broadly rounded, base + narrowly rounded, glabrous; pedicel 1.5-2.0 mm long;
perianth at anthesis cleft to nearly ¥2-way, hard-fleshy, not collapsing on drying,
valves 0.2-0.3 mm thick at sutures, perianth towards the base 0.5-0.7 mm thick.
Androecium subglobose-obovoid, c. 0.6 X 0.6 mm, rather sharp-triangular in
transverse section; anthers 5-7 (i.e. 10-14 thecae), slightly curved, almost entirely
connate, concealing an apical cavity in the central column reaching c. “4-'2-way;
androphore slightly tapering towards the base, 0.1-0.2 mm long. Female flowers
and fruits not seen.
Distribution. Borneo, W Kalimantan; only known from b.b. 28128, Melawi,
Bukit Kelawai, 80 m alt., CO fl., 20 May 1939; no further ecological or fieldnotes
given.
NOTES
1. The present species is described as new because it does not fit into any of the
species which it resembles in general habit, i.e., the leafy twig and flowers. The
separate male inflorescence is extremely stout, with the main axis (rachis) c. 5-8
mm thick and with shortish and stout lateral axes, and by this character it keys out
easily. Most of the numerous flowers are submature, but the largest ones apparent-
ly are nearly full-grown. These highly resemble superficially those of the group of
species with H. polyspherula (especially those of var. sumatrana) or H. laticostata,
also in the distinctly triquetrous androecium. However, in the H. polyspherula-
group of species the (sub)erect anthers are mutually largely free, at least for c.
'2-way, whereas in our present species the anthers are (almost) completely con-
nate. Also in general habit and the colour of the leafy twig, it agrees little with
the H. polyspherula group. The connate anthers (and the general appearance of
the leaves) point to the group of species keyed out around H. fragillima, and the
specimen on which the present new species is based would then key out beside H.
New account of Horsfieldia 4 7
borneensis (because of the distinctly articulated pedicels), a species which does not
appear related at all.
2. The only collection on which the present species is based, b.b. 28128, consists
in L of a leafy twig and, in a separately attached envelope, a part of an infloresc-
ence, apparently broken off from the specimen in Kew. The K-specimen only
consists of a stout, slightly submature inflorescence, cut-off close to its base, and
there are no leaves. I have not seen the BO and SING duplicates. The collection
was identified by Sinclair (1975, p. 12) as H. brachiata var. laticostata (my present
H. laticostata), but Sinclair remarks: ‘somewhat approaching robust specimens of
var. sumatrana but probably best here’.
Unfortunately West Kalimantan is still strongly undercollected, as is also appa-
rent from the specimen Hallier 624 from the same area; this latter seems to
represent a new species as well, as is discussed in note 3 under H. valida.
The collection b.b. 28340, sterile, from the same locality as the type (Melawi, B.
Melaban Kenjit, 470 m), resembles the type and may be conspecific; this was
determined by Sinclair as H. fragillima.
74. Horsfieldia ridleyana (King) Warb. Fig. 1C(74)
Myristica ridleyana King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 311, pl. 145 — Horsfieldia ridleyana
(King) Warb., Mon. Myrist. (1897) 331; Gamble, Mat. Fl. Mal. Pen. 5, 23 (1912) 221; Ridley, FI.
Mal. Pen. 3 (1924) 60; Burk., Dict. 1 (1935) 1199; Sinclair, Gard. Bull. Sing. 16 (1958) 432, fig. 52; 28
(1975) 108 — Type: Malaya, Cantley 1768 (K, syntype); King’s Coll. 10917 (CAL, n.v. iso: BM, K, L;
syntype); Scortechini (12) (CO fl., lectotype, annotated by King; BM, K, iso:L), 862 (Q fls. syntype:
CAL, n.v.; iso: K).
Horsfieldia karengasicola Sinclair, in sched. (Borneo material).
Tree 5-25 m. Twigs terete or somewhat angular, towards the apex 1.5-3.5(-5) mm
diam., dark brown, generally early-glabrescent, tomentum rusty, with hairs c.
0.2-0.4 mm long, bark lower down finely to coarsely striate, lenticles either small
and distinct, or absent; older bark not flaking. Leaves in 2 rows, thinly chartaceous
to coriaceous, elliptic-oblong to lanceolate, broadest at about the middle, 5-15(-16)
x 2-4.5 cm, base attenuate, top acute to acute-acuminate; upper surface glabrous,
drying olivaceous to brown, lower surface glabrous, drying light brown to reddish-
brown, sometimes rather contrasting with upper surface, without larger blackish
dots; midrib above flat, sunken or slightly raised, and often inconspicuous, glab-
rous; nerves 7-15 pairs, above sunken to flattish, inconspicuous or often hardly
visible; tertiary venation forming a lax network, not or hardly visible; petioles 7-15
x 1.5-2.5 mm, early glabrescent; leaf bud slender, 6-14 x 1-2 mm, densely
pubescent with hairs 0.2-0.5 mm long. Inflorescences rather sparsely pubescent
with hairs 0.1-0.3 mm, sometimes subglabrous or glabrescent, in O: c. 3 (or 4)
times ramified, rather many-flowered, 2-6 x 2-4 cm, common peduncle (2-) 5-15
mm long, the flowers in loose clusters of 3-6 each; Q-inflorescences fewer-
flowered, c. 1.5-4 cm long; bracts not seen, caducous; perianths 3-or 4-(rarely 2-)
valved, glabrous, pedicels slender, glabrous, at base articulated. Male perianth
subglobose, or + shortly ellipsoid, 1.0-1.2 x 0.8-1.2(-1.3) mm, top rounded, base
rounded or short-attenuate, glabrous; pedicel 1.0-2.0 mm long, slender; perianth at
anthesis cleft to c. ¥3- nearly 2, not or slightly collapsing on drying, valves 0.2-0.4
mm thick. Androecium (incl. androphore) broadly obovoid, 0.6-0.8 x 0.4-0.7 mm,
8 Gard. Bull. Sing. 39(1) (1986)
in transverse section generally sharply 3- or 4-angular (rarely + ellipsoid in 2-valved
flowers); anthers 4-6 (i.e., 8-12 thecae), mutually almost entirely free (Malaya,
Borneo), or free only nearly for upper half (part of the material from Malaya),
sub-erect, acutish, c. 0.3-0.4 mm long; androphore relatively long, + tapering, up
to nearly the length of anthers, 0.2-0.4 mm long. Female perianth ellipsoid, c.
1.5-1.8 xX 1.5 mm, glabrous, cleft at anthesis to 4- nearly 2-way, valves 0.2-0.4(-
0.5) mm thick; pedicels 1.0-2.0 mm long; ovary ellipsoid, 1.2-1.3 x 0.6-0.7 mm, +
grooved at one side, glabrous, stigma shallowly 2-lipped, c. 0.1-0.2 * 0.6 mm.
Fruits 1-6 per infructescene, ellipsoid, top and base rounded, 1.5-2.0 x 1.0-1.4 cm,
glabrous, drying brown, without lenticel-like tubercles, pericarp c. 1.5 mm thick;
stalk 2-5 mm long; perianth not persisting.
Distribution. Malaya (Perak, Kelantan, Trengganu, Pahang, Selangor, Malac-
ca), Borneo (Sarawak, Brunei, Sabah); apparently not in Sumatra (see notes).
MALAYA. FRI 7404, 8334, 14321, 14729, 20023, 20626; Kep. 94376, 98171, 99117; King’s Coll.
10917; Ridley 16177; Scortechini s.n. (12), 862.
BORNEO. Sarawak: Purseglove P. 5607; S 15925, 20925, 21540, 32200, 37632, 37681, 38438, 38579;
Sinclair & Kadim 10406, 10406A.
BRUNEI: (Ashton) BRUN 3277; Sinclair & Kadim 10438 — Sabah (Beaufort Dist.): San. 31427,
49267.
Ecology. Usually in forest on poor soils: heath forest (with Dacrydium beccarii),
kerangas forest, ridge-forest, quartzite conglomerate-ridges, exposed ridges; sand-
stone with very shallow soil, with Gymnostoma, Tristania, Cotylelobium; sandstone
ridges with Dipteris; sandstone summits with Dacrydium, Gymnostoma-forest;
0-1100 m alt. Flowers throughout the year, but in Borneo most collections June to
October; fruits throughout the year.
NOTES
1. Fieldnotes. Slender or crooked trees, once recorded on a hill side as having
many buttresses. Bark dark brown to red brown, shallowly (rectangular) fissured,
shallowly cracked, flaky, or shallowly dippled and fissured. Slash bark reddish,
fibrous, laminated, once recorded as notably dense; sapwood pale, whitish,
creamy pink. Flowers yellow; androecium (stamens) pink; ovary pale green. Fruits
glossy, green turning yellow-green.
2. Specimens Kep 99117 and FRI 20023, both from Pahang, Malaya, have
proportionally many 2-valved flowers in the inflorescences. Such flowers slightly
deviate by their shorter androecium, with shorter androphore, and the androecium
not conspicuously triquetrous but rather subellipsoid in transverse section.
3. The material from Malaya generally has less coriaceous leaves and thicker
tomentum on the leaf-bud (c. 0.4 mm) as compared with most of the material from
Sarawak, Brunei, and Sabah. Specimens from Borneo generally are from heath or
kerangas forest, those from Malaya from ridges and hill slopes.
4. An excluded specimen. I have to exclude the specimen Sinclair & Kadim
10453 from Brunei, which Sinclair probably used for the description of the fruits,
c. 3-4 xX 2-2.8 cm, in his publication of 1975, p. 109. In my opinion H.
ridleyana has essentially smaller fruits, c. 15-20 mm long. The specimen Sinclair &
New account of Horsfieldia 4 9
Kadim 10453 differs furthermore by its much thicker pericarp (c. 8 mm), thinner
leaves, the slightly raised lateral nerves on the upper leaf surface, and rather
differing aspect of the bark of the older twigs; I cannot match it with any species
known to me, and it is described here as a new species, H. disticha.
5. The only specimen seen by me from Sumatra, which may represent H.
ridleyana, is b.b. 6479, a sterile specimen from W. Coast at 1000 m. This differs
from the material from Malaya (which it resembles most) by the coarser tomentum
on leaf bud and twig apex, and the midrib rather raised above. Possibly H.
ridleyana does not o¢cur in Sumatra, and the specimen b.b. 6479 may belong
instead to a new species H. triandra which is based on Forbes 2465.
6. Vegetatively H. ridleyana may be confused with H. penangiana. In Borneo H.
ridleyana may resemble H. oligocarpa, but the latter has distinctly raised nerves on
the upper leaf surface.
75. Horsfieldia obtusa de Wilde, sp. nov. Fig. 1C(75)
Gemma pilis 0.2-0.4 mm longis obtectum. Folia coriacea, oblonga, 8-10 cm longa, apice rotundata,
nervis c. 10 paribus, supra planis. Perianthium masculum subglobosum, c. 1.5 mm diam., 3-valvatum.
Androecium triquetrum, antheris 9 vel 10, suberectis, in parte dimidia superiore liberis. Pedicellus
basi articulatus. — Type: Sarawak, (Native Collector) Bureau of Science 821 (L).
Tree. Twigs terete or faintly angular, towards the apex 2.5-4(-6) mm diam.,
grey-brown, rather late glabrescent, tomentum dark rusty, with hairs 0.2-0.4 mm
long, bark lower down rather coarsely striate, when older finely, longitudinally
cracking; lenticels present but small and inconspicuous. Leaves in 2 rows, thinly
coriaceous, oblong, broadest at about the middle, 8-10 x 3-3.5 cm, base attenuate,
top rounded; upper surface glabrous (except midrib), drying dark olivaceous, lower
surface glabrous (early glabrescent), drying somewhat purplish brown, provided
with many paler usually pale yellowish enlarged hair-scars but not dotted; midrib
above moderately raised, late glabrescent, nerves c. 10 pairs, flat or slightly raised
above, well-visible, the submarginal arches rather regularly shaped and well-
visible; tertiary venation forming a lax network, not or hardly visible; petioles c. 10
x 2 mm, rather late glabrescent; leaf bud c. 12 X 3 mm, densely dark rusty
pubescent, hairs 0.2-0.4 mm. Inflorescences situated behind the leaves, densely
pubescent, rusty hairs 0.2-0.4(-0.5) mm long, in CG: c. 3 times ramified, many-
flowered, 5-9 x 3-5 cm, common peduncle 3-10 mm long, the flowers in clusters of
3-6 each; 9 inflorescences not seen; bracts not seen, caducous. Flowers 3(or
4)-valved, perianth glabrous, pedicel glabrous, terete, at base articulate. Male
perianth subglobose, 1.3-1.5 x 1.5-1.6 mm, top and base broadly rounded; pedicel
1-1.5 mm long, slender; perianth at anthesis cleft to nearly /2-way deep, + woody-
brittle, not collapsing on drying, valves at the top c. 0.2 mm, at base c. 0.4 mm
thick. Androecium (incl. androphore) broadly obovoid, c. 0.8 x 1.0 mm, in trans-
verse section 3 (or 4)-quetrous; anthers 9 or 10 (thecae 18 or 20), c. 0.5 mm long,
suberect, free in the upper half, acutish; androphore + tapering to below, c. 0.3
mm long; apical cavity rather distinct, depth to about the base of the anthers.
Female flowers and fruits not seen.
Distribution. Only known from the type from Sarawak, precise locality not
indicated.
Ecology. Not known.
10 Gard. Bull. Sing. 39(1) (1986)
NOTES
1. Obviously a member of the group of species with H. polyspherula, because of
the articulated pedicel and the angular androecium with the anthers distally free. It
is distinguished by its generally flat nerves and the rather large male flowers with 9
or 10 anthers. Because of the smallish coriaceous leaves with rounded tip it is
reminiscent of H. montana, but that species has quite different male flowers.
Similar large, pale yellowish hair-scars on the leaves can be found 1 in H. xanthina, a
species quite different in various ways.
2. Sinclair identified the type specimen in 1959 as H. montana, a species which in
1975 he reduced to H. glabra, but the present type-collection was not included
among the cited specimens.
76. Horsfieldia disticha de Wilde, sp. nov.
Ramuli non-angulati, primum pilis 0.2-0.3 mm longis obtecti deinde glabrescentes, deorsum cortice
longitudinaliter fisso. Folia disticha, chartacea, oblongo-lanceolata, 8-13.5 cm longa, longe acumina-
ta, nervis supra prominentibus. Fructus ovoideo-ellipsoidei, c. 3 cm longi, glabri, pericarpio (in sicco)
8-10 mm crasso. — Type: Brunei, Sinclair (& Kadim) 10453 (L; iso: BM, K; A, B, E, NY, SAR,
SING, n.v.).
Tree 20 m. Twigs terete to faintly angular, not ridged, towards the top 2-3(-4)
mm diam., in fruit-bearing portion 11-13 mm diam., dark grey-brown or dull
reddish-brown, very early glabrescent, tomentum dull rusty, of hairs 0.2-0.3 mm,
bark lower down coarsely longitudinally cracking, lenticels rather sparse but dis-
tinct. Leaves in 2 rows, chartaceous, oblong-lanceolate, broadest at about the
middle or + parallel-sided, 8-13.5 x 2.5-3.5 cm, base + rounded to short-
attenuate, tip long-acute-acuminate (acumen c. 15 mm); upper surface drying
olivaceous-brown, glabrous, lower surface pale chocolate, glabrous, without larger
scattered dots; midrib raised above, glabrous; nerves 9-13 pairs, raised above, the
marginal arches indistinct; tertiary venation faint or invisible on both surfaces;
petioles relatively long and slender, glabrous, 10-15 x 1-1.5 mm; leaf bud slender,
c. 10 x 2 mm, densely dull rusty pubescent with hairs 0.2-0.3 mm long. Male and
female flowers not seen but perianth apparently 3-valved as judged from the
perianth-scars on the fruits. Fruits 3-6 in infructescences measuring 3-6 X 2-4 cm,
borne on the older wood behind the leaves, glabrous (glabrescent), the fruits
ovoid-ellipsoid, top narrowly rounded, base broadly rounded, 2.8-3.2 x 2.1-2.5
cm, glabrous, drying dark brown, not tubercled nor lenticellate, pericarp hard-
woody when dry, 8-10 mm thick; stalk 10-15 mm long; perianth not persistent.
Distribution. Borneo: Brunei, known only from the type.
Ecology. Lowland forest, at side of new road, Andulau Forest Reserve (West);
fruits in August.
NOTES
1. Fieldnotes. Bark with longitudinal, shallow furrows as in H. wallichii. Twigs
slender, leaves distichous, dark green above, paler beneath, dull on both surfaces.
Fruits unripe, pear-shaped, large.
New account of Horsfieldia 4 it
2. According to the general habit of the specimens, and especially the leaves
with the nerves raised above, this species obviously belongs to the group of H.
polyspherula. It was, together with some other specimens, provisionally identified
as H. disticha by Sinclair. In his posthumous publication these specimens were
listed as H. ridleyana, and | agree, except for the collection Sinclair & Kadim 10453
on which the present new species is based. H. disticha differs from H. ridleyana in
its general habit, the raised nerves, the nature of the bark on the twigs, and the
much larger fruits with a conspicuously thick pericarp. In fact, the above specimens
were used by Sinclair to describe the fruits of H. ridleyana, but it seems to me that
the fruits of that species are different, being smaller and with a much thinner
pericarp.
77. Horsfieldia tenuifolia (Sinclair) de Wilde, stat. nov. Fig. 1C(77); 28
Horsfieldia polyspherula var. tenuifolia Sinclair, Gard. Bull. Sing. 28 (1975) 105. — Type: Haji Bujang
S 13686 (SING, n.v.; iso: K, L; S, SAR, n.v.).
Tree 5-15 m. Twigs terete, towards the apex 1-3(-4) mm diam., early glabrescent,
tomentum greyish-rusty, hairs c. (0.1-)0.2 mm, bark of older twigs striate, lower
down neither cracking nor flaking; lenticels inconspicuous or absent. Leaves in 2
rows, membranous to thinly chartaceous, elliptic to oblong-lanceolate, broadest at
about the middle, 6.5-16.5 x 3-6.5 cm, base attenuate, top acute-acuminate; upper
surface drying greyish olivaceous to dark greyish brown, glabrous, lower surface
dull greyish brown, glabrous, without larger blackish brown dots; midrib raised
above, glabrous; nerves 5-11 pairs, raised above, marginal arches regular but rather
faint; tertiary venation forming a lax network, indistinct or hardly visible; petioles
8-16 X 1.5-2 mm, glabrous; leaf bud 5-8 x 1-1.5 mm, densely pubescent with hairs
c. 0.2(-0.3) mm. Inflorescences moderately to sparsely pubescent with stellate hairs
0.1-0.2 mm, in CO: rather many-flowered, c. 3 times ramified, 3-5 x 2-4 cm,
common peduncle 2-10 mm; in Q: rather slender, few-flowered, c. 2-3.5 x 1 cm;
bracts elliptic to oblong, 2-3 mm, pubescent, caducous. Flowers in loose clusters of
3-8 each, perianth 3-(or 4-)valved, glabrous, pedicels glabrescent or with few
scattered hairs up to 0.2 mm in the lower half, at base indistinctly or not articulate.
Male perianth globose to broadly obovoid, 0.8-1.3 x 1.0-1.5 mm, top broadly
rounded, base rounded to subattenuate, glabrous; pedicels 1-2 mm long, some-
what tapering; perianth at anthesis cleft to depth of c. 4 to nearly ¥2-way, valves
0.2-0.3 mm thick, at base up to 0.4 mm thick. Androecium + obovoid to ellipsoid,
0.5-0.7 x 0.5-0.8 mm, in transverse section triquetrous; anthers 4-6, sub-erect,
0.3-0.4 mm long, mutually free for at least /2-way, central column largely hollow,
towards the base continued into the somewhat tapering androphore 0.2-0.3 mm
long. Female perianth ellipsoid, c. 2.0 X 1.5 mm, cleft at anthesis to c. 43 or slightly
over, valves c. 0.3 mm thick, pedicel c. 1.5 mm long, at base + articulate or
inarticulate, glabrescent, hairs 0.2-0.3 mm; ovary ellipsoid, c. 1.3 x 0.8-0.9 mm,
glabrous, stigma 2-lobed, c. 0.2 mm high. Fruits 1-3 per infructescence, ellipsoid,
top rounded, base rounded or shortly tapered, 1.7-2.0 x 1.4-1.5 cm, glabrous,
drying dark brown, not lenticel-like tubercled, pericarp 1-1.5 mm thick; stalk 3-5
mm long; perianth not persistent.
Distribution. Borneo: Sarawak (1st. Div.), Sabah (Beaufort Hill, Jesselton).
BORNEO. Sarawak: S 12774, 13686, 24914, 24945, 26599, 34528; Sinclair & Kadim 10179 — Sabah:
Jesselton, Kep. 71665; Beaufort Hill, San. 44531.
12
Horsfieldia tenuifolia (Sinclair) de Wilde.
a. habit of twig with leaves and male inflorescences, x 2; b, ditto with female inflorescence,
x 1/2; c, mature male flower, lateral view, x 12; d, ditto, opened, showing androecium, X 12;
e, mature female flower, x 12; f; ditto, opened, showing glabrous ovary and minute 2-lipped
stigma, X 12; g, portion of twig with infructescence, fruits mature, X /2— a, c, d, from §
24945, b, e, f, from S 34528; g, from S 24914.
New account of Horsfieldia 4 13
Ecology. Lowland dipterocarp forest, 0-300 m alt.; soil a yellow clay, yellow
sandy clay, yellow loam, brownish soil; on ridges and slopes. Flowers and fruiits
May to September.
NOTES
1. Fieldnotes. Slender tree, once recorded as buttressed. Bark dark brown to
greenish brown, narrowly fissured, not flaking; inner bark dark red, with red latex;
sapwood whitish. Flowers greenish yellow. Fruits yellowish.
2. Sinclair regarded the present species as a variety of H. polyspherula, and
some specimens were included by him in the type-variety, H. polyspherula var.
polyspherula.
Our present species is marked by quite a different habit; it has the twigs glabrous
to the top, the leaf bud is small and slender, covered with hairs only c. 0.2 mm long;
the leaves are membranous to thinly coriaceous, not brittle, drying to a greyish
tinge and the colour of the upper and the lower surfaces not markedly different; the
petioles are relatively long and slender; the inflorescences and flowers are compa-
ratively small. According to the architecture of the male flowers the species is
undoubtedly closely related to H. polyspherula. The present species is an under-
storey tree of the lowland dipterocarp forest on richer soils.
3. § 34528, with female flowers, has the twigs rather pale and contrasting with the
darker colour of the dry leaves and petioles; by this feature, it may vegetatively key
out to the group of species, incl. H. oligocarpa, characterized by a pale colour of the
dry twigs.
A much related species is H. macilenta, also with thin membranous leaves, but it
has much more pubescent twigs and inflorescences
78. Horsfieldia macilenta de Wilde, sp. nov Fig. 1C(78)
Ramuli tenues. Gemma pubescens pilis c. 0.5 mm longis. Folia membranacea, oblonga usque ad
oblongo-lanceolata, 10-18 cm longa, basi attenuata, nervis lateralibus paribus 10-15, supra prominen-
tibus. Perianthium masculum subglobosum, c. 1 mm diam., 3-valvatum. Androecium triangulare ad
sectionem transversam, antheris 5-7, suberectis, in parte dimidia superiore liberis. Pedicellus basi
articulatus. — Type: Sabah, A. Gibot SAN. 37103 (L).
Tree 4-17 m. Twigs terete, towards the apex 1-3(-12) mm diam., rather late
glabrescent, tomentum pale rusty, composed of stellate-dendroid hairs (0.2-)0.5
mm long, bark of older twigs rather finely striate, not cracking; lenticels small and
usually inconspicuous. Leaves in 2 rows, membranous, oblong to oblong-
lanceolate, broadest at about the middle, 10-18(-27) x 3-6.5 cm, base (rounded to
short-)attenuate, tip acute-acuminate; upper surface drying dull olivaceous, glab-
rous, lower surface rather bright light brown, glabrous except midrib and the very
base; without larger, dark brown dots; midrib raised above, on lower surface late
glabrescent; nerves 10-15(-18) pairs, raised, the marginal arches fairly regular,
distinct or not; tertiary venation forming a lax network, + indistinct; petioles 10-15
x 1.5-2.0 mm, rather late glabrescent; leaf bud 6-8 x 1.5 mm, densely pubescent
with hairs (0.2-)0.5 mm. Inflorescences rather sparsely woolly-pubescent with hairs
(0.2-)0.5 mm, in ©’: many-flowered, 2 or 3(-4) times ramified, 4-9(-10) x 3-5(-7)
14 Gard. Bull. Sing. 39(1) (1986)
cm, common peduncle 3-12 mm long; 9 inflorescences similar to the males (see
notes);bracts oblong to lanceolate, 2-5 mm, pubescent, caducous. Flowers in CO in
clusters of 5-8, perianth 3-valved, glabrous or sometimes a few minute hairs less
than 0.1 mm are present towards the base of the pedicel; pedicel terete, glabrous or
sparsely pubescent in the lower portion with hairs 0.1 mm or less, distinctly
articulate at base. Male perianth globose to depressed globose, 0.7-1.2 x 1.0-1.2
mm, top and base (broadly) rounded; pedicels c. (0.8-) 1.0(-1.3) mm long; perianth
at anthesis cleft to nearly ’2-way, valves 0.2-0.3 mm thick. Androecium subglo-
bose, (incl. androphore) 0.4-0.5 x 0.5-0.7 mm, in transverse section triangular;
anthers 5-7, suberect, 0.3-0.4 mm long, at least the upper half mutually free,
column largely hollow and passing into the broad + tapering androphore c. 0.2-0.3
mm long. Female perianth (see notes) subglobose, c. 1.5 mm diam., cleft at
anthesis to c. 12-way, valves c. 0.4-0.5 mm thick, pedicel c. 1.5 mm, subglabrous,
only in the lower part with a few scattered hairs less than 0.1 mm long, base
articulate; ovary globose, glabrous, 1.0-1.2 mm diam., stigma minutely 2-lipped,
(0.1-)0.2 mm long. Fruits (in Sumatra) 2 or 3 together in a short infructescence 2-3
cm long, the fruits glabrous, ellipsoid, 2.2-2.4 x 1.5-1.6 cm, top rounded, base
contracted into a 2-4-mm long narrowed portion, pericarp c. 2 mm thick, drying
brown, not or sparingly tubercled; stalk c. 5 mm long; perianth not persistent.
Distribution. Sumatra, Malaya, Borneo (Sarawak, Sabah).
Cultivated Bot. Garden Bogor (orig. unknown): sub IV.H. 29 (Oct. 1912), in herb. L.
MALAYA (Kelantan, Johore): Whitmore FRI 4415; Kadim & Noor KN. 185.
SUMATRA (Jambi Prov.): Roos & Franken T.F.B. 1999.
BORNEO. Sarawak (4th Div.): Hirano & Hotta 1206; S 39027 — Sabah: B.N.B. For. Dept. 4204; A.
Gibot SAN 37103; SAN 53268.
Ecology. Primary lowland mixed dipterocarp forest, swamp forest; 0-200 m alt.
Flowers from June to November.
NOTES
1. Fieldnotes. Outer bark whitish or green-yellow, inner bark red with latex red,
cambium yellowish. Flowers greenish yellow to yellow.
2. Clearly much related to H. polyspherula especially to the slender specimens of
var. polyspherula, according to the structure of the male flowers. The present
species differs by many details in general habit such as its more slender and tiny
build: the slender twigs, the thinly membranous leaves and the slender and tiny
inflorescences.
3. I am not quite sure whether the only known female flowering specimen (5S.
39027) belongs here. Its leaf nervation is a trifle more projecting, and the marginal
arches are regular and distinct, the pubescence on the leaf bud, twig apex, and
inflorescence is shorter than in the four male specimens presently known, viz. c. 0.2
mm long versus c. 0.5 mm in the male specimens. Remarkable are the female
inflorescences which are as much-branched as the male; all flowers are mature and
purely female. In the related H. polyspherula the female inflorescences are always
noticeably smaller as compared with the males. In the fruiting specimen T.F.B.
New account of Horsfieldia 4 15
(Roos & Franken) 1999, from Sumatra, the infructescence is, as expected, only 2-3
cm long.
4. Specimens of the present species were included by Sinclair (1975) in H.
polyspherula; he did not see others.
79. Horsfieldia laticostata (Sinclair) de Wilde, stat. nov. Fig 1C(79)
Horsfieldia brachiata (King) Warb. var. laticostata Sinclair, Gard. Bull. Sing. 28 (1975) 12 — Type:
Sinclair 10265 (K; iso: L; A, E, SAR, SING, n.v.).
Tree 12-35 m. Twigs terete, towards the top 3-8(-15) mm diam., grey-brown to
blackish, early glabrescent, tomentum grey-brown to rusty, composed of hairs
0.1-0.4 mm, bark lower down coarsely striate, when older almost flaking, lenticels
usually distinct. Leaves in 2 rows, thinly to thickly coriaceous, obovate-oblong to
oblong, broadest at or somewhat above the middle, (15-)20-33 x 6-12 cm, base
attenuate, top (short-)acute-acuminate; upper surface drying dull olivaceous to
greenish brown, glabrous, lower surface drying chocolate brown to rusty, rather
contrasting with the upper surface, without blackish dots or markings, early
glabrescent; midrib above raised, + slender to broad but towards the base flattish
and conspicuously broadened to 3-4(-5) mm wide at the transition to the petiole,
glabrous or sometimes with some remnants of tomentum towards the base; nerves
11-20(-24) pairs, largely raised but towards the blade margin sunken, the marginal
arches usually + distinct, sunken; tertiary venation forming a lax network, flat or
sunken, distinct or not; petioles stout, 6-15 x 5-8 mm, early glabrescent; leaf bud
15-23 X 3-5 mm, densely grey-brown to rusty pubescent with hairs 0.1-0.4 mm long.
Inflorescences rather sparsely pubescent with hairs 0.2-0.4 mm long, in CG: stout, 4
or 5 times ramified, many-flowered, 10-25 x 8-22 cm, main axis towards base 3-4
mm diams., common peduncle 5-35 mm long; Y inflorescences 3-7 cm long, 2 or 3
times ramified, rather few-flowered; bracts elliptic, acute, 2-8 x 1-4 mm, pubescent
with hairs c. 0.3 mm, + glabrescent, caducous; flowers 3-valved, in clusters of up to
10, perianth glabrous, pedicel glabrous except sometimes towards base a few hairs
are present, 0.1-0.2 mm, at base distinctly articulate. Male perianth broadly obo-
void to subglobose, 1.3-1.5 mm diam., top broadly rounded, base rounded or
sometimes + attenuate, glabrous; pedicel (1.0-)1.5(-2.0) mm long, slender;
perianth at anthesis cleft to c. /2-way, not collapsing on drying, valves 0.2-0.3 mm,
towards base of perianth c. 0.5 mm thick. Androecium including androphore
broadly obovoid in outline, c. 0.7 mm diam., sharp-triangular in transverse section;
anthers 6 (thecae 12), suberect, acutish, mutually largely free, c. 0.3(-0.4) mm long;
androphore + tapering, c. 0.3-(0.4) mm long. Female perianth broadly ellipsoid-
globose, c. 2.5 mm diam., glabrous or with a few minute hairs c. 0.1 mm long, cleft
at anthesis to c. ¥%, valves 0.3-0.4 mm thick, pedicel 0.5-1 mm long; ovary broadly
ovoid, c. 1.5 mm long, glabrous, stigma c. 0.1 mm long, shallowly 2(-4)-lobed.
Fruits 1-4 per infructescene, ellipsoid, top and base rounded, 2.8-4.0 x 2.2-2.5 cm,
glabrous, drying brown, with or without warts or lenticels, pericarp c. 4-5 mm
thick; stalk 2-4 mm long; perianth not persisting.
Distribution. Borneo: Sarawak, Sabah, N.E. Kalimantan.
BORNEO. Sarawak: Chew Wee-Lek CWL. 637; Haviland 3074; S 13689, 17252, 18519; Sinclair 10265
— Sabah: B.N.B. For. Dept. 2487; San A 2995, 32602 N.T. 223, 67215, 74585, 80429, 83709 — E. & NE.
Kalimantan: Kostermans 4355, 8777, 9010, 9321.
16 Gard. Bull. Sing. 39(1) (1986)
Ecology. Primary peat swamp forest, forest on sandy acid soils, waterlogged sand
soils, heath forest, karengas, poor forest; 0-400 m alt. Flowers and fruits through-
out the year.
Vernacular name. Piasau piasau (Kedayan lang., Sabah).
NOTES
1. Fieldnotes. Tree to 35 m, without buttresses. Branches predominantly hori-
zontal. Bark hard, fissured or flaky, brown to reddish-brown; inner bark reddish,
laminated, sapwood whitish. Flowers yellow, smelling of Peru balsam. Fruits
yellow, orange or red; once recorded as “‘fruiting abundantly throughout the crown
along the smaller branches’. Seed shining white, spotted.
2. According to Sinclair (p. 12, 13) this is a robust ecological form of H.
brachiata var. sumatrana (in this treatment H. polyspherula var. sumatrana). In-
deed, the present species seems closely related to H. polyspherula, especially to the
varieties sumatrana and maxima, but I prefer it to keep it a separate species as it
can usually be satisfactorily distinguished and as such it does not further complicate
the still very variable H. polyspherula-complex; similarly I have treated H. oligo-
carpa aS a separate species, which is also very close to H. polyspherula, but
separable on various grounds, including a distinct habitat in heath forest.
The present species seems largely confined to forests on poor soils; peaty and
sandy grounds. It is characterised by the stout twigs, large coriaceous leaves with
the midrib very broad and flat at the transition to the petiole, the usually stout,
broad petioles, the fruits of moderate size (c. 30-40 mm), and the large, stout, male
inflorescences. In size and architecture of the male flowers (perianth and
androecium) it is very similar to H. polyspherula.
80. Horsfieldia nervosa de Wilde, sp. nov. Fig. 1C(80)
Gemma pilis c. 0.5 mm longis obtectum. Folia tenuiter coriacea, oblonga, 16-28 cm longa, basi
subrotundatis, nervis later-alibus paribus 16-19, supra prominentibus. Perianthium masculum globo-
sum, c. 1.2 mm diam., 3-valvatum. Androecium triangulare ad sectionem transversam, antheris 5 vel
6, erectis, in parte dimidia superiore liberis. Pedicellus basi articulatus. — Type: Sarawak, Ilias Paie &
P.S. Ashton S 16652 ((L; iso: K; S, NB, A, M, B, n.v.).
Tree 13-16 m. Twigs terete or towards the apex subterete, 3.5-6(-8) mm diam.,
grey-brown, not early glabrescent, tomentum rusty, with hairs 0.3-0.6 mm long,
bark lower down coarsely striate, with the lenticels rather inconspicuous, older
bark slightly flaking or not. Leaves in 2 rows, thinly coriaceous, oblong, broadest +
at the middle, 16-28 x 5.5-9 cm, base short-attenuate to rounded, top acute-
acuminate; upper surface drying olivaceous to greenish, largely glabrous but midrib
and nerves late glabrescent, lower surface chocolate colour, conspicuously con-
trasting in colour with the upper surface, without larger blackish dots, glabrescent
but tomentum of hairs c. 0.5 mm often partly remaining on nerves especially the
midrib; midrib slender above, at transition of petiole c. 1.5 mm wide, distinctly
raised, late-glabrescent; nerves 16-19 pairs, above conspicuously raised, glabres-
cent, the lateral arches flattish,.indistinct; tertiary venation forming a lax network,
indistinct or hardly visible; petioles 15-22 x 3.5-4.5 mm, (late) glabrescent; leaf bud
16-20 x 3.5-4.5 mm, densely rusty pubescent with hairs c. 0.5 mm long. Infloresc-
New account of Horsfieldia 4 17
ences seen only on the older wood behind the leaves, densely pubescent with
woolly-shaggy hairs up to 0.6 mm long, in GC: c. 4 times ramified, many-flowered.
c. 10 X 10cm, common peduncle 10-20 mm long, the flowers in clusters of 2-6 each:
Q inflorescences fewer-flowered, c. 4.0 xX 3.5 cm; bracts pubescent, caducous:
flowers 3-valved, perianth glabrous, pedicel towards base pubescent with hairs
0.1-0.2 mm, articulate at base. Male perianth globose, c. 1.2 mm diam., top and
base + rounded, glabrous; pedicel c. 1.5 mm long: perianth at anthesis cleft to
nearly 2-way, not collapsing on drying, valves c. 0.2 mm thick, at base of perianth
+ fleshy-coriaceous, 0.5-0.6 mm thick. Androecium incl. androphore + broadly
obovoid, c. 0.6-0.7 X 0.6 mm, triquetrous in transverse section: anthers 5 or 6.
(thecae 10 or 12), at least the upper half free, + erect, c. 0.3-0.4 mm long; column
deeply hollowed; androphore + tapering to the base. c. 0.3 mm long. Female
perianth subglobose, c. 2.5 mm diam., subglabrous with some scattered hairs c. 0.1
mm towards base, cleft at anthesis to slightly over 43, valves c. 0.5 mm thick.
pedicels c. 1.5(-2.0) mm long, + pubescent with hairs 0.1-0.2 mm: ovary broadly
ovoid, c. 1.5 mm diam., glabrous, stigma c. 0.3 X0.6 mm, 2-lobed, the lobes very
shallowly 5 or 6-lobulate. Fruits not seen.
Distribution. Borneo (Sarawak, Ist Div.); known from two collections only.
Ecology. Primary forest on yellow podsolic soil; c. 70 m alt. Flowers in
November.
NOTES
1. Fieldnotes. Bark pale ochre and brown-mottled, smooth. Buttresses thin,
small, to c. 35 cm tall. Flowers pale yellow.
2. This species is a close relative of H. polyspherula; its male flowers are
practically identical. The known specimens, S$ 16651 (2), and 16652 (C’). both from
Bt. Gaharu, Serian Dist., are somewhat different especially in the leaves: those of
H. polyspherula are generally smaller, with fewer lateral nerves and generally an
attenuate base. In our present species the nerves on the upper leaf surface are very
strong and markedly raised and distinct, and midrib and nerves on both surfaces
remain for rather a long time covered with the indumentum. Sinclair did not seem
to have seen these two specimens.
81. Horsfieldia polyspherula (Hook. f. emend. King) Sinclair Fig. 1C(8l)
Myristica polyspherula Hook. f., Fl. Brit. Ind. 5 (1886) 108 (p.p., see notes by Sinclair, 0.c., 1958, p.
425; 1975, p. 103); King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 312, pl. 146, emend. — Horsfieldia
polyspherula (Hook. f. emend. King) Sinclair, Gard. Bull. Sing. 16 (1958) 422, fig. 47, pl. XII B; 28
(1975) 101, p.p.. for the type-variety only. — Lectotype (Sinclair, p. 103): Griffith 4354 (K: iso: BM,
P, U; C, CAL, CGE, FI, G, S, n.v.).
For further synonyms see under the varieties.
Tree 4-35(-40)m. Twigs terete or subterete, never distinctly lined nor ridged,
2-5(-13) mm diam., grey-brown to blackish, early to rather late glabrescent, tomen-
tum dark rusty ‘mealy’, composed of dendroid hairs 0.1-0.6 mm long; bark lower
down finely to coarsely striate, lenticels distinct or not, older bark neither cracking
nor flaking. Leaves in 2 rows, thickly membranous to chartaceous, usually brittle,
18 Gard. Bull. Sing. 39(1) (1986)
(elliptic-)ovate-oblong to oblong-lanceolate, broadest usually at the middle, 7-28 x
2.5-9 cm, base subrounded to usually attenuate, top acute-acuminate; upper sur-
face glabrous, drying greenish, olivaceous, or greenish-brown, the midrib glabrous
(early glabrescent), lower surface drying light brown to red or chocolate-brown,
usually much contrasting with the upper surface, without any larger brown or
blackish dots, early glabrescent but with the midrib Often late glabrescent; midrib
slender; nerves 6-15 pairs, above very distinctly raised (except close to the blade
margin), glabrous, the lateral arches usually indistinct above; tertiary venation
forming a lax network usually not or hardly visible above; petioles 6-15 x 1.5-3.0
mm, early to rather late glabrescent; leaf bud 6-17 xX 1.5-3.0 mm, pubescent by
rusty dendroid hairs 0.1-0.6 mm long. Inflorescences rather sparse to densely
woolly-pubescent with hairs c. 0.6 mm long, sometimes glabrescent, in CO’: 3-5 times
ramified, many-flowered, 4-15 x 3-12 cm, common peduncle 3-15 mm long, the
flowers usually in clusters of up to 8 each; 9 inflorescences few- to many-flowered,
generally smaller than the males, up to 8 cm long; bracts oblong to lanceolate, c.
1.5-7 mm long, densely pubescent, caducous; perianths 3-valved, glabrous or in 9
glabrescent, pedicel glabrous or minutely pubescent towards the base which is
articulate. Male perianth globose or broadly obovoid, 1.0-1.8 mm diam., top
(broadly) rounded, base rounded or rarely + tapering into the pedicel, glabrous;
pedicel slender, sometimes tapering, 0.8-1.5(-2.0) mm long; perianth at anthesis
cleft to 2-44, not collapsing on drying, valves 0.2-0.4, at base up to 0.6 mm thick.
Androecium (incl. androphore) + broadly obovoid in outline, 0.5-0.8 x 0.6-1.0
mm, usually sharp-triangular in transverse section; anthers (3 or) 4-7 (thecae 6-14,
free), almost completely free, at least the upper half, + curved or suberect, 0.3-0.5
mm long, apex acutish, at base attached to the short bowl-shaped column which
continues into the relatively long + tapering androphore (0.2-)0.3-0.4 mm long.
Female perianth broadly ellipsoid-obovoid, 2.0-3.0 x 1.8-2.8 mm, glabrous or
sparingly pubescent (hairs c. 0.1 mm), cleft at anthesis to /3-'2, valves towards base
c. 0.5 mm thick, pedicels 1-1.8 mm long, minutely pubescent; ovary ovoid-
ellipsoid, 1.2-1.5 x 1.0-1.5 mm, glabrous, stigma shallowly 2-lobed, c. 0.2-0.3 x 0.5
mm. Fruits 1-6 per infructescence, subglobose to ellipsoid, top rounded, base
rounded or slightly attenuate, 1.9-6.0 x 1.4-5.0 cm, glabrous, drying light to dark
brown, not lenticel-like tuberculate, pericarp 2-15 mm thick; stalk 1-4 mm long;
perianth not persisting.
Distribution. Malaya, Sumatra, Borneo, Philippines (Mindanao, only the var.
polyspherula) 0-1100 m altitude.
NOTES
1. A very variable species. After having segregated some closely related species
such as H. oligocarpa, H. tenuifolia, and H. laticostata (all 3 from Borneo), H.
brachiata (Sumatra to Borneo), and H. majuscula (Malaya), I found that the
remaining specimens from the same island-areas are still heterogenous. Much
variability is found in the habit of the plants, i.e., the thickness of the twigs, size
and texture of leaves, denseness of the indumentum on leaf bud and twig apex, and
especially in fruit size and thickness of the pericarp. On the basis of mainly fruit
characters, three rather heterogenous varieties are presently recognized. There are
slight differences in the size of the male perianth, attributed mainly to the varying
thickness of its valves; the number of anthers is possibly the same in all three
varieties.
New account of Horsfieldia 4 19
2. The anthers are composed of two rather widely separated, almost mutually
free thecae, giving the impression as if there are twice as many anthers as are
present. Thus, Sinclair gave the number of anthers as 9-12 for H. polyspherula
(1958, p. 22) and 8-13 for H. subglobosa (sensu Sinclair, l.c. p. 426, but H.
polyspherula as treated here).
KEY TO THE VARIETIES
la. Fruits (when dry) (3.5-)4.0-6.0 cm long, the pericarp 5-15 mm thick. Leaves 9-20 cm long, nerves
Ee Fiat Ri PROPS OE ROWE og eres optic gn <2 = sqnacgene es ece ee cessanacecege anes sghanas c. var. maxima
Bo Preis an to 3.5 cil lone) mic peticarp 2-S MINER 201220... 2
2a. Fruits (2.5-) 2.8-3.5 cm long. Male perianth 1.2-1.8 mm diam.; anthers 6 or 7. Leaves 13-28 cm long,
OE ee eons Se RE ee eee ere ee b. var. sumatrana
b. Fruits 1.9-2.5(-2.8) cm long. Male perianth 1.0-1.5 mm diam.; anthers (3 or)4-7. Leaves 7-19 cm
Megas, Tee es | pete ere a. tks. adn AE hs ROR tet ee chee a. var. polyspherula
a. var. polyspherula Fig. 1C(81)
Horsfieldia lemanniana auct. non (A. DC.) Warb: Warb., Mon. Myrist. (1897) 326 (type of basionym
Myristica lemanniana excluded).
Myristica globularia auct. non. Bl.: Hook.f. & Th., Fl. Ind. (1855) 160; A. DC., Prod, 14, 1 (1856)
202, p.p., for the specimens from Malacca.
Twings 2-3 mm diam. towards the apex; leaf bud covered with hairs (0.2-)
0.3-0.6 mm long. Leaves 7-19 x 2.5-6 cm, lateral nerves 6-15 pairs. Male perianth
1.0-1.5 mm diam.; anthers (3 or 4)-7. Female perianth 2.0-2.5 mm long. Fruits
1.9-2.5(-2.8) X 1.4-2.0; cm, pericarp 2-4 mm thick.
Distribution. Malaya, Singapore, Sumatra, Borneo (Sarawak, rare; Sabah, E.
Kalimanthan), Philippines (Mindanao, | collection).
MALAYA. (Kedah, Perak, Trengganu, Pahang, Selangor, Malacca, Johore): Derry 1216; FRI 0491,
0642, 0650, 2795, 4348, 4357, 4381, 4435, 4483, 11348, 13172, 14304, 16448, 17279, 021619, 023913,
27532, 28361; Griffith 4354; Hassan Rani H. 86; Kadim & Noor KN. 185, KN. 422; KEP 21593, 38129,
71903, 83478, 85230, 85231, 98919, 99802; King’s Coll. 3309, 7526, 10431; Maingay 1002, 1003A, 1003,
1286; Shah & Noor MS. 1947; Ridley 4162, 7629; Scortechini (21la) 91la; SFN, 29366, 31976, 32109,
36122, 40288.
SINGAPORE. Maxwell 82-226; SFN (Ngadiman) 34630, (Sinclair) 39601, 40681, 40711
SUMATRA. Northern (Langkat): de Wilde & de Wilde-Duyfjes 19390, 19484, 19485 — East Coast:
Bartlett 7304 — Central (Paryakumbuh): Maradjo 285 — Riau: Soepadmo 10] — Djambi: Posthumus
833, Roos & Franken 1968 — Palembang; b.b. E. 85] —Bangka: Kostermans & Anta 593.
BORNEO. Sarawak (lst, 4th & 7th Div.): S 38598, 39112, 40980, 41279 — Sabah: Puasa Angian 3918;
San 17186, 17244, 21159, 21487, 30450, 31129, 32209, 32283, 37103, 49349, 53020, 62802, 73995, 76070,
80798, 80957, 82599 — Kalimantan: West (Landak), Teysmann s.n. — E & NE.: Kostermans 8658,
9285; Meijer 2137, 2398; Nedi 719 — SE.: Hubert Winkler 2526.
PHILLIPPINES. NE. Mindanao (Agusan Prov.): Mendoza 61-409 (= PNH 42247).
Ecology. Lowland forest, most often on sandy soils; also Casuarina forest (E.
Borneo), fresh water swamp forest, ridge-top forest, kerangas forest (Sarawak,
rare); 0-900 m alt. Flowers throughout the year, but most collections June to
September; fruits throughout the year.
Vernacular names. Tjemanding (Palembang, Sum.), Manggoe mangkiras (Land-
ak, West Kalimantan).
20 Gard. Bull. Sing. 39(1) (1986)
NOTES
1. Fieldnotes. Slender tree with narrow crown, the branches often almost in
whorls. Bark fissured, rarely flaky; inner bark mostly yellow to reddish, fibrous.
Wood whitish to ochre-brown. Flowers at first jade-green, at anthesis yellow to
orange-yellow with faint sweet odour when crushed. Fruits greenish yellow to
orange.
2. The type-variety var. polyspherula agrees with Sinclair’s H. polyspherula as
treated in Gard. Bull. Sing. 16, 1958, p. 422 and largely with H. polyspherula var.
polyspherula as treated in Gard. Bull. Sing. 28, 1975, p. 102.
3. The leaves, as in var. sumatrana are usually brittle when dry and often
fragmented in herbaria. Dry leaves are usually greenish above and contrast well
with the light to chocolate-brown of the lower surface. The tomentum of the leaf
bud and young twig apex is rather long, the hairs 0.2-0.6 mm, 1.e., on the average
slightly longer than in the other varieties.
4. As var. polyspherula is mainly characterized by smaller fruits, it is often
difficult to tell whether specimens with fruits just over 25 mm are those of var.
sumatrana or var. polyspherula.
5. Specimens from the kerangas forest of Sarawak (e.g., S 35598) somewhat
deviate by their stouter habit and less contrasting colour of the dry leaves, being
more brownish yellow rather than greenish. Var. polyspherula is apparently rare in
Sarawak.
b. var. sumatrana (Miq.) de Wilde, comb. nov.
Myristica glabra Bl. var. sumatrana Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 49 — Horsfieldia
brachiata (King) Warb. var. sumatrana (Miq.) (Sinclair ex Whitmore, Tree Flora Malaya 1 (1972)
325, nom. inval., basionym wrongly cited and without literature ref.) Sinclair, Gard. Bull. Sing. 28
(1975) 13 (p.p., excl. syn. H. majuscula and H. bartlettii) — Type: Korthals s.n. (L; iso: K, U; Bt; A,
BP, BR, MEL. S, n.v.), W. Sumatra.
Myristica integra Wall., Cat. (1832) no. 6799, nom. nud.
Myristica collecttiana King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 312, pl. 147 — Syntype: Malaya, King
3620 (CAL, n.v.), 3899, lecto (CAL, n.v.; iso: K, P: FI, G, n.v.), 6672 (CAL., n.v.; iso: PDA, SING,
n.v.), 6737 (CAL, n.v.; iso: BM. K.L).
Horsfieldia subglobosa auct. non (Mig.) Warb.: Warb., Mon. Myrist. (1897) 328; Gamble, Mat. FI.
Mal. Pen. 5, 23 (1912) 220; Ridley, Fl. Mal. Pen. 3 (1924) 60 — H. subglobosa [non (Miq.) Warb.]|
var. subglobosa auct.: Sinclair, Gard. Bull. Sing. 16 (1958) 425, 426, fig. 48-50, SLA-D (p.p., excl.
syn. H. majuscula).
Twigs 2-4 mm diam. towards apex; leaf bud covered with hairs 0.1-0.4 mm long.
Leaves 13-28 x 4.5-9 cm, lateral nerves 12-15 pairs. Male perianth 1.2-1.8 mm
diam., anthers 6 or 7. Female perianth c. 3 mm long. Fruits (2.5-) 2.8-3.5 x 2.2-2.7
cm, the pericarp 3-5 mm thick.
Distribution. Malaya, Singapore, Sumatra, Borneo.
MALAYA (specimens seen from Kedah, Perak, Kelantan, Pahang, Selangor, Negri Sembilan,
Johore): FRI 1138, 1690, 2637, 3026, 4012, 5776, 6900, 6959, 7851, 9353, 11610, 12339, 12498, 13823,
20400, 25263; KEP 95006, 98976, 100129, 104913, 104996, 115988; King’s Coll. 3899, 6004, 6737; Shah
& Kadim 362; Md Nur 34117; SFN 32064, 35779; Soepadmo c.s. F.S.C. 831.
New account of Horsfieldia 4 Bh
SINGAPORE. SFN 39599, 39697, 39989, 40043; Wallich 6799.
SUMATRA. W. Coast: Korthals s.n. — East Coast: Bartlett 6867; Krukoff 4138; Soepadmo 15, 193
— Siberut Isl.: Jboet 31] — Belitung: Vorderman (50).
BORNEO. Sarawak: S 12751, 13987, 16956, 20888, 34059, 34813, 37289, 37881, 38576, 39574, 39761
— Brunei: Fuchs & Muller 21168; Van Niel 4053; Sinclair & Kadim 10414 — Sabah: Elmer 21338, 21364;
San. 16284, 25045, 28152, 30403, 32932, 46674, 63866, 72321, 73228, 78219, 82022. — Kalimantan.
West: Hallier 21/41; East: Endert 2114, 5088, 5091; Kostermans 4448, 6093, 7721, 10176, 10696, 12613;
Wiriadinata 1194; NE. (Nunukan Isl.(: Kostermans 8746; Meijer 2032, 2259; Payments 5.
Ecology. Lowland mixed dipterocarp forest, ridge forest, montane forest, mossy
dipterocarp forest; also in heath forest, peat swamp forest, swamp forest and
kerangas; on sandy and sandy-loamy soils, ‘red’ soil, yellow clayey soil; 0-1100 alt.
Flowers and fruits throughout the year.
Vernacular names. Kajang (Lubok Antu reg., Sarawak, 2nd. Div.); Kumpang
lusoh (Sarawak, Semengoh F.R., Ist Div.; it means ‘lazy kumpang’ which refers to
the slow combustion of wood which is not dry.
NOTES
I. Fieldnotes. Tree usually slender, bole straight, without buttresses, crown
slender, branching monopodial, branches horizontal. Bark generally dark brown,
rather smooth, shallowly to fairly fissured, sometimes flaky (strips 10-20 mm wide).
Inner bark reddish, fibrous, laminated, kino profuse, colourless then deep red;
slash wood (sap-wood) whitish to cream; cambium whitish; heartwood pinkish.
Flowers yellow to waxy yellow; fruits glossy green, turning greenish yellow, yellow,
or orange; aril bright orange-red to red.
2. The tomentum of the leaf-bud in specimens from Sumatra (incl. the type of
var. sumatrana), is quite short, the hairs only c. 0.1 mm long; the tomentum in
specimens from Malaya and Borneo is usually longer, and rough with hairs c. 0.4
mm long.
3. Where fruits are less than 30 mm long, leaves smallish and twigs rather
slender, specimens may be difficult to distinguish from var. polyspherula.
4. Some specimens from East Kalimantan such as Kostermans 7721 (Samarin-
da), 10696 (Central Kutai; both & flowers), and 126/73 (W. Kutai, fruits) slightly
deviate in habit by their thinner, membranous leaves, which dry to a paler brownish
colour; the flowers and fruits are not different, although Kostermans remarks that
the fruits of 26/3 are wine red, a colour as yet unrecorded for the other, fairly
abundant, fruiting collections.
5. Deviating specimens. Two, King’s Collector (Goping) 6004 (syntype of H.
majuscula, not the lectotype), from Perak, Malaya, c. 200 m alt., with male
flowers, and Md, Nur 34117, from Selangor, at low alt., also with male flowers,
obviously belong to the H. polyspherula-complex according to the subglobose
shape of the male perianth, and shape and architecture of the androecium. As
regards their general habit both specimens agree with var. sumatrana, but differ by
their larger perianth, c. 1.8-2.0 mm diam, and, accordingly, larger androecium, c.
1.0 x 0.8 mm; there are 5 or 6 anthers (10 or 12 thecae). Possibly they link up with
the normal flowers of var. sumatrana, which is mainly characterised by fruit-size,
and of which the true variation in flower size I am not certain of.
22 Gard. Bull. Sing. 39(1) (1986)
c. var. maxima de Wilde, var. nov.
Ramuli haud prominulo-lineati. Folia nervis lateralibus supra prominentibus instructa. Fructus sicco
(3.5-)4-6 x 3-5 cm, pericarpio (5-)8-15 mm crasso. — Type: Sarawak, 7th Div., Paul Chai S 36228 (L;
iso: K; SAR, KEP, MO, SAN, n.v.).
Twigs towards apex 2-4 mm diam.; leaf bud covered with hairs 0.2-0.5 mm long.
Leaves 9-20 x 3.5-7 cm, lateral nerves (6-)9-16 pairs. Flowers not seen. Fruits
(3.5-)4.0-6.0 x 3.0-5.0 cm, the pericarp (5-)8-15 mm thick.
Distribution. Borneo
BORNEO. Sarawak: S 17027, 32244, 36228, 37188, 38498 — Sabah: San. A 1735, 25593, 48793,
66747, 86049. — E. Kalimantan: Kostermans 13023; Leighton 908.
Ecology. Mixed forest, Agathis forest; on sandy water-logged soil, sandy loam or
yellow clay-loam soil; 50-500 m alt. Fruits throughout the year.
NOTES
1. Fieldnotes. Tree recorded both as with and as without buttresses. Bark shal-
low boat-shaped fissured. Inner bark pinkish. Wood medium soft, whitish yellow.
Fruits yellow to red.
2. The present variety differs mainly by its conspicously large and almost globose
fruits with a very thick pericarp. It may be confused with H. majuscula from
Malaya, a species which was reduced by Sinclair (I.c., p. 13) to H. brachiata var.
sumatrana (the latter presently treated as H. polyspherula var. sumatrana). H.
majuscula differs from H. polyspherula by the size of the male flowers (the flowers
of var. maxima are unknown) with a differently shaped androecium, by the diffe-
rent leaf colour and by the slightly different fruits with a thinner pericarp.
Fruiting specimens of H. polyspherula var. maxima may also be confused with H.
punctatifolia, which differs by its typical punctate leaves.
82. Horsfieldia oligocarpa Warb. Fig. 1C(82)
Horsfieldia oligocarpa Warb., Mon. Myrist. (1897) 354, t. 22 fig. 1-3 — Myristica oligocarpa (Warb.)
Boerl., Handl. Fl. Ned. Ind. 3. 1 (1900) 87 — H. polysherula (Hook. f.) Sinclair var. oligocarpa
(Warb.) Sinclair. Gard. Bull. Sing. 28 (1975) 104 — Syntype: Beccari 2066 (fruits, FI acc. no. 7620,
lecto; iso: K. P), s.n. (CO fl., FI acc. no. 7621, n.v.).
Tree 4-20 m. Twigs terete, towards the apex 2-3(-4) mm diam., at the very apex
dark grey-brown, early glabrescent, lower down bark striate, generally pale yellow-
ish or whitish-brown, contrasting with the dark colour of the dry petioles, lenticels
small, at first distinct, lower down inconspicuous, bark when older not flaking.
Leaves in 2 rows, chartaceous, elliptic-oblong to oblong, broadest at the middle,
7-16 X 2.5-6 cm, base attenuate, top acute-acuminate, upper surface glabrous,
drying usually grey-greenish, dull, lower surface generally bright rusty or choco-
late-brown, contrasting much with the upper surface, glabrous including midrib,
without blackish marks; midrib above raised, glabrous; nerves 8-11 pairs, above
slender, raised; tertiary venation indistinct or invisible on both surfaces: petioles
6-12 x 1.5-2.5 mm, drying blackish brown; leaf bud 7-10 x 2-3 mm, covered with
New account of Horsfieldia 4 23
dense grey-rusty tomentum of hairs c. 0.2 mm long, the tomentum early shed in the
form of small crust-like pieces. Inflorescences rather sparsely pubescent with
stellate hairs 0.2-0.3 mm long, sometimes glabrescent, in C’: 2 or 3 times ramified,
3-6 X 1.5-4 cm, common peduncle 5-10 mm long, few-flowered, the flowers in
clusters of 3-8 each; 2 inflorescences not seen, according to the infructescences c.
1.5-4 cm long; bracts not seen, caducous; flowers 3-valved, perianth glabrous,
pedicel glabrous, at base articulate. Male perianth globose to broadly obovoid,
1.0-1.7 mm diam., top broadly rounded, base + rounded, glabrous; pedicel 1-1.5
mm long, slightly tapering; perianth at anthesis cleft to nearly Y%2-way, +
coriaceous, not collapsing on drying, valves towards base up to 0.5 mm thick.
Androecium (incl. androphore) + obovoid, c. 1.0 x 1.0 mm, in transverse section
triangular; anthers 6 or 7 (thecae 12 or 14), anthers and thecae largely mutually
free, c. 0.5 mm long, tip acutish, at base attached to a short column which continues
into the somewhat tapering androphore c. 0.5 mm long. Female perianth not seen.
Fruits 1-4 per infructescence, ellipsoid, top and base rounded, 1.8-2.7 x 1.4-1.9 cm,
glabrous, drying brown to dark brown, not to hardly lenticellate, pericarp 2-3 mm
thick; stalk 2-5 mm long; perianth not persisting.
Distribution. Borneo (Sarawak, Brunei)
BORNEO. Sarawak (lst, 3rd & 4th Div.): Beccaris.n. (Fl acc. no 7621, n.v.), 2066; Native Coll. 821;
S 16219, 19470; Yacub 8255 — Brunei: (Ashton & Whitmore) BRUN 398, 635; Brunig S 4402; Sinclair &
Kadim 10414, 10430, 10452, 10503 (= H. bicolor Sincl. in sched.).
Ecology. Forest on poor soils: white sand, white podsolic sand, yellow sand,
sandstone, ‘terraces’, sand and peat, once recorded from a ridge; heath forest; 0-50
m alt. Flowers in June, August; fruits August to October.
Vernacular name. Kumpang puteh.
NOTES
1. Fieldnotes. Shrub or tree, buttresses absent. Bark grey and dark brown, to
reddish brown, fissures c. ¥4-'2 in. wide; c. 9 in. long; inner bark pink brown, soft,
c. 1 cm thick, sap red; sapwood pink-yellow, soft. Timber firm. Leaves pale green,
dull, not glaucous beneath. Fruit pale yellow to orange; pear-shaped, + pointed at
apex. Flowers light brown.
2. This species is identical with H. polyspherula var. oligocarpa as treated by
Sinclair (p. 104), who regards it as a distinct ecological variety. I agree that it is
extremely close to H. polyspherula, especially to var. polyspherula, as it has fruits
of about the same size. Because I accept three varieties within the polymorphous
H. polyspherula, mainly on the basis of different fruit-sizes (though some sizes
overlap), the present H. oligocarpa could be treated as a variety based on quite
different grounds. I prefer to keep it apart as a separate species because it has a
strikingly different general appearance and its own distinct ecology. H. oligocarpa
seems to be restricted to low forest and heath forest on very poor sandy and peat
soils at low altitudes. The plant stands out by the overall pale colour; the older
twigs are pale, whitish brown to straw; the leaves dry a dull pale green above,
contrasting strongly with the bright brown, or rusty, copper, or chocolate-brown of
the lower leaf surface (more contrasting than is usually the case with H. polyspher-
ula); the inflorescences are rather small and not many-flowered, + glabrescent; the
flowers are markedly coriaceous (?always). Sinclair named some of the specimens
24 Gard. Bull. Sing. 39(1) (1986)
as H. bicolor Sincl. in sched. H. polyspherula grows generally on richer soils, and
the twigs dry always to grey-brown to brown; the leaves are generally of a less paler
green; and the perianths are not coriaceous or slightly so.
83. Horsfieldia endertii de Wilde, sp. nov. Fig. 1C(83)
Ramulorum apices atque gemmae pubescentes pilis scabris 0.3-0.6 mm longis. Folia coriacea. Perian-
thium masculum 3-valvatum, ellipsoideum, 2.5-3.5 x 2-2.5 mm. Androecium ellipsoideum, sessile,
antheris 10-14, sessilibus, pedicello basi non-articulato. — Type: E. Kalimantan, Endert 3996 (L; iso:
K; A, BO, n.v.).
Tree 4-25 m. Twigs terete, towards the top 2.5-4(-8) mm diam., dark or blackish
brown, early to rather late glabrescent, tomentum rough, deep rust, composed of
hairs 0.3-0.6 mm; bark coarsely striate, with coarse and conspicuous, paler len-
ticels, older bark tending or not to flake. Leaves in 2 rows, coriaceous to strongly
coriaceous, elliptic-oblong to oblong-lanceolate, broadest generally at the middle,
8-17(-26) x 3-6(-10) cm, base rounded to (short-)attenuate, tip rounded to sub-
acute; upper surface glabrous but base of midrib late glabrescent, drying oli-
vaceous or yellowish to dark-brown, lower surface drying pale brown to chocolate,
not much contrasting with the upper surface, without larger blackish marks but
with usually large and conspicuous, pale yellowish hair-scars (lens, X 60), glabrous
but the midrib towards the base sometimes late glabrescent; midrib above relatively
broad, raised, glabrous but towards base late glabrescent; nerves 8-15 pairs, above
sunken, flattish, or slightly raised, the submarginal arches fairly regular-shaped and
sometimes distinct; tertiary venation forming a lax network, generally indistinct or
invisible on both surfaces; petioles 6-16 x 2-3.5(-4) mm, glabrescent; leaf bud 10-20
x 3-4 mm, with hairs 0.3-0.6 mm. Inflorescences densely and rather shaggy-
pubescent with rusty hairs 0.5-1.0 mm long, in ©’: 2 or 3 times ramified, not very
many-flowered, (1.5-) 3-10 x (1-) 2-5 cm, common peduncle 5-20 mm long, the
flowers in clusters of (1-)2-6 each; 9 inflorescences + few-flowered, | or 2 times
ramified, 2-4 cm long; bracts broadly ovate-ellipsoid, 3-7 mm long, densely pubes-
cent, caducous; flowers 3-(or 4-) valved, perianth glabrous (glabrescent), pedicel
minutely puberulous, hairs 0.1-0.3 mm, especially in the lower half, at base inar-
ticulate. Male perianth + obovoid (when immature) to ellipsoid, 2.5-3.5 x 2.0-2.5
mm, top and base rounded, glabrous; pedicel 2-3 mm long; perianth at anthesis
cleft to 4-'2-way, only slightly collapsing on drying, valves c. 0.3 mm thick.
Androecium + sessile, obovoid to truncate-ellipsoid, 2.0-2.8 x 1.4-1.6 mm, in
transverse section sub-triangular; anthers (10-)12-14 (c. 24-28 thecae), almost
completely sessile, 2.0-2.8 mm long, free at apex for only 0.1 mm or less, column
solid except for the narrow apical hollow or slit reaching to “5-3 deep; androphore
narrow, (0-)0.1-0.3 mm long, hidden by the anthers. Female perianth broadly
ovoid-ellipsoid, 2.5-3 x 2-2.5 mm, glabrous, split at anthesis to 43-2, valves 3, at
sutures 0.3-0.5 mm thick, pedicels 1.5-2.0 mm long, towards the base with hairs c.
0.2 mm, inarticulate, ovary ovoid-ellipsoid, c. 2 x 1.5 mm, glabrous, stigma c. 0.2
mm high, minutely 2-lobed. Fruits 1-4(-8) per infructescence, ellipsoid, base round-
ed, top rounded to acutish, 3.0-4.2 * 1.6-2.4 cm, glabrous, drying grey-brown to
dark brown, without or with a few lenticel-like tubercles, pericarp 2-4 mm thick;
stalk 3-5 mm long; perianth not persisting.
Distribution. Borneo: Sarawak, Sabah, E. Kalimantan.
BORNEO. Sarawak (4th & Sth Div., Baram Dist.): Anderson 4258: Nielsen (Mulu Exp.) 792;
S 15083, 33027, 35486 — Sabah (mainly Kinabalu): Chew, Corner & Stainton 142; Clemens 29558,
New account of Horsfieldia 4 25
32500, 32605, 50721; (Wyatt-Smith & Wood) Kep 80353; Nooteboom 1044; San 33940, 36778, 65314,
76815, 79585 — E. Kalimantan (W. Kutai): Endert 3996.
Ecology. A montane species; in ridge forest, mossy forest, mountain forest,
dwarfed forest on wind-swept crests; on sandy soil, black or brownish soil; 1200-
2100 m alt. Flowers and fruits throughout the year.
Vernacular name. Binarak (Murut lang., Sarawak).
NOTES
1. Fieldnotes. Tree of medium size, sometimes dwarfed. Bark cracked to finely
fissured, brown-black to dark brown; inner bark reddish; cambium whitish, sap-
wood whitish; exudate turning reddish. Fruit orange-yellow, pink, pink-red, or
orange-red, aril bright orange. Flowers yellow.
2. At first Sinclair identified most specimens (incl. the type) as H. polyspherula
var. montana, or as H. montana Airy Shaw; later on he included H. montana as
well as the superficially similar H. xanthina in his polymorphous concept of H.
glabra. In the present revision I have kept H. montana and H. xanthina as distinct
species differing from H. glabra and from the material here described as a new
species. H. glabra and H. xanthina differ in the flowers, and vegetatively by the
much shorter tomentum on the leaf bud, twig-apex and inflorescences; H. montana
differs by the much smaller globose flowers. H. endertii is characterized by fairly
large ellipsoid male flowers with an androecium distinctly longer than broad, the
pedicels inarticulated at base, very coriaceous leaves, rough-haired leaf buds and
inflorescences. On the lower leaf surface there are practically always distinct, large,
pale-yellowish coloured hair-scars, well visible when magnified 60 times. H. ender-
tii usually has + rounded leaf tips, as in H. montana, but the leaves in the latter
usually dry to a blackish colour and are generally smaller and thinner.
3. The type, Endert 3996, with male flowers, is from W. Kutai, the only speci-
men from Kalimantan. The remaining specimens from Sarawak and Sabah some-
what differ in general appearance; their twigs are often less roughly hairy towards
the top, the leaves are somewhat more coriaceous with the lateral nerves ascending
at a slightly sharper angle from the midrib; all male flowering specimens from
Sarawak and Sabah appeared to be in a juvenile state, except the BM duplicate of
Clemens 29558 (Tenompok), of which the identity with the type is evident.
4. The collection Chew, Corner and Stainton RSNB 142 from Kinabalu some-
what deviates by the relatively large, broad and very coriaceous leaves; it has
immature male inflorescences but the lower leaf surface has typical, large, golden
or whitish hair scars.
84. Horsfieldia valida (Miq. ) Warb. Fig. 1C(84)
Myristica valida Miq., Fl. Ind. Bat. 1 (2), 1 (1858) 67; Suppl. 1 (1860) 156 — Horsfieldia valida (Miq.)
Warb., Mon. Myrist. (1897) 349; Heyne, Nutt. Pl. (1927) 638. — Type: Sumatra, West Coast,
Teijsmann 479 (U; iso: BO, n.v.) (sterile, the fruit said to be as large as a goose egg).
Tree 10-15 m. Twigs terete, towards apex 2.5-5 mm diam., at insertion of
inflorescences up to 10 mm diam., brown to grey-brown, rather early glabrescent,
26 Gard. Bull. Sing. 39(1) (1986)
tomentum rusty woolly, with hairs 0.4-0.7 mm long; the bark lower down coarsely
striate, rather densely set with conspicuous pustulate lenticels; older bark not
flaking. Leaves in 2 rows, chartaceous, (ob)ovate-oblong, broadest at or somewhat
above the middle, 20-35 x 8-13 cm, base short- to long-attenuate, tip subobtuse to
acutish; upper surface glabrous, drying dark- to olivaceous-brown, lower surface
drying bright brown, without blackish marks, early glabrescent but midrib rather
late glabrescent; midrib above rather broad towards the base, glabrescent, slightly
raised; nerves 20-25 pairs, raised above; tertiary venation forming a rather lax
network, flat, indistinct or invisible on both surfaces; petioles 7-12 x 3-4.5 mm,
glabrescent; leaf bud c. 15-20 x 4mm, densely ferrugineous-pubescent with hairs c.
0.4-0.7 mm. Inflorescences situated just behind the leaves, + woolly pubescent
with hairs 0.3-0.5 mm, subglabrescent, in O: 2 or 3 times ramified, not very
many-flowered, 5-6 X 3-4 cm, common peduncle 10-15 mm long; @ inflorescences
not seen, according to the infructescences c. 5 cm long; bracts not seen, caducous;
flowers 4-(or 3-)valved, in loose clusters of 3-6 each, perianth glabrous, pedicel
glabrous and inarticulate at the base. Male perianth globose, 2.5-3 x 3-3.5 mm, top
and base rounded, pedicel c. 1.5 mm long; perianth at anthesis cleft to c. %, slightly
collapsing on drying, valves c. 0.3-0.4 mm thick. Androecium + depressed globose,
0.8-1.2 xX 1.4-6 mm (much smaller than the perianth); in transverse section +
irregularly rounded or faintly 4-angular; anthers 12-14, curved and largely connate
with the broad column, at apex free for only c. 0.2 mm; column not or hardly
hollowed; androphore rather narrow, 0.1-0.2 mm long. Female flowers not seen
(but see under fruits). Fruits 2-9 per infructescence, ellipsoid, top and base +
rounded, c. 8.0(-9.0) x 5.0(-6.0) cm, glabrous, drying brown with surface wrinkled
and + warted, pericarp c. 15 mm thick; stalk c. 5 mm long; perianth persistent
under young fruits in Maradjo 449, 4-valved, c. 3 mm long.
Distribution. Sumatra (E. and W. Coast; possibly Palembang, see notes), prob-
ably W. Borneo (see note 3).
SUMATRA. West Coast: Maradjo 449; Teijsmann 479 — East Coast: Lorzing 5896, 15557 —
Palembang: Dumas 1649 (sterile).
BORNEO. West Kalimantan, Mt. Damoes: H. Hallier 624 (doubtful, see notes).
Ecology. Primary forest, ravine forest; (?200m, see the notes —) 900-1100 m alt.
Flowers in March and August, fruits in August.
Vernacular names. Simar mudar-mudar (Timor lang. of Sumatra), Lundang (W.
Coast), Pijangoe pématang (Palembang).
NOTES
1. Fieldnotes. Tree erect, branches wide-spreading and arching. Flowers tinged
_ yellow, brown, sweet smelling.
2. The specimen Dumas 1649, from Pelambang, is sterile; it deviates from the
other specimens (all from 900-1100 alt.) by the shorter tomentum on the leaf bud
(hairs only c. 0.3 mm long) and its lower (c. 200 m.) altitude.
3. Deviating specimen from W. Borneo. Hallier 624, with Cf fl., of Mt. Damoes,
is obviously taxonomically closely related to H. valida, but it is markedly different
in several ways. Apparently it represents a new undescribed species. It keys out
New account of Horsfieldia 4 27
beside H. valida because its male perianths are cleft at anthesis to c. “% or deeper,
and the pedicels inarticulate at the base. It cannot be H. fragillima (pedicels also
inarticulate, and keys out next), because that species has the perianths at anthesis
cleft at most to 2-way deep, and a saucer-shaped androecium. Our specimen
differs from H. valida by the much smaller flowers, globose perianth (which is
possibly submature) measuring 1.2-1.5 x 1.5-1.6 mm; smaller androecium, 0.5-0.6
x 0.8-1.0 mm with the column at apex broadly hollowed to about '2-way deep,
anthers + completely sessile, + incurved and concealing the hollow (in H. valida
the column not or hardly hollow at the apex); there are fewer anthers, 9-11 (in H.
valida 12-14). It has predominatly 4-valved perianths as in H. valida from Sumatra.
The C inflorescence is 21 cm long, those of H. valida only c. 6 cm. I have not found
Q-flowering or fruiting specimens which match Hallier 624, and I have refrained
from describing it as a new species. The specimen was determined by Sinclair as H.
fragillima.
4. Sinclair (p. 150) accepted H. valida in a much wider sense. As distribution he
included China, most of Borneo, the Philippines, and Celebes. Most of the speci-
mens cited by him for Sumatra have at present been included in H. valida; other
specimens have been assigned by me to various species. The only one from Borneo
(see under note 3, deviating specimen) was included by Sinclair in H. fragillima.
5. Warburg (p. 349) remarks that it is vegetatively very alike H. macrothyrsa,
differing almost only in the fruits which were recorded as unusually large. I disagree
as H. macrothyrsa differs in many ways, such as the punctate leaves and the much
larger male flowers.
85. Horsfieldia borneensis de Wilde, sp. nov. Fig. 1C(85)
Ramulorum apices atque gemmae pubescentes pilis ferrugineis 0.2-0.4 mm longis. Folia tenuiter
coriacea, subtus praedita punctis atque lineis brevibus sparsis fusco-brunneis. Perianthia mascula
3-valvata, subglobosa ad late obovoidea, 1.3-1.8 X 1.2-1.7 mm, androecio subgloboso sessili, antheris
7-10, sessilibus, pedicello glabro, basi articulato. — Type: Sarawak, Bojang bin Sitam S 14610 (L iso:
K).
Tree 10-30 m. Twigs subterete, not ridged, (1.5-)2-4(-10) mm diam. towards the
apex, usually dark grey-brown, sometimes blackish, not conspicuously hollow,
early to rather late glabrescent, tomentum rusty, composed of hairs 0.2-0.4 mm,
bark lower down faintly finely striate, not distinctly lenticellate, when older finely
cracking or not, flaking slightly or not. Leaves in 2 rows, chartaceous to thinly
coriaceous, elliptic-oblong to oblong-lanceolate, broadest usually at the middle,
7-18 X 2-6 cm, base (short-) attenuate, top acute to (short-) acuminate; upper
surface drying dull olivaceous to (partially) blackish-brown, glabrous, the midrib
not or somewhat late glabrescent, lower surface drying pale brown to bright reddish
brown or chocolate, glabrescent, tomentum of shortish, densely-branched den-
droid hairs, 0.3-0.4 mm (especially on midrib), always with scattered, usually
subcircular, larger, blackish dots or marks (cork warts); midrib slightly raised
above, early or late glabrescent; nerves 10-16 pairs, slender above, usually flat or
sunken (or only close to the midrib slightly raised) or in thinner-leaved specimens
slightly raised, glabrous, the lateral arches + regularly shaped, not very distinct;
tertiary venation hardly or not visible on both surfaces; petioles relatively long,
12-25 X 1.5-2.5 mm, early to rather late glabrescent; leaf bud 10-17 x 2-4 mm,
densely pubescent with hairs c. 0.3 mm long. Inflorescences behind the leaves,
ps: Gard. Bull. Sing. 39(1) (1986)
densely short-woolly pubescent with rusty hairs up to 0.7 mm long, in CG: fairly
large, many-flowered, c. 4 times ramified, (8-)13-20 x (5-)10-18 cm, common
peduncle 15-35 mm long; @ inflorescences not seen, infructescences up to 13 cm
long; bracts elliptic to elliptic-oblong, pubescent as the inflorescences, 1.5-5 mm
long, caducous; male flowers in loose clusters of 2-6 each; perianths glabrous,
3-valved, pedicels glabrous, distinctly articulate at base. Male perianth subglobose
to broadly ellipsoid or broadly obovoid, 1.3-1.8 * 1.2-1.7 mm, top broadly round-
ed, base (narrowly) rounded, glabrous; pedicels 1.0-1.5 mm; perianth at anthesis
cleft to c. 4% (to nearly 2), not collapsing on drying, valves 0.2-0.3 mm thick.
Androecium broadly ellipsoid to subglobose, 0.7-1.2 x 0.6-1.3 mm, the top broadly
rounded, slightly impressed in the centre, base rounded, in cross-section subcircu-
lar; anthers 7-10 (thecae 16-20), almost completely sessile and mutually closely
appressed, at apex incurved over the cavity which reaches to c. “%-/% (or less) deep;
free apicies + none; column broad; androphore narrow, c. 0.1 mm or less long.
Female flowers not seen; as judged from remnants under very young fruits:
perianth c. 2.5 mm long, ovary glabrous. Fruits 1-7 per infructescence, ovoid,
somewhat laterally flattened and slightly flanged, top and base rounded, 4.0-6.0 x
3.0-4.5 cm, glabrous, drying brown and often with a glaucous tinge, rather smooth,
pericarp (measured dry) 10-15 mm thick; stalk stout, 4-6 mm long; perianth not
persisting.
Distribution. Borneo: Sarawak, Sabah, E. & NE. Kalimantan.
BORNEO. Sarawak (all Kuching & vicinity, Ist. Div.): Asah Arb. no. 715; S 12771, 14610, 14759,
25241, 34706 — Sabah (Beaufort Dist.): San 16838, 31413 — E & NE. Kalimantan (W. Kutai,
Balikpapan, Nunukan Isl.): b.b. 16516, 18180, 29301, 29315, 29373, 34325; Kostermans 7043, 8617,
9782, 9947, 13623; Schut K. 31.
Ecology. Primary lowland dipterocarp forest; on sandy soils, flat clay soil,
sandstone, sandy ridges; 0-200 m alt. Flowers in April, August and September,
fruits throughout the year.
NOTES
1. Fieldnotes. Bark of trunk rough, deeply fissured, flaking in squares, usually
dark brown or reddish or blackish; strips or flakes up to 5 cm wide, up to 1 cm
thick. Living bark 5-10 mm thick, red-brown; sap red; sapwood c. 10 cm, reddish
white to pale red, heartwood red-brown. Bark of tree c. 30 cm diam. recorded as
deeply fissured, but with a smooth appearance, the strips with rounded edges.
Fruits bluish green, turning green-yellow to yellow or reddish, pericarp pink inside,
aril orange.
2. Sinclair included most of the specimens of the present new species in H.
wallichii, a few were determined by him as H. polyspherula s.1. or H. aff. ridleyana
(e.g. Achmat b.b. 34325). Our present species has in common with H. wallichii the
characteristic blackish dots on the lower leaf surface, the dull upper leaf surface
with largely sunken nerves, and practically similar fruits though they have a
persistent perianth in H. wallichii. However, H. wallichii, which also occurs in
Borneo, is generally stouter, has much larger leaves, often with a persistent
tomentum. Above all, it differs in the flowers, the general appearance and shape
and structure of the androecium, and the pedicel which is inarticulate at the base.
3. On account of the dotted lower leaf surface, H. borneensis belongs to the
group of species including H. wallichii and H. pulcherrima whereas the structure of
New account of Horsfieldia 4 29
the androecium links it up with species such as H. pulcherrima, H. flocculosa, H.
grandis etc.
4. Variation. A few specimens (especially San 16838, 31413, from Beaufort
Dist., Sabah) have relatively thin leaves with the lateral nerves on the upper surface
raised; the dry leaves and twigs have a relatively dark blackish colour. These
specimens may be mistaken for H. polyspherula.
86. Horsfieldia fragillima Airy Shaw Fig. 1C(86); 29
Horsfieldia fragillima Airy Shaw, Kew Bull. 1939, no. 10 (1940) 542 — Type: Sarawak, 4th Division.
Mt. Dulit.Richards 2602 (K; iso: A, SING, n.v.).
Tree 10-30 m. Twigs stoutish, terete or faintly angular, 2.5-7(-15) mm diam.
towards the apex, early glabrescent, tomentum with hairs c. 0.3 mm, bark of young
twigs dark brown, often with conspicuously pale lenticels, bark of older twigs
usually paler brown, coarsely striate with a tendency to crack longitudinally and to
flake somewhat; lenticels distinct, on older wood inconspicuous. Leaves in 2, or
sometimes (partly) in 3 rows, membranous to thinly coriaceous, oblong to oblong-
lanceolate, broadest at or somewhat above the middle, 20-45 x 6.5-12 cm, base
narrowly rounded to short-attenuate, but blade rather tapering below the middle,
base rarely long-attenuate, tip acute or acute-acuminate; upper surface drying
olivaceous to brown, lower surface early glabrescent, often with some minute
tomentum remaining at and near the base of the midrib; without blackish dots:
midrib above raised, towards the base rather broad and flat; nerves 20-30 pairs,
above raised, the marginal arches + indistinct; tertiary venation forming a lax
network, faint to nearly invisible above; petioles stout, relatively short and broad
and often + pulvinate, 4-13(-20) x 3-8 mm, glabrous; leaf bud 12-20 x 3-5 mm,
densely pubescent with hairs c. 0.3 mm. Inflorescences behind the leaves, rather
sparingly set with hairs (0.1-)0.2-0.4 mm, in CG: large, very many-flowered, 4 or 5
times ramified, 15-30 x 10-20 cm, common peduncle 25-60 mm long; in Q: 6-13 cm
long, stout, rather few-flowered (known from infructescences only); bracts up to 12
x 5 mm, tomentulose, caducous. Flowers 3-(or 4-)valved, glabrous, in © in
clusters of 2-6; pedicels glabrous, at base inarticulate. Male perianths globose or
sometwhat depressed-globose, 1.4-2.0 x 2.0-2.5 mm, top broadly rounded, base
rounded; pedicels 1-1.5(-2.0) mm long; perianth at anthesis cleft nearly to ¥2-way.
valves c. 0.2(-0.3) mm thick. Androecium strongly depressed-globose, almost
saucer-shaped, much and broadly depressed in the centre, (sub-)circular in trans-
verse section, 0.5-1.0 x 1.0-1.5 mm; anthers 7-9, almost completely sessile, in-
curved towards the top, free apices up to 0.1 mm; column broadly saucer-shaped.,
with a broad apical hollow to about '2-way deep, androphore (if present) narrow.
up to 0.1 mm long. Female flowers not seen; perianth 3- or 4-valved, 4-5 x 3-4 mm
according to the remnants under the fruit. Fruits up to 8 per infructescence, broadly
ellipsoid, rounded at top and base, possibly slightly flattened, c. 6.0-8.0 x 4.0-6.0
cm, glabrous, drying dark brown, the surface wrinkled and usually with small and
large pustules or warts, the dry valves 10-20 mm thick; perianth persistent for a long
time under the fruit; stalk 2-4 mm long.
Distribution. Borneo (Sarawak, Brunei, Sabah, C. Kalimantan; possibly E.
Kalimantan, see note).
BORNEO. Sarawak: Anderson et al. 15449; Hj. Bakar 4361; Chew Wee-lek CWL. 606; Richards
2602; S 13594, 15137, 16614, 16985, 19535, 23348, 28083, 32141, 34358, 34621, 34968, 35202, 35442,
Fig. 29 Horsfieldia fragillima Airy Shaw
30
a. leafy twig apex, x 2; b, portion of older twig with male inflorescence axilliary to leaf scar,
x 2; c, mature male flower, lateral view, x 12; d, ditto, longitudinally opened, showing
androecium, X 12; e, androecium, longitudinal section, schematic, x 12; f, twig portion with
infructescences, fruit mature, note persistent perianth, x /2— a-e, from S 34358; f, from §
16985.
New account of Horsfieldia 4 31
35756 — Brunei: BRUN 137 — Sabah: B.N.B. For. Dept. A 865; SAN A 3449, A.4211, A 4713, 16519,
16648, 27424, 39622, 50063, 66735, 73695, 74417, 75942, 78243, 83457, 83670, 92578 — Central
Kalimantan: Veldkamp 8451 — E. Kalimantan (Balikpapan): b.b 25571 (doubtful, see notes).
Ecology. Primary lowland dipterocarp forest; hill slopes, flat land, alluvial land,
once in seasonal swamp; most commonly recorded from sandy soils, leached clayey
loam over sandstone, sandy clay, often along or near streams, riverine forest; 0-400
m alt. Flowers February to May, fruits collected all through the year.
Vernacular name. Kumpung Pango (Iban lang.)
Uses. Fruits recorded as edible, very acid and resinous.
NOTES
1. Fieldnotes. Trunk without buttresses, then often broadly gullied, rounded-
fluted, or only with short and rounded buttresses; bark chocolate-brown, reddish-
brown, or brown-blackish, recorded either as not furrowed, slightly fissured, mostly
with few flakes, or scaly, or peeling into thin flakes. Inner bark c. 3 mm thick, pale
pink or reddish to yellowish; sapwood soft, whitish, yellow-pink or pinkish white.
Branches somewhat drooping. Flowers yellow. Fruits ramiflorous, large, up toc. 10
x 7cm. (seed c. 3 X 2 cm), greenish yellow turning rose-pink to red; pericarp thick,
to over 2 cm; aril reddish-orange.
2. On the whole a homogeneous species, only a few specimens from Sarawak
(1st and 4th Div.) markedly vary, viz. Bakar 4361 (CO), and S 13594, 15137, 16985,
34621, 35202 (all in fruit); these deviate from the rest of the specimens by a
somewhat stouter habit. The flowers in Bakar 436] are somewhat larger, reaching
to c. 2.3(-2.5) mm. diam., whereas male perianths of the other specimens from the
same localities reach only to c. 2 mm diam.
3. In the sterile state, H. fragillima may be confused superficially with H.
laticostata, as the former may have a similarly broad basal part of the midrib. H.
laticostata differs by its architecture of the male flowers, much smaller fruits, and
different texture and colour of the leaves.
4. H. fragillima seems most related to H. splendida and H. pulcherrima, which
have resembling androecia (though a narrower hollow), and with an inarticulate
pedicel-base. The two species differ, however, strongly from H. fragillima in
general habit, including their markedly hairy lower leaf surface.
5. In H. fragillima the leaves in fertile plagiotropic shoots are usually arranged in
2 rows, but in a few specimens they are distinctly in 3 rows. In most
Horsfieldias the leaves of plagiotropic shoots are always distichous. The ramiflor-
ous fruiting twigs may reach a width of 25-30 mm.
6. The present species is here accepted largely in the same sense as by Sinclair. |
have to exclude Hallier 624 which may represent a new species; that specimen is
discussed in the notes under H. valida.
7. The collection b.b. 25571 from E. Kalimantan is sterile and deviates by the
rather coriaceous leaves; no other specimen from E. Kalimantan has been col-
lected.
a2 Gard. Bull. Sing. 39(1) (1986)
87. Horsfieldia androphora de Wilde, sp. nov. Fig. 1C(87); 30
Ramulorum apices atque gemmae pubescentes pilis 0.3-0.6 mm longis. Folia membranacea, subtus non
punctata. Perianthia mascula 3-valvata, globosa, 1.4-2 mm diam., synandrio depresso-globoso, c. 1
mm diam., antheris 7-11, androphoro distincto, gracili, 0.3-0.8 mm longo, pedicellis basi non-
articulatis. — Type: Sarawak, Nooteboom & Chai 01710 (L).
Tree 7-20 m. Twigs terete, 2-4(-5) mm diam. towards apex, rather late glabres-
cent, tomentum rusty with hairs 0.3-0.6 mm, lower down the bark rather finely but
distinctly striate, blackish brown, when older not flaking; lenticels small, not
conspicuous. Leaves in two rows, membranous, elliptic to oblong, broadest at
about the middle, 9-18 x 3.5-6.5 cm, base attenuate, tip acute-acuminate; upper
surface drying dark brown or blackish brown, not minutely pustulate, glabrous,
lower surface glabrescent (except midrib), without scattered larger blackish dots;
midrib above raised, beneath with some vestigial tomentum or late glabrescent;
nerves 9-13 pairs, raised above, the marginal arches on the lower surface not very
regularly shaped and not very conspicuous; tertiary venation forming a lax net-
work, distinct or not; petiole 10-12 x 1.5-2 mm, glabrescent; leaf bud 8-12 x 2-3
mm, rusty pubescent with hairs 0.3-0.6 mm. Inflorescences rather densely pubes-
cent with hairs 0.2-0.6 mm, in C: rather many-flowered, 3 (or 4) times ramified,
6-14 x 3.5-9 cm, common peduncle 10-20 mm long; Q-inflorescences (as seen from
infructescences) 3-4 cm long; bracts densely short-pubescent, ovate-elliptic, acut-
ish, c. 3 mm long, caducous. Flowers in © in clusters of 2-6 each, 3-valved,
glabrous; pedicels glabrous or with a few minute hairs 0.1 mm or less at the very
base, at the base inarticulate. Male perianth globose, 1.4-2.0(-2.2) mm diam., base
and apex rounded, pedicels (0.5-) 1.0-2.0 mm, slender; perianth at anthesis cleft to
a depth of 4% to nearly 2-way, valves c. 0.2 mm thick. Synandrium (androecium
minus androphore) depressed-globose, somewhat flattened or impressed at apex
and/or at base, in transverse section subcircular, (0.6-)0.8-1.0 x (0.8-)1.0-1.3 mm,
androphore slender, (0.3-)0.4-0.8 mm long; anthers 7-11 (thecae 14-22), almost
completely sessile and interconnate, curved towards the top and concealing the
apical hollow, hollow 0.2-0.3 mm deep, occupying c. %5-% of the broad column.
Female flowers not seen. Fruits 2-5 per infructescence, ellipsoid, top subacute to
rounded, base rounded or shortly narrowed, 2.4-3.0 x 1.4-2.0 cm, glabrous, drying
dark brown, finely tubercled but without lenticels, dry valves 1.5-2 mm thick; stalk
c. 2 mm long; perianth not persisting.
Distribution. Borneo: Sarawak, Sabah.
BORNEO. Sarawak (Bukit Goram, 4th Div.; Kelabit Highlands, 7th Div.): Nooteboom & Chai
01710; (Chai) S 35443, 36146 — Sabah (Bukit Kinasaraban); Sinclair (Kadim & Kapis) 8977.
Ecology. Montane forest, mossy-forest, wooded sandstone ridges; 800-1200 m
alt. Flowers in March and October, fruits in March, June.
Vernacular name. Li-ang (Kelabit lang., Sarawak, 4th Div.).
NOTES
1. Fieldnotes. Bark surface chocolate- to reddish-brown, bark narrowly cracked
or longitudinally furrowed and cut into rectangular blocks; latex of bark watery,
latex once recorded as more or less colourless (tree in flower), once as blood red
(tree in fruit). Twigs chocolate brown, pubescence rusty. Flowers yellow; fruit
smooth, not pubescent, orange, aril orange, outer seed coat whitish grey.
Fig. 30.
Horsfieldia androphora de Wilde.
a, portion of branch with leafy twig and male inflorescence, x 2; b, mature male flower, X
12; c, ditto, longitudinally opened, showing androecium, X 12; d, androecium, longitudinal
section, schematic, X 12; e, portion of twig with infructescence, fruits mature, aril complete,
x Y% —a-d, from Nooteboom & Chai 01710 (Type); e, from Sinclair, Kadim & Kapis 8977.
33
34 Gard. Bull. Sing. 39(1) (1986)
2. According to the general architecture of the androecium, this species belongs
to the group of H. grandis, but it seems closest related to H. tomentosa from S.
Thailand and Malaya; it has the long-stalked synandrium in common with that
species. H. tomentosa differs by the generally larger flowers, tomentose lower leaf
surface, and smaller fruit which are pubescent or glabrescent. Our present species
is mountainous; H. tomentosa is exclusively found in the lowland.
H. androphora keys out beside H. fragillima (also with inarticulate pedicels), but
the latter differs in many characters: general habit, fruit-size, and quite a different
saucer-shaped androecium.
3. Three of the known specimens of the present species were yet uncollected
when Sinclair revised the genus; his own collection Sinclair 8977 (fr.) was not
recognized as a new species and was included in H. polyspherula var. montana. In
this revision I consider that H. montana.
88. Horsfieldia amplomontana de Wilde, sp. nov. Fig. 1C(88)
Ramulorum apices atque gemmae tomento conspicuo pilorum 0.3-1.5 mm longorum. Folia 15-35 x 5-11
cm, subtus non punctata. Inflorescentiae masculae magnae, usque ad 20 cm longae. Perianthia
mascula 3-vel 4-valvata, subglobosa, 1.5-2 mm diam.; androecio depresso-globoso, 0.6-1.0 x 1.1-1.8
mm, antheris 10-13, sessilibus, androphoro brevi atque angusto, usque ad 0.4 mm longo. Fructus sicci
ellipsoidei, 7-8 cm longo, perianthio persistente. — Type: J. & M.S. Clemens 30536 (L; iso: BM. K).
Tree 10-20 m. Twigs terete, towards the apex 3.5-6(-10) mm diam., grey-brown
to dark brown, early to rather late glabrescent, tomentum brown to rusty, com-
posed of hairs 0.3-1.0(-1.5) mm long; bark coarsely striate, lenticels small, not
contrasting in colour and inconspicuous, older bark not flaking. Leaves in 2 rows,
membranous to chartaceous, elliptic-oblong to oblong-lanceolate, broadest at a-
bout the middle, 15-35 x 5-11 cm, base short-attenuate to narrowly rounded, tip
acute-acuminate; upper surface glabrous (i.e., glabrescent, except towards the base
of the upper midrib in young leaves), drying olivaceous to brown, when dry
contrasting or not with the colour of the lower surface, lower surface glabrous
(glabrescent), without larger blackish or brown dots, without large pale hair-scars;
midrib rather slender above, flattish to moderately raised; nerves 11-12 pairs,
above flattish to raised, the submarginal arches not distinct; tertiary venation
forming a lax network, + distinct or not; petioles 8-15 x 2.5-3.5 mm, glabrescent;
leaf bud 15-22 x 3-4 mm, densely brown to rusty pubescent with hairs 0.5-1.0(-1.5)
mm long. Inflorescences behind the leaves, shaggy-pubescent with rusty hairs c. 0.5
mm, in CG: 3 or 4 times ramified, many-flowered, 10-21 x 10-16 cm, common
peduncle 15-40 mm long, the flowers in CO in loose clusters of 5-10; Q-
inflorescences not seen, in fruit c. 7-8 cm long; bracts broadly oval-ellipsoid, 2-7
mm long, finely pubescent, caducous; flowers 3-(or 4-)valved, perianth glabrous,
pedicel generally glabrous, at base not articulate or + articulate in only some
flowers (see notes). Male perianth globose or slightly depressed-globose, 1.5-2.0 x
2.0-2.3 mm, top and base (broadly) rounded, glabrous; pedicel 0.8-1.5(-2.0) mm
long; perianth at anthesis cleft to 2-73, not or only slightly collapsing on drying,
valves c. 0.2 mm thick. Androecium depressed-globose, above and beneath broad-
ly rounded, or sagged at base, 0.6-1.0 x 1.1-1-8 mm, in transverse section (sub-)
circular; anthers 10-13, almost completely sessile, c. 0.8-1.2 mm long, free apices
up to 0.1 mm, curved, concealing the + 3-radiate apical slit or cavity (0.2-1)0.3-0.5
mm deep; column broad, solid; androphore rather narrow, (0.1)0.2-0.4 mm long,
New account of Horsfieldia 4 35
completely or partly hidden by the anthers. Female perianth not seen (according to
persistent perianth under the fruit 3-valved, c. 3 mm long). Fruits 1-3 per in-
fructescence, ellipsoid, top and base rounded, c. 7.0-8.0 x 4.5-5.0 cm, glabrous,
drying dark brown, finely to coarsely tubercled, pericarp c. 15 mm thick; stalk c. 3
mm long, at base inarticulate; perianth persisting under the fruit.
Distribution. Borneo (Sabah, Mt. Kinabalu).
BORNEO. Sabah: Clemens 26971, 30536, 31579; (Mujin) San. 18843.
Ecology. Primary and secondary forest, ridge forest; on sandstone; 1000-1500 m
alt. Flowers November, December, February; fruits in November.
NOTES
1. Fieldnotes. Large tree. Bark grey, fissured; outer bark soft, c. 5 mm thick;
inner bark white, soft, c. 5 mm; cambium pale; sapwood white. Exudate from bark
stickly. Flowers golden. Ripe fruits orange, aril red.
2. The present species is close to H. montana according to the almost similar
male flowers, but differs considerably by its stouter habit; stouter twigs, larger
leaves, larger male inflorescences, and very much larger fruits with a thick pericarp;
in H. montana the fruits only measure 20-27 mm in length, and the perianth on
these do not persist.
3. The pedicels are generally inarticulate at the base; however, some of the
flowers of San. 18843 seem to have an articulation which may be an artifact caused
by drying the specimen.
4. Sinclair identified the specimens of the present new species as H. valida,
which was in his interpretation very variable and heterogenous, but which is
presently regarded as a species confined to Sumatra.
89. Horsfieldia montana Airy Shaw. Fig. 1C(89)
Horsfieldia montana Airy Shaw, Kew Bull. 1939, no. 10 (1940) 542 — Type: Sarawak, Dulit Range,
Richards (Native Collector) 2509 (K).
Tree (3-)7-24 m. Twigs terete or faintly angular, towards the apex 1.5-4(-7) mm
diam., grey brown to dark brown, early to rather late glabrescent, tomentum
short- to long-shaggy, composed of hairs 0.2-1.0 mm long; bark coarsely striate,
lenticels small and inconspicuous or absent, older bark with or without a tendency
to flake. Leaves in 2 rows, chartaceous to coriaceous, elliptic to elliptic-oblong,
broadest at or slightly above the middle, 4-14 x 2-6 cm, base (short-) attenuate, tip
rounded to (sub)acute; upper surface glabrous except the midrib which is late
glabrescent towards the base, drying olivaceous-brown to blackish, lower surface
glabrous except the very base and midrib (late) glabrescent, drying (chocolate-)
brown, not much contrasting with upper surface, without larger blackish marks
and without conspicuously large yellowish hair scars; midrib raised above; nerves
6-11 pairs, flattish to raised above, submarginal arches usually not distinct; tertiary
venation forming a lax network, faint or invisible on both surfaces; petioles 5-16 x
1.5-2.5 mm, rather late glabrescent; leaf bud 5-10 x 1.5-4 mm with hairs 0.2-1.0 mm
36 Gard. Bull. Sing. 39(1) (1986)
long. Inflorescences situated just behind the leaves, either rather sparsely pubes-
cent with hairs 0.2-0.4 mm, or densely shaggy-pubescent with hairs 0.5-1.0 mm
long, in ©’: 3 or 4 times ramified, moderately to many-flowered, 4-12(-16) x 3-10
cm, common peduncle 5-20 mm long, the flowers arranged in loose clusters of 3-10
each; 9-inflorescences few-flowered, 2-6 cm long; bracts ovate to ellipsoid, pubes-
cent, 2.5-6 mm long, caducous; flowers 3- or 4-valved, perianth glabrous, pedicel
glabrous, in C& inarticulate at base, in 9 more or less distinctly articulate (always?;
see notes). Male perianth globose to subglobose, (1.2-)1.4-2.0 mm diam., top and
base rounded, glabrous; pedicel 1-1.5 mm long; perianth at anthesis cleft to c. 4 to
nearly '2-way, not or only slightly collapsing on drying, valves (0.2-)0.3 mm thick.
Androecium globose or depressed globose, above rounded, at base rounded or +
truncate or sagged, (0.5-)0.6-1.1 x 0.8-1.1 mm, in transverse section subcircular;
anthers (8-)9-13 (thecae 18-26) almost completely sessile, 0.8-1.2 mm long, +
curved, free apices up to 0.1 mm; column broad, solid except for a shallow apical
cavity or slit 0.1-0.2 mm deep; androphore narrow, 0.3-0.5 mm long, completely
clasped:and hidden by the anthers thus making the androecium look sessile. Female
perianth ellipsoid-obovoid, c. 0.2 X 1.8 mm, glabrous, cleft at anthesis to c. 4,
valves 3, at sutures c. 0.3 mm thick, pedicel c. 1.5 mm long, glabrous, articulate at
base (see notes), ovary ellipsoid, c. 1.5 x 1.3 mm, glabrous, stigma minutely
2-lobed, 0.1-0.2 mm high. Fruits 2-9 per infructescence, ellipsoid, base rounded, +
contracted toward the stalk, top rounded to acutish, 2.0-2.7 x 1.3-1.7 cm, glab-
rous, drying brown, with or without few small lenticel-like tubercles, pericarp
1.5-2.0 mm thick; stalk 2-4 mm long, at base + articulate; perianth not persisting.
Distribution. Borneo: Sarawak, Brunei, Sabah.
BORNEO. Sarawak (2nd & 5th Div.): Richards (Native Collector) 2509; S: 32858, 33025, 33057,
3359] — Brunei: (Ashton) BRUN 1053 — Sabah (mainly Kinabalu & Pinosok Plateau): Clemens s.n. (9.
II. 1933), 29558, 32642, 32800, 50505, 50513; Native Coll. 821; Poore 12; SAN 28914, 29239, 32283,
36190, 38257, 38294, 38425, 28957, 49301, 76424, 76458, 76803, 76939; Sinclair, Kadim & Kapis 8987.
Ecology. Crest forest, kerangas forest on ridges, montane forest, moss forest,
Agathis forest; on black soil or sandy soil; (800-)1300-2000 m alt. Flowers and fruits
throughout the year.
NOTES
1. Fieldnotes. Low or medium-sized tree, without buttresses. Bark (slightly)
longitudinally, shallowly fissured, or sometimes falky, chocolate, red-brown, dark
brown or dark grey; inner bark pale yellowish, reddish, or brownish, with red
watery exudate or not; sapwood pale orange or whitish. Flowers yellow; staminal
disc orange; fruits yellow or red. Flowers sweet-scented.
2. Variation. All specimens from Sabah were collected on Kinabalu and its
vicinity, and they differ rather markedly from those from Sarawak (2nd and 5th
Div.) and from Brunei in the nature of the tomentum on the leaf bud, the apical
portion of the twigs, and the inflorescences. The tomentum on the specimens from
Brunei and Sarawak (including the type) is short, composed of compact dendroid
hairs only c. 0.2(-0.5) mm in length; the Kinabalu specimens have the tomentum
composed of hairs 0.5-1.0 mm long, sometimes with even longer emerging hairs,
and the tomentum of these specimens render the Kinabalu plants much ‘“‘rougher”’
in appearance. This character, however, seems incorrelated with any difference in
flower structure, and I have refrained from recognizing it as a different variety.
New account of Horsfieldia 4 37
3. Articulation of pedicels. Whether the pedicel at the base is articulated or not
has appeared to be a reliable character for many species, and this holds for both the
male and female flowers. In the present species the pedicels of male plants are
always inarticulate; in the only female flowering specimen seen, S$. 32858 (Sarawak,
5th Div.) the pedicels look distinctly articulate, as is the case in several collections
with immature fruit from Sabah. However, I cannot find other grounds to separate
these specimens.
4. Sinclair formerly regarded H. montana as a variety of the variable H. polys-
pherula, but in his posthumous publication (1975, p. 36, 42) the taxon was sub-
merged in his polymorphous H. glabra.
90. Horsfieldia punctata de Wilde, sp. nov. Fig. 1D(90)
Folia subtus dense praedita punctis nigrescentibus, ut in Horsfieldia punctatifolia Sinclair, differt foliis
minoribus eodem tempore coriaceis, 4-12 cm longis, apice obtusis, fructibus angustioribus, c. 2 cm
longis, pericarpio in sicco c. 1.5 mm crasso. Perianthia mascula 3-valvata, globosa, c. 1 mm diam.,
androecio globoso, ad sectionem transversam circulari, antheris c. 11, sessilibus. — Type: Malaya,
Burgess FRI 9014 (L; iso: K).
Tree 10-25 m. Twigs terete, not ridged, towards the apex 2.5-5(-8) mm diam.,
dark grey-brown, early to rather late glabrescent, tomentum rusty, with hairs
0.1-0.4 mm long; bark coarsely striate, lower down with a tendency to crack
longitudinally and to flake; lenticels + absent. Leaves in 2 rows, coriaceous,
elliptic-oblong to oblong, broadest at about the middle, 4.5-12.0 x 2.0-5.0 cm, base
(short-)attenuate, top rounded to subacute; upper surface drying olivaceous-
brown, lower surface rufous-brown, glabrous, but midrib remaining pubescent for
some time, with densely regularly spaced brown-black dots c. 0.1-0.4 mm diam.;
midrib flat or only slightly raised above; nerves 5-12 pairs, thin and flat or only
slightly raised above, late glabrescent, the marginal arches indistinct; tertiary
venation invisible on both surfaces; petioles 6-12 x 1.5-2.5 mm, late glabrescent;
leaf bud 7-13 x 2-3.5 mm, pubescence dense grey-brown to rusty, with hairs 0.1-0.4
mm. Inflorescences densely rusty pubescent, with hairs 0.2-0.3 mm, in GC: c. twice
ramified, not many-flowered, 2-3 x 0.5-1.0 cm, common peduncle c. 10 mm long;
Q inflorescences only once ramified, 2-3 cm long, glabrescent in fruit; bracts
elliptic-oblong, 1-3 mm, pubescent, caducous. Flowers 3-valved, in & in small
clusters of 3-5 each, the perianth glabrous, pedicels towards the base minutely
pubescent with hairs c. 0.1 m or less long, at base inarticulate. Submature male
perianths globose, c. 0.8 x 0.8-1.0 mm; pedicels c. 1.0 mm long, slender; perianth
cleft (sutures) to c. (%-)*4, valves c. 0.2 mm thick. Androecium depressed-globose
to globose, c. 0.4-0.5 x 0.5 mm, top and base broadly rounded, transverse section
+ circular; anthers c. 11 (c. 22 thecae, see notes), sessile, towards the top incurved
and concealing a shallow, narrow, apical cavity 0.2(-0.3) mm deep; androphore
narrow, up to 0.1 mm long. Female flowers not seen. Fruits 2-4 per infructescence,
ovoid, top rounded to narrowly rounded, base rounded, glabrous, 2.0-2.3 x 1.7-1.9
cm, drying dark brown, finely granulate, not tubercled; dry valves c. 1.5 mm thick;
stalk c. 2 mm long; perianth not persisting.
Distribution. Malaya (Cameron Highlands; Fraser’s Hill, Genting, Gunung Bun-
ga Bua).
MALAYA: FRI (Whitmore) 0306, 4539; (Burgess) 9014.
38 Gard. Bull. Sing. 39(1) (1986)
Ecology. Lower montane forest on granite, ridge forest; at c. 1000 m. alt.
Flowers in March, fruits in November.
NOTES
1. Fieldnotes. Bole straight, with good form, no buttresses. Bark deep-brown to
mid-brown, grid-cracked with rather chunky scales or finely fissured with ridges
firm, or bark thick and corky, finely longitudinally fissured; outer cut of slash bark
brown, inner bark bright red, layered, separated by blade line. Slash wood white to
fawn, speckled red; exudate red, blood-like. Fruits greenish-yellow, slightly
glaucous.
2. The available flowers are clearly immature, and hence the anthers are difficult
to count; possibly there are c. 11 thecae and in reality only 5 or 6 anthers.
3. Obviously closely related to H. glabra from S. Sumatra and Java, and to H.
montana, H. punctatifolia and H. subalpina. Horsfieldia montana (from Borneo) is
very similar in general habit as well as in the architecture of the male flowers and
their fruit-shape, but lacks the characteristic blackish punctation of the leaves; H.
punctatifolia, a species with a wider distribution in Sumatra, Malaya and Borneo
differs in its membranous leaves and larger fruits with a very thick-leathery peri-
carp; H. subalpina is also related, but that species lacks the punctation of the
leaves, has larger and less coriaceous leaves, and larger fruits. H. punctatifolia has
glabrous pedicels, in H. subalpina the pedicels are finely puberulous in the lower
part, similar as in our present species. Generally H. glabra has a much shorter
tomentum.
4. Specimens of this species had not yet been collected when Sinclair revised the
genus.
91. Horsfieldia costulata (Miq.) Warb. Fig. 1D(91)
Myristica costulata Miq., Ann. Mus. Bot. Ludg.,-Bat, 2, 1 (1865) 48 — Horsfieldia costulata (Miq.)
Warb., Mon. Myrist. (1897) 350 — Type: Celebes, de Vriese & Teijsmann s.n. (L).
H. pachythyrsa Warb., Mon. Myrist. (1897) 618: Koorders, Fl. N.O. Celebes in Med. Lands. Pl. Tuin 19
(1898) 570 (‘‘crassithyrsa’’) — M. pachythyrsa (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 86; 87
(““Myristica crassithyrsa’’) — H. minahassae auct. non (Warb.) Koord.: Koord., Fl. N.O. Celebes,
etc. (1898) 70, p.p. quoad Koorders 18158 — Syntype: Koorders 181568 (0', L. lecto), 18158 (L.),
181708 (2, L).
H. confertiflora Merr., Ph.J. Sc. C. Bot. 13, 5 (1918) 285 — Type: Ahern’s Coll. F.B. 3183 (PNH, n.v.;
iso: K; P; BO, NY; ve v2):
H. megacarpa Merr., Ph. J. Sc. C. Bot. 13, 5 (1918) 286 — Type: Ramos B.Sc. 16527 (PNH, n.v.; iso:
BM, & LP? BOSNY; SING.“US: #.7.).
H. villamilii Elmer ex Merr., En. Phil. Fl. Pl. 2 (1923) 182, nom. nud.
H. vulcanica Elmer ex Merr., En. Phil. Fl. Pl. 2 (1923) 182, nom. nud.
Tree 9-30 m Twigs terete or subterete, not ridged, towards the apex 2.5-5.0
(-10.0) mm diam., greyish to dark brown, early glabrescent, tomentum grey-brown
to light rusty, with hairs 0.1(-0.2) mm, lower down the bark finely to coarsely
New account of Horsfieldia 4 39
striate, not flaking, lenticels small, generally inconspicuous. Leaves in 2 rows,
membranous, elliptic-oblong to oblong-lanceolate, broadest at or somewhat above
the middle, 15-30 x 5-13 cm, base narrowly rounded to attenuate, top acute-
acuminate; upper surface drying olivaceous to dark brown, sometimes with whit-
ish marks as in H. irya; lower surface early glabrescent, without regularly scattered
larger, brown to blackish dots; midrib above flat or slightly raised, early glabres-
cent, also towards the base; nerves 14-21 pairs, above thin, flattish to usually
raised, marginal arches generally indistinct; tertiary venation forming a lax network
indistinct or invisible on both surfaces; petioles 7-14 x 2-4 mm; leaf bud slender,
8-14 x 2-2.5 mm, densely grey-brown to rusty pubescent with hairs 0.1(-0.2) mm.
Inflorescences situated behind the leaves, rather densely to sparsely pubescent with
hairs 0.1-0.2 mm, in CG: 3 or 4 times ramified, many-flowered, 6-14 x 5-13 cm,
common peduncle 10-30 mm long; @ inflorescences 2-6 cm long, shortly branched;
bracts broadly triangular to elliptic-oblong, 2-4(-5) mm long, densely short-
pubescent, caducous. Flowers 3(or 4)-valved, in & in rather dense clusters of 5-10
each, in 9 fewer, glabrous, pedicels glabrous, at base inarticulate. Male perianth
globose to + depressed-globose, glabrous, 1.5-1.8 xX 1.5-2.0 mm; pedicel rather
slender, short, 0.4-0.6(-0.7) mm long; perianth at anthesis cleft to c. /2-way, valves
c. 0.2 mm thick. Androecium depressed-globose or broadly ovoid or subglobose,
0.5-0.8 X 0.7-1.1 mm, circular in transverse section, anthers 7-10 (thecae 14-20),
completely sessile, free apices up to 0.1 mm, incurved over the rather narrow apical
cavity which is c. (0.1-) 0.2 mm deep; androphore rather stout, 0.2-0.4 mm long,
completely or partly hidden by the anthers. Female perianth subglobose, 2.3-2.5
mm diam., glabrous, cleft at anthesis to a depth of c. 4% to nearly '2-way, valves
0.3(-0.4) mm thick; pedicel 0.5-1.0 mm long; ovary ovoid, glabrous, 1.2-1.5 mm
diam., stigma minutely 2-lobed, 0.1-0.2 mm. Fruits 1-3 per infructescence, subglo-
bose to broadly ellipsoid or broadly obovoid, top and base broadly rounded,
glabrous, c. 3.5-6 x 3-4 cm, surface finely granulate, drying bright brown to
blackish brown, dry valves 8-10 mm thick; stalks 2-4 mm long; perianth not
persisting.
Distribution. Philippines, Celebes.
PHILIPPINES (Luzon, Pinay, Leyte, Mindanao, Basilan): BS 16527, 46059, 48080; Elmer 17045;
For. B. 3183, 11885, 18893; Jacobs 7604 (Spirit Coll. 5719); PNH 2685, 10995, 42247; Santos 4246; Vidal
1676, 3563; Wenzel 920.
CELEBES (Minahasa, North, Central): van Balgooy 3432, 3572; b.b. 17630, 17967, 28217; Koorders
181568, 181588; Meijer 9718; de Vogel 2602, 5523, 5626, 6217; de Vriese & Teijsmann s.n.
Ecology. Mixed rain forest, primary dipterocarp forest; recorded from alluvial
soil and volcanic soil, with Eucalyptus deglupta dominant; 250-1200 m alt. Flowers
and fruits throughout the year, but flowers mainly July to September.
Vernacular names. Kaya Ra (Ra=blood; C. Celebes), Kajura (C. Celebes).
NOTES
1. Fieldnotes. Tree with or without low buttresses, c. 30 x 10 cm. Bark fissured
or with longitudinal grooves, often peeling off. Heartwood reddish. Sap from bark
first clear, turning red to brown-red. Flowers yellow. Fruits yellow to red, on the
larger branches. Jacobs, referring to no. 7604, with the fruits looking as though
mature, had remarked: fruits not yet ripe, but aril orange, apparently dehiscent.
40 Gard. Bull. Sing. 39(1) (1986)
2. H. confertiflora (type from Rizal Prov., Luzon) seems a form with rather
chartaceous leaves.
3. At first I had intended to describe Jacobs 7604, from the Sierra Madre,
Luzon, as a separate variety. Its pickled fruits measure c. 7 X 5 cm, and have the
pericarp (in spirit) 14-16 mm thick. When dried, however, these fruits attained the
size of c. 6 X 4 cm, with the pericarp c. 10 mm thick, linking up with the only
slightly smaller fruits of the other material. Its mature-looking (solid) seeds are
ellipsoid, c. 4 cm long.
4. H. costulata, incl. the synonyms H. confertiflora and H. megacarpa, was
reduced by Sinclair to H. valida, sensu lato.
92. Horsfieldia subalpina Sinclair Fig. 1D(92)
Horsfieldia subalpina Sinclair, Gard. Bull. Sing. 16 (1958) 410; 28 (1975) 131. — Type: Malaya, Wray
467 (K; iso: L).
Tree 6-30 m. Twigs terete, not ridged, towards the apex 2.5-5(-12) mm diam.,
dark grey-brown, early glabrescent, tomentum greyish-brown with hairs c. 0.1 mm
long or less; bark finely to coarsely striate, lower down not flaking; lenticels usually
conspicuous. Leaves in 2 rows, membranous to chartaceous, elliptic-oblong to
oblong, broadest at or somewhat above the middle, 15-27 x 5-10 cm, base attenu-
ate, top acute to acute-acuminate, sometimes + bluntish; upper surface drying
olivaceous brown to dark brown, lower surface glabrous, without regularly scat-
tered larger blackish brown dots; midrib above flat or slightly raised; nerves
9-18(-20) pairs, thin and flat or slightly raised above, marginal arches not distinct;
tertiary venation forming a lax network not or slightly visible on both surfaces;
petioles 5-15 x 2-3 mm, glabrous; leaf bud slender, 12-20 x 2-3 mm, with dense
grey-brown to rusty tomentum of hairs c. 0.1 mm long or less. Inflorescences
situated behind the leaves, sparsely (subsp. kinabaluensis) to rather densely pubes-
cent with hairs c. 0.1 mm, in CG: rather stout, c. 3 (or 4) times ramified, many-
flowered, 5-14 x 3-10 cm, common peduncle (6-)10-30 mm long; 9 inflorescences 2
(or 3) times ramified, 2-7.0 x 1.5-4 cm, fewer-flowered than in C’; bracts ellipsoid
to oblong, acutish, densely short-pubescent, 2-5 mm long, caducous. Flowers 3- (or
4-)valved, in the © in loose clusters of 2-5 each, perianth glabrous, pedicels
glabrous (subsp. kinabaluensis) or thinly pubescent entirely or only in the lower
half (subsp. subalpina), at base inarticulate. Male perianths broad-ellipsoid or
subglobose, 1.6-2.3 mm long; pedicels 1.5-2 mm, slender; perianth at anthesis cleft
to c. 2-way, valves 0.2(-0.3) mm thick. Androecium globose or broadly ellipsoid,
1.0-1.5 mm long, transverse section circular; anthers 8-12, almost completely
sessile, free apices c. 0.1 mm, curving towards the top and concealing the rather
narrow, apical cavity, which is 0.3-0.5 mm deep, the column otherwise solid;
androphore narrow, 0).2-0.3 mm long, largely hidden by the anthers; see further
under the subspecies. Female perianth ellipsoid, 2.0-2.5 x 1.8-2.1 mm, glabrous
(subsp. kinabaluensis), cleft at anthesis to a depth of c. (3-)’2, valves c. 0.2(-0.3)
mm thick; ovary subglobose to broadly ellipsoid, 1.2-1.5 mm long, glabrous, stigma
minutely 2-lobed, c. 0.1 mm high; pedicel c. 1.5 mm long, at base inarticulate.
Fruits 2-6 per infructescence, subglobose to broadly ellipsoid to ellipsoid-oblong,
2.5-5 cm long; perianth not persisting; stalk 3-7 mm long; see further under the
subspecies.
New account of Horsfieldia 4 4]
Distribution. Two subspecies, one in mountainous Malaya, one in the Kinabalu
area in Sabah.
NOTE
When working on the group of species with H. subalpina and H. obscura,
there remained a number of fruiting specimens which I could not assign satisfactori-
ly to any of the known taxa. More information on their affinities can only be
obtained from such male flowering specimens which have vegetative characters
matching closely those of the fruiting material. Such taxa are probably closely
related to H. subalpina or H. obscura. They are enumerated and discussed from A
to H under H. obscura (p.44).
KEY TO THE SUBSPECIES
la. Pedicels of male flowers pubescent, at least in the lower half. Male perianth subglobose;
androecium subglobose, anthers 9-12. Fruits subglobose to broadly ellipsoid, 2.5-3 cm long
a. subsp. subalpina
ROE EEE EEE EEE HERE EEE EEE EEE EEE EEE EEE HEE HERE TEETH EEE EEE HEE HEHEHE HEHE EHH HEE THEE EEE E EE
b. Pedicels of male flowers glabrous. Male perianth (broadly) ellipsoid; androecium (broadly) ellip-
soid, anthers 8 or 9. Fruits ellipsoid, 3-5 cm long .....................cceeeeeeeee ees b. subsp. kinabaluensis
a. subsp. subalpina Fig. 1D(92)
Inflorescences rather densely pubescent. Male pedicels, at least in the lower half
thinly pubescent with hairs c. 0.1 mm. Male perianth subglobose to broadly
ellipsoid, 1.6-2.3 x 1.6-2.2 mm; androecium of similar shape, 1.0-1.5 x 1.0-1.2
mm; anthers 9-12. Female flowers not seen. Fruits subglobose to broadly ellipsoid,
2.5-3.0 xX 2.0-2.3 cm, top and base rounded, drying brown-blackish, without or
with a few tubercles; dry valves 3-4 mm thick.
Distribution. Malaya (Perak, Pahang, Selangor; Fraser’s Hill, Genting High-
lands).
MALAYA: Burkill & Holttum 8679; FRI 3884, 4539, 4565; 10972, 16161, 20485; Purseglove P. 4212;
SFN 23646; Md Shah MS 1096; Wray (jr) 467.
Ecology. Mountain forest, 800-1500 m alt. Flowers April, June; fruits Jan,
August to December.
Fieldnotes. Bark smooth or with shallow distant fissures; slash reddish, with red
sap; slash wood whitish. Flowers yellow, fruits greenish yellow and glaucous, or
yellow; aril orange, seed white. The fruits of Shah MS 1096 were recorded when
fresh as measuring 4.5-5 cm long, dry as 3.3-4 cm long; on the Leiden specimen the
dry mature fruits reach only c. 3 cm long.
b. subsp. kinabaluensis de Wilde, subsp. nov.
Gemma tomento e pilis 0.1 mm longis vel minus composito praedita. Differt a subsp. subalpina
pedicellis florum masculorum glabris, perianthio masculo atque androecio late ellipsoideis, antheris 8
vel 9, fructibus in sicco ellipsoideis, 3-5 cm longis. — Type: Sabah, Clemens 33136 (L; iso: BM, K;
A, B, BO, M, NY, SING, UC, n.v.)
Inflorescences very sparsely pubescent. Male pedicels glabrous. Male perianth
(broadly) ellipsoid, 1.7-2.0 x 1.5-1.8 mm; androecium broadly ellipsoid, 1.1-1.2 x
42 Gard. Bull. Sing. 39(1) (1986)
0.8-1.0 mm; anthers 8 or 9. Female flowers as described under the species. Fruits
ellipsoid to broadly ellipsoid, 3.0-5.1 x 1.7-2.5 cm, top + narrowly rounded, base
rounded, drying bright brown to blackish brown, without tubercles; dry valves 4-5
mm thick.
Distribution. Borneo: Sabah (Mt. Kinabalu and vicinity), one doubtful collection
from E. Sarawak, see notes.
BORNEO. E. Sarawak (Kalabit Highland, 4th Div.): Chai S 35461, doubtful — Sabah (mainly Mt.
Kinabalu and Pinosok Plateau) Clemens 26863, 28305, 28730, 32204, 33136, 50721; Kokawa & Hotta
5477; M.E.D. Poore 0906; Chew & Corner RSNB 4116, 4209, 4530, 7000; San 26793, 28962, 38081,
49691; SFN (Carr) 27351, 27450.
Ecology. Mountain forest, montane oak forest; clayish soil; 1400-2000 m alt.
Flowers and fruits throughout the year.
NOTES
I. Fteldnotes. No buttresses. Bark slightly fissured, reddish brown; inner bark
fibrous, whitish turning brown, or soft, and then yellowish; cambium pale yellow;
wood white to yellowish, medium hard, heartwood not differentiated. Flowers
bright yellow. Fruits yellow-red.
2. Chai § 35461 from Kalabit Highland, c. 1250 m alt., Baram Dist., 4th Div.,
Sarawak, with immature fruit, possibly belongs here. It deviates by having narrow
leaves and small fruits which are albeit immature. The specimens also might belong
to H. xanthina.
3. Sinclair determined specimens of the present subspecies as H. glabra and H.
valida.
93. Horsfieldia obscura de Wilde, sp. nov. Fig. 1D(93)
Tomentum gemmae e pilis 0.1-0.2 mm longis compositum. Folia subtus non praedita punctis nigrescenti-
bus. Androecium ad sectionem transversam circulare. Affinis Horsfieldia subalpina Sinclair, differt in
anthesi perianthio masculo usque ad 4 fisso, atque fructibus in sicco majoribus, 5-5.5 cm longis. —
Type: E. Kalimantan, Kostermans 13773 (L; iso: K, P; BO, A, SING, PNH. NY, n.v.).
Tree c. 20 m. Twigs terete, not ridged, towards the apex 2-3(-4) mm diam., dark
grey-brown, early glabrescent, tomentum greyish-brown to rusty, with hairs 0.1-0.2
mm long; bark finely striate, lower down not flaking; lenticels moderately large,
not much contrasting. Leaves in 2 rows, membranous, elliptic-oblong to oblong,
broadest at or slightly above the middle, 10-15 x 4-7 cm, base short-attenuate, top
shortly acute-acuminate; upper surface drying dark brown, lower surface pale
brown to bright brown, glabrous, without scattered larger blackish dots; midrib
above flat to slightly raised; nerves 10-13 pairs, very thin and flat above, marginal
arches invisible; tertiary venation forming a lax network, very faint or invisible on
both surfaces; petioles 11-15 x 1.5-2.5 mm, glabrous; leaf bud slender, 7-10 x 1.5-2
mm, densely grey-brown to rusty pubescent with hairs c. 0.1-0.2 mm. Infloresc-
ences situated behind the leaves, + sparsely minutely pubescent with hairs c. 0.1
mm, in &: 3(or 4) times ramified, rather many-flowered, 7-10 x 5-8 cm, common
peduncle (2-)6-12 mm long; @ inflorescences c. 3.5 cm long (very stout, as seen in
New account of Horsfieldia 4 43
the infructescences of S. 36305, (see notes); bracts caducous, not seen. Flowers 3-(or
4-) valved, in CO in clusters of c. 3-5, perianths glabrous, pedicels glabrous or with
some minute hairs towards the base, not or sometimes faintly articulate. Male
perianth subglobose, 2.0-2.2 x 2.0-2.3 mm; pedicels (1-)1.5-2 mm long, slender;
perianth at anthesis cleft 73-*/4(-%), valves 0.2-0.3 mm thick. Androecium subglo-
bose to depressed globose-ovate, at the top narrowly rounded, base broadly
rounded, c. 1.0 < 1.0-1.2 mm, transverse section (sub)circular; anthers c. 8, almost
completely sessile, the free apices sometimes + sterile, up to 0.1(-0.2) mm long,
curving over and concealing the narrow, shallow, apical cavity, cavity c. 0.1-0.2 mm
deep, the rest of the column solid; androphore narrow, up to 0.2 mm long, hidden
by the anthers. Female flowers not seen. Fruits (of S. 36305, see note 3) 2 or 3 ona
short, stout infructescence c. 3.5 cm long; fruits broadly ellipsoid, 5-5.5 x 3.5-4.0
cm, top and base rounded, glabrous, drying dark brown, with a few, small tuber-
cles, dry valves 7-10 mm thick, perianth not persisting.
Distribution. Borneo: E. Kalimantan, possibly Sarawak, see note 3.
BORNEO. E. Kalimantan: Kostermans 6044, 13773 — Sarawak (7th Div., Kapit): S 36305
(doubtful).
Ecology. Ridge forest on limestone, coral limestone, yellow sandy soil in lowland
dipterocarp forest; 150-730 m. alt. Flowers (E. Kalimantan) in August & Novem-
ber; fruits (Sarawak) in May.
NOTES
1. Fieldnotes. Trunk irregular. Bark red-brown, rough, 5 mm thick, peeling off
irregularly in strips. Living bark 10 mm, red to brown-red; sap red. Wood reddish
to brown-red. Flowers yellow to dark yellow, smelling of Peru-balsam. Fruits bright
orange, seed (aril) red.
2. The two flowering specimens on which the present species is based, Koster-
mans 6044 and 13772, were identified by Sinclair as Horsfieldia, probably glabra.
and H. glabra respectively.
3. The specimen S. 36305 from Kapit, 7th Div. Sarawak is in fruit and agrees
vegetatively with the two flowering specimens from E. Kalimantan. However, it
being the only one in fruit, collected rather far from the limestone site in E.
Kalimantan, provenance of the two male specimens, I am not completely sure
whether the two lots are conspecific.
4. Horsfieldia obscura, with male flowers, keys out beside H. subalpina, which is
accepted for Borneo as the subspecies kinabaluensis, restricted to the montane
forest in the Kinabalu area. That differs from our present species by the more
elongate male perianth, cleft at anthesis to only nearly /2-way, the short-ellipsoid
androecium, the more rigid leaves, and the smaller fruits with the pericarp less
thick. The present species appears to be closest to the typical H. subalpina, from
Malaya, which differs in general habit (larger leaves, more distinct lenticels on
twigs, larger inflorescences), male flowers cleft to c. /2-way deep, and by smaller
fruits.
5. Some deviating specimens which are in fruit, all from Borneo, will most likely
key out beside H. subalpina and H. obscura.
-- Gard. Bull. Sing. 39(1) (1986)
When I was working on the group of species included by Sinclair in his large
concept of H. glabra and resembling species, there remained some ten sheets of
fruiting material which obviously or likely belong to this alliance, but which cannot
be satisfactorily matched with any of the species accepted by me. These specimens
are enumerated and discussed below (in 8 groups, A to H), because I presume that
if male flowers were available, they would key out on or near H. obscura or H.
subalpina. They are not identical with these species, however, because of some
features of general habit (twigs and leaves), and because of the characters of the
leaves, fruits, and possibly flowers. As a matter of fact, the listed entities below also
differ clearly one from the other. They may represent distinct species, but male
flowers are needed to affirm this.
A. Kostermans 7414, 7461. Both were collected in July 1952 on the peak of
Balikpapan (Besikan Balu), E. Kutei, E. Kalimantan, at 700 and 900 m in a mossy
forest on sandstone. Vegetatively they come very close to H. subalpina, but they
strongly deviate by the enormous fruits measuring c. 7-9 X 4.5-5.5 cm when dry,
with the pericarp c. 25 mm thick. The bark on the twigs is rather pale and rather
smooth, with only small inconspicuous tuberculate lenticels; the nerves are flat
above. They were annotated as trees of 12-15 m, bark light brown to blackish,
superficially fissured, living bark brown-red, 5-6 mm, wood white with red streaks,
fruits orange-brown, aril yellow to orange. These specimens were in 1959 identified
by Sinclair as H. subalpina and in 1975 as H. valida.
B. Kostermans 4355. This was collected in Nov. 1950 in E. Borneo, Balikpapan
Bay, Muan region near Sg. Riko, at 20 m. alt. on a low ridge with sandy soil with
lime. In 1954, in the herbarium at Leiden Sinclair identified it as H. subalpina; in
1975 as H. valida. It is annotated as a tree, 28 m, with buttresses 6 m high, c. 50 cm
over the ground and 30 cm thick; bark rough, fissured, dark brown, containing red
sap. Fruit orange. A rather rare tree. The specimens have large, broad, brittle
leaves, with the midrib above being rather broad at the base, and therefore, it was
perhaps the late Mr. Hildebrand who wrote the identification as probably H.
brachiata var. laticostata, i.e., the present H. laticostata. However, the large fruits
of c. 5.5 xX 4.0 cm, with thick woody pericarp do not agree. The tomentum of the
leaf bud is fairly coarse, with hairs c. 0.2 mm long, somewhat reminiscent of that of
H. valida, a species presently regarded as restricted to Sumatra.
C. Ashton BRUN 766 and Murthy & Ashton §. 23348 are probably identical.
The first was collected in Brunei in Nov. 1957 in a forest on alluvial soil at the top of
a riverbank at c. 150 m altitude and the second, in Sarawak, 4th Div., March 1965,
rather inland on the banks of Sg. Baloi, on clay-sand soil. Both have large fruits,
long-ellipsoid, c. 6.5 cm long, blackish brown, with a rather woody pericarp 6-8 mm
thick (dried), when fresh orange-red and orange-yellow respectively. The leaves
dry bright dark brown, the nerves are rather raised above. The twigs of BRUN 766
dry rather pale and lenticels are inconspicuous. Both specimens have rather large
leaves, those of §. 23348 measuring up to 34 x 12 cm. They somewhat superficially
resemble H. fragillima.
D. Tong and Ilias § 32763 from Sarawak, Sth Division (Ulu Sg. Pandarasan, —
March 1973), a kerengas forest at c. 800 m on sandy soil, has large broadly ellipsoid
to subglobose fruits c. 5.5 X 4.5 cm, with apparent thick pericarp, orange when
fresh. The fruit resembles somewhat but is smaller than that of species A, B, and C.
Vegetatively the specimen agrees best with A in that the leaves are of about similar
New account of Horsfieldia 4 45
size and texture, and the nerves flattish above, and in the bark of the twigs,
relatively pale and lenticels inconspicuous. A, however, has very much larger fruits
(see above). In Murut language it is called ‘“‘Bidarak”’.
E. Sinclair (& Kadim & Kapis) 9278 at Leiden is a leafy twig with the female
inflorescences in an envelope. The field label states that fruits are large oblong,
orange outside, 9 X 6 cm, pink inside, the aril orange, the seed 5.5 X 3 cm. It was
collected in June 1957, Ranau Dist., Sabah, altitude unrecorded. Sinclair deter-
mined it as H. punctatifolia but its leaves are devoid of dots, and it does not belong
there. These are of moderate size, up to 16 X 5.5 cm, with nerves on the upper
surface flat; the bark on the twigs is rather pale.
F. §. 28083 (Ilias Pai’e). I feel strongly that this represents an undescribed
species. It was collected in Sarawak, Serian Dist., Ist. Div., South of Kuching,
Sept. 1968, in Bukit Selabor, at c. 800 ft. alt., near a stream below Bukit Selabor; a
tree. c. 15 m tall, on yellow clay soil, the fruits reddish-green. This locality is in an
area with several other locally endemic Horsfieldia species. The L-specimen consists
of a twig with several large thinly chartaceous leaves c. 33 X 14 cm, which dried
light olivaceous-brown; nerves 16-18 pairs, raised above. The leaf bud is densely
pubescent with hairs c. 0.2 mm long, the twig is conspicuously lenticellate and there
is a ramified infructescence of c. 8 cm long with two large fruits c. 6.5 X 4.5 cm,
drying dark brown, with some coarse tubercles, the dry pericarp is c. 10 mm thick.
Possibly the alliance of the specimen is with species like H. fragillima or, less likely,
with H. grandis or H. reticulata (which have the leaves pubescent beneath), rather
than with the H. subalpina and the H. obscura-group.
G. Chew Wee-lek CWL 687 from Sarawak, Kuching Dist., from limestone at c.
500 ft. has mature fruits (July 1963) which are ellipsoid, c. 3 x 2 cm, the pericarp c.
2.5 mm thick, finely pale warted-punctate. The leaves are not punctate-dotted
beneath. In habit and in fruit, this specimen resembles H. subalpina which is a
montane species in Malaya and Borneo and I have accepted the Bornean material
as subsp. kinabaluensis which has larger fruits and which is restricted to Mt.
Kinabalu at 1400-1800 m.
H. SAN 41826 (Mikil) trom Sabah, Tambunan Dist., 6000 ft, and S$ 33061 (Tong
& Jugah) from Sarawak, Lawas, 5th Div., at 4500 ft. are probably conspecific. Both
are in fruit and do not agree with any known species or with the specimens A to G.
The twigs have small but rather conspicuous lenticels, the tomentum of the leaf bud
is composed of hairs c. 0.1 mm long, the leaves are of moderate size, up to c. 17 cm
long, + membranous, with the nerves distinctly raised above; the fruits are ellip-
soid, 3.5-4 cm long, with the pericarp drying hard, 3-4 mm thick. The specimens are
reminiscent of H. xanthina, but the nerves in the latter are not raised above, and
of H. androphora and H. montana, but these have the tomentum of the leaf bud
composed of remarkably longer hairs, i.e., c. 0.5 mm long, a character which has
proved to be constant and valuable in the taxonomy of Horsfieldia. Possibly they
belong to an undescribed species close to H. androphora or H. montana.
94. Horsfieldia xanthina Airy Shaw Fig. 1D(94)
Horsfieldia xanthina Airy Shaw, Kew Bull. 1939, no. 10 (1940) 541 (441) — Type: Mt. Dulit, Sarawak,
Richards 1927 (K; iso: A, SING, n.v.).
Tree 10-30 m. Twigs terete or faintly angular at apex, 2.5-6(-16) mm diam.,
46 Gard. Bull. Sing. 39(1) (1986)
drying dark brown, early glabrescent tomentum of hairs c. 0.1 mm, lower down
bark coarsely striate and with a tendency to flake, lenticels conspicuous or not.
Leaves in 2 rows, coriaceous, elliptic to oblong-lanceolate, broadest at about the
middle, 8-35 x 3.5-13 cm, base attenuate to short-rounded, top subacute to
acute-acuminate; upper surface drying olivaceous-brown to (dark) brown, usually
with rather distinct hair-scars, lower surface drying usually with a reddish-brown
tinge, without blackish dots, early glabrescent; midrib above slightly raised, glab-
rous; nerves 8-20 pairs, above slender, flat or slightly raised, glabrous, the lateral
arches very indistinct above; tertiary venation forming a lax network, indistinct or
invisible on both surfaces; petioles 7-12 x 2.5-3 mm, early glabrescent; leaf bud
8-13 x 2-3.5 mm, covered by hairs 0.1(-0.2) mm long. Inflorescences behind the
leaves, rather densely pubescent by hairs c. 0.1 mm, in GC’: rather short and robust,
(2-)4-20 cm long, (2 or) 3 times ramified, not many-flowered, common peduncle
5-8 or 15-20 mm (subsp. macrophylla) long, the flowers in loose clusters of 3-5(-8),
reflexed or not; Q inflorescences (as seen in infructescences with young fruits) 1-1.5
or 5-7 (subsp. macrophylla) cm long, few-flowered; bracts not seen, caducous;
tlowers 3- or 4-valved, perianth glabrous, pedicels glabrescent or with a little
tomentum of hairs, c. 0.1 mm towards the base, at base inarticulate. Male perianth
broadly ellipsoid to ovoid-subglobose, 2.5-2.8 mm long, top rounded, base shortly
rounded and somewhat tapering into the pedicel; pedicel thickish, somewhat
tapering or not, straight or + curved (flowers reflexed), 1-2.5 mm long; perianth at
anthesis cleft to c. 2 (-%), slightly wrinkled but not collapsing on drying, valves
0.4-0.8(-1.0) mm thick. Androecium somewhat laterally flattened, + broadly obo-
void in outline, 1.0-1.3 mm long; anthers 5-8, erect, 0.9-1.1 mm long, largely sessile
with free apices erect, 0.1-0.3 mm; column rather broad, solid, at apex shallowly
hollowed (0.1-0.3 mm); androphore + tapering, rather broad, (0.1-)0.2-0.3 mm
long. Female inflorescences and flowers known only in subsp. macrophylla; ovary
glabrous. Fruits 2-6 per infructescence, when immature (subsp. xanthina) ellipsoid-
obovoid or broadly ovoid, glabrous; stalk stout, c. 3 mm long; perianth not
persisting.
Distribution. Borneo: Sarawak, Sabah; two subspecies, both montane.
KEY TO THE SUBSPECIES
la. Twigs moderate, towards the apex 2.5-4.5(-8) mm diam. Leaf blades 8-18 x 3.5-7 cm. Male
inflorescences 2-5 cm long, the flowers often + reflexed; anthers 5S or6. ........... a. subsp. xanthina
b. Twigs stout, towards apex 3.5-6(-16) mm diam. Leaf blades 22-35 x 7-13 cm. Male inflorescences
10-20 em; flowers efect, anthers: OF B.A J. Nanos hireren Pek tc Ath ree ene b. subsp. macrophylla
a. subsp. xanthina Fig. 1D(94)
Tree 10-17 m. Twigs towards apex 2.5-4.5(-8) mm diam., bark with a tendency of
flaking. Leaf blades 8-18 x 3.5-7 cm. Male inflorescences (2-)4-5 x 2-3.5 cm, the
rhachis at base 2-3 mm diam.; flowers often + reflexed. Male perianth subglobose
or broadly ellipsoid, c. 2.5 X 2.2-2.5 mm, cleft at anthesis to c. 2-74; pedicels 2-2.5
mm long. Androecium slightly laterally compressed, 1.0-1.2 x 0.8-1.0 x 0.4-0.5
mm, anhers 5 or 6. Female flowers not seen. Fruits immature, ellipsoid-obvoid, c.
1.4 xX 1.2 cm.
New account of Horsfieldia 4 47
Distribution. Sarawak.
BORNEO. Sarawak (Mt. Dulit, Baram Dist.) Richards 1927; S 19396, 30821, 34876.
Ecology. Kerangas forest, heath forest, submontane forest; on sandy soils,
yellow sandy soil, sandstone, or “‘on higher flanks of limestone mountain among
huge limestone boulders with vegetation and organic layer entwined between
boulders”; 800-1150m. Flowers in August, September; young fruits in September.
Vernacular names. Kumpang; Getah merah (Sarawak); Buah itek (Kenyah,
Sarawak).
NOTES
1. Fieldnotes. Flowers yellow. Fruit glaucous green or green and shiny.
2. Male flowers known only from the type, and § 34876. A third specimen, $
30821, bears immature fruits. A fourth, S 19396 is annotated as having fruits, but I
have not seen these in the L-duplicate; this latter specimen slightly deviates by the
somewhat longer hairs on the leaf buds, being c. 0.2 mm long rather than c. 0.1 mm
or less as in the other specimens.
3. H. xanthina was included in H. glabra by Sinclair (1.c. p. 36, 41, 48). I accept it
aS a separate species, characterized by the flaky bark of the twigs, the coriaceous
and often + reddish-brown tinged leaves, and the coriaceous rather large male
flowers with a typical androecium of only 5 or 6 anthers and a rather marked
androphore. Possibly the species is confined to Kerangas at rather high altitudes.
I agree with Airy Shaw that the species is close to H. majuscula, from Malaya and
Sumatra, which differs by the non-flaking bark of twigs, membranous leaves, 7-9
anthers, the pedicel articulate at base, and possibly by larger fruits.
b. subsp. macrophylla de Wilde, subsp. nov.
Perianthium masculinum crasse coriaceum; androecium quam latum longius. A subspec. xanthina
ramulis robustioribus apicem versus 3.5-6(-16) mm diam., foliis maioribus 22-35 cm longis, 7-13 cm latis,
antherisque 7 vel 8. — Type: Mt. Kinabalu, J. & M.S. Clemens 50050 (K; iso BM, L).
Tree 25-30 m. Twigs towards the apex 3.5-6(-16) mm diam.; bark not flaking.
Leaf blades 25-35 x 7-13 cm. Male inflorescences 10-20 x 7-11 cm, the rachis at
base 3.5-4.5(-5.0) mm diam.; flowers erect. Male perianth broadly ellipsoid, 2.6-2.8
x 2.5-2.7 mm; valves 3, splitting the bud to nearly ’2-way; pedicel 1-1.5 mm long.
Androecium c. 1.3 X 0.8 mm, subtriquetrous in transverse section; anthers 7 or 8.
Female inflorescences 5-8 cm long, once or twice ramified. Female perianth ovoid-
ellipsoid, 4.0-4.5 x 3.0-3.5 mm, valves 3, splitting the bud to nearly '/2-way, valves
c. 0.8(-1.0) mm thick. Pedicel 1.5-2.0 mm, glabrous or with some minute hairs
towards the base. Ovary ovoid, c. 2.2 x 2.0 mm, glabrous, stigma broadly 2-lipped,
c. 0.3 X 1.0 mm. Fruits immature, broadly ovoid, c. 1 cm long.
Distribution. E. Sarawak, Sabah (Mt. Kinabalu).
BORNEO. Sarawak (East; Baram Dist., Kapit Dist.): Anderson & Ilias bin Paie S 28523; Chai S
35442 — Sabah (Mt. Kinabalu): J. & M.S. Clemens 50050.
48 Gard. Bull. Sing. 39(1) (1986)
Ecology. Mountain forest, ridge forest, at 1100-1300 m; on igneous derived
(andesitic) soils. Flowers July and November; young fruits in November.
Vernacular names. Kumpang lusoh, Kumpang parawan (Baram Dist.).
NOTES
1. Fieldnotes. Poorly developed low buttresses, once recorded; bark brown and
grey, fissures boat-shaped; exudate light red, watery. Flowers yellow or orange;
ovary pale purple. Young fruits green.
2. The type, with male flowers, was determined by Sinclair as H. valida, a
species presently regarded as restricted to Sumatra; the other specimens of the
present new subspecies were not yet collected in Sinclair’s time.
3. Fruit of the iso-type, in BM, is annotated as being in the fruit collection; I
have not seen them and doubt that they belong to the same species because the
mounted specimen is male.
95. Horsfieldia majuscula (King) Warb. Fig. 1D(95)
Myristica majuscula King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 310, pl. 143 — Horsfieldia majuscula
(King) Warb., Mon. Myrist. (1897) 315; Gamble, Mat. Fl. Mal. Penins. 5, 23 (1912) 215; Ridley, FL.
Mal. Pen. 3 (1924) 57 — Lectotype: Malaya, King’s Coll. 5039 (BM, L; see notes).
Horsfieldia bartlettii Merr., New Sumatran Plants IV, Papers Mich. Acad. Sc. Arts & Letters 24, 1 for
1938 (1939) 71 — Type: Sumatra, Rahmat si Boeea 8772 (A, n.v.; iso:L).
Tree 6-25 m. Twigs terete, towards the top 2.5-5(-8) mm diam., grey-brown,
early glabrescent, tomentum greyish brown, of hairs c. 0.1(-0.2) mm; bark lower
down rather coarsely longitudinally striate, with smallish, sometimes rather incon-
spicuous, lenticels, older bark not flaking. Leaves in 2 rows, membranous to thinly
chartaceous, elliptic-oblong to oblong, broadest usually at the middle, 15-27 x
5.5-9.5 cm, base attenuate, top acute-acuminate; upper surface glabrous, drying
dull olivaceous brown, the midrib glabrous, lower surface drying brown, not very
conspicuously contrasting in colour of the upper, without blackish larger dots, early
glabrescent including midrib; midrib above slender, glabrous, raised; nerves 11-19
pairs, raised above; tertiary venation forming a lax network generally indistinct or
invisible on both surfaces; petioles 10-20 x 2-2.5 mm, early glabrescent; leaf bud
slender, 10-15 x 2-3 mm, pubescent with grey brown hairs c. 0.1(-0.2) mm.
Inflorescences not very densely pubescent with stellate hairs c. 0.1 mm long, in C’:
2 or 3 times ramified, not very many-flowered, c. 8 X 4-5 cm, common peduncle
10-15 mm, the flowers in clusters of 2-5; Q-inflorescences 2-5 cm long; rather
few-flowered, twice ramified; bracts ovate-oblong, c. 4 x 2 mm, densely pubescent
with hairs c. 0.2 mm, caducous; flowers 3-(or 4-)valved, perianth glabrous, pedicel
glabrous or sparsely pubescent, immature ones with hairs c. 0.1 mm, at base
articulate. Male perianth broadly ellipsoid or obovoid, (2.0-)2.2-3 x (1.8-)2.2-2.5
mm, top rounded, base short-attenuate, pedicel 2-3 mm long, slightly tapering;
perianth at anthesis cleft to 4 to nearly /2-way, not or slightly collapsing on drying,
valves thick, c. 0.3 mm, towards the base to c. 0.8(-1.0; mm thick. Androecium
(incl. the 0.2-0.5-mm long androphore) ellipsoid to ellipsoid-oblong, (1.4-) 1.7-2.0
< 0.8-1.0 mm, in transverse section subtriangular to subelliptic; anthers 7-9 (i.e., 14-18
thecae), slightly curved, (1.2) 1.3-1.8 mm long, free at apex for c. 3-/%, 1.e., c.
New account of Horsfieldia 4 49
(0.2-)0.3-0.5 mm, androphore rather broad, shortly tapering, 0.2-0.5 mm long.
Female perianth broadly obovoid, c. 2.5 X 2.5 mm, glabrous, cleft at anthesis to c.
3, valves at base c. 0.6-0.8 mm thick, pedicels 2-3 mm long, glabrous, articulated,
ovary ellipsoid, 1.8-2.0 x 1.3-1.5 mm, glabrous, stigma 2-lipped, c. 0.3 mm high.
Fruits 1-5 per infructescence, ellipsoid, top and base rounded, 4.3-6.5 x 3.0-4.5 cm,
glabrous, drying rusty-brown, finely granulate but not or inconspicuously warted or
lenticellate, pericarp 4-11 mm thick; stalk 2-7 mm long; perianth not persisting.
Distribution. Malaya (Perak, Pahang, Kelantan), Sumatra (W. & E. Coast,
Palembang).
MALAYA: FRI 2885, 4506, 4767, 5656, 10835, 16511; Kadim & Noor KN. 610; KEP 104863; King’s
Coll. 5059; Scortechini 837.
SUMATRA. West Coast: Beccari 791; Meijer 6776 — East Coast: Lérzing 15557; Rahmat si Boeea
8772, 9331 — Palembang: van Steenis 3384.
Ecology. Dry-land forest and freshwater swamp-forest of lowland and montane
areas; rocky stream banks, river valleys; up to 1000 m alt. Flowers and fruits
throughout the year.
Vernacular name. Kajoe andorodong (E. Coast Sumatra, Asahan).
NOTES
1. Fieldnotes. Trees without buttresses; bark distantly shallowly fissured or
forming shallow, rectangular flakes. Slash wood whitish, red-flecked. Fruits (Mezer
6776, W. Sumatra) up to 5 per infructescence, when fresh up to 7 X 6 cm, pericarp
c. 1.5 cm thick. Fruits yellow to bright red, aril orange. Flowers yellow.
2. H. majuscula has formerly been mixed up with what is in my present treat-
ment H. polyspherula var. sumatrana (in Sinclair’s as H. brachiata var. sumatrana)
and the present new variety H. polyspherula var. maxima. Horsfieldia polyspherula
var. sumatrana differs by the usually coarser hairs on the leaf bud, more contrasting
colour of the two surfaces of dry leaves, smaller globose male perianths, an
essentially different androecium, and smaller fruits, up to 35 mm long. The var.
maxima, only known from Borneo, differs by the same characters, but has fruits of
similar size as H. majuscula, with the pericarp up to c. 15 mm thick.
3. Lectotypification. Sinclair (1975, p. 16-18) placed H. majuscula entirely (1.e.,
including the whole of King’s syntype) in the synonymy of his H. brachiata var.
sumatrana, a taxon which in this treatment is accepted in a much more restricted
sense as a variety of H. polyspherula. However, Sinclair did comment on the
heterogeneity of King’s syntypes.
King (p. 311) cites 7 syntype-specimens. These certaintly belong to more than
one species. Although King placed his Myristica majuscula in the section /rya on
the characters of male flowers, and although within this section it is keyed out
mainly on characters of the male flowers and the male inflorescences, I have chosen
King’s Collector 5059 (in L, iso K), a fruiting specimen, as the lectotype for the
following reasons.
50 Gard. Bull. Sing. 39(1) (1986)
My analysis of King’s syntypes together with some information from the rest of
the protologue is:
King’s Coll. 5059: fine fruiting collection, seen in K and L; lectotype; identical
with the present circumscription of H. majuscula.
King’s Coll. 6004: male flowering, seen in BM and L; this is a somewhat
aberrant specimen, because of its relatively large flowers, to be treated under H.
polyspherula var. sumatrana, see there.
King’s Coll. 7965: in submature fruit, seen in BM, K;; this is possibly the present
H. majuscula.
King’s Coll. 10413: submature fruit, seen in K, P. This is possibly H. polyspher-
ula var. sumatrana.
Wray 2218, 2705: specimens in CAL, SING, not seen; according to Sinclair both
collections link up with H. polyspherula var. sumatrana in the restricted sense.
(?) Hullet (590), from Singapore: fruiting, in herb. K; the fruits are small,
identical with my present H. polyspherula var. sumatrana.
However, the male plant figured on King’s plate 143 apparently belongs to true H.
majuscula as presently accepted by me, according to the smallish few-flowered
male inflorescence, and the longer-than-broad shape of the perianth and
androecium. Possibly it is drawn from Scortechini 837, a male collection, which was
certainly in King’s hands, but which is not cited among the syntype specimens of
Horsfieldia majuscula. Thus, most likely, this collection served for the drawing,
whereas King’s description of the male flowers (perianth globose with rather
depressed androecium) apparently was taken from King’s collector 6004, a some-
what abberrant specimen presently treated in the notes under H. polyspherula var.
sumatrana.
From the above it will be obvious that the present lectotypification prevents the
proposal of a new name, and the name H. majuscula is retained.
4. The specimen van Steenis 3384 (Palembang Province in Sumatra, near Lake
Ranau at c. 600 m) has immature male flowers; the perianth is slightly less
coriaceous and only c. 0.4 mm thick, the androphore short, only 0.1-0.2 mm, and
the pedicels are sparsely pubescent towards the base. I suppose that these slight
differences, if compared with the other specimens, can be attributed to the juvenile
stage of the flowers.
96. Horsfieldia coriacea de Wilde, sp. nov. Fig. 1D(96)
Habitu atque fructibus Horsfieldiae costulatae (Miq.) Warb. similis, differt floribus masculis perianthio
crasse coriaceo, androecio elongato, antheris 5 vel 6. — Type: b.b. Cel. III-27 (L).
Tree 8-25 m. Twigs terete, towards the apex 2.5-4(-10) mm diam., dark grey-
brown, early glabrescent, tomentum greyish brown, of hairs c. 0.1 mm; bark lower
down finely striate, older bark not flaking, lenticels conspicuous or not. Leaves in 2
New account of Horsfieldia 4 51
rows, membranous to thinly chartaceous, elliptic-oblong to oblong, broadest at or
slightly above the middle, 14-27 x 5-10 cm, base attenuate, top acute-acuminate;
upper surface glabrous, drying olivaceous brown to blackish brown, the midrib
glabrous but towards the base in younger leaves finely pubescent, lower surface
glabrous, without larger, brown or blackish dots; midrib moderately raised above;
nerves 13-18 pairs, raised above, marginal arches not distinct; tertiary venation
forming a lax network little or not visible on both surfaces; petioles 12-16 x 2.5-3.5
mm, early glabrescent; leaf bud slender, 12-17 x 2-3 mm, densely grey-brown to
rusty pubescent with hairs c. 0.1 mm long. Inflorescences situated behind the
leaves, rather sparsely pubescent with hairs c. 0.1 mm; in CG: 2 or 3 times ramified,
not many-flowered, 4-10 x 3-5 cm, common peduncle 10-20 mm long, the flowers
in loose clusters of 3-5; Q-inflorescences (according to infructescences) c. 2-5 cm
long; bracts not seen, caducous. Flowers 3- or 4-valved, perianth glabrous, pedicel
glabrous, at base inarticulate. Male perianth subglobose to broadly ovoid, 2.0-2.5
x 2.0-2.3 mm, top shortly rounded to subacute, base rounded; pedicel 1.5-2.0 mm
long; perianth at anthesis cleft to c. 2-*4, not or only slightly collapsing on drying,
valves 0.4-0.5 mm thick, coriaceous. Androecium subellipsoid, c. 1.5-1.6 x 0.8-0.9
mm, in transverse section + blunt-triangular; anthers 5 or 6, at the base curved,
and towards the top erect or somewhat curved, c. 1.6 mm long, largely sessile, free
apices 0.1-0.2(-0.3) mm, apical cavity narrow, 0.2-0.3(-0.5) mm deep, androphore
narrow, 0.1-0.2 mm long, hidden by the anthers. Female flowers not seen. Fruits
1-5 per infructescence, ellipsoid, top and base rounded, 4.0-4.2 x 2.5-3.2 cm,
glabrous, drying rust-brown, finely granulate and without or with a few tubercles or
lenticels, pericarp rather coriaceous, 3.5-8 mm thick; stalk 2-4 mm long; perianth
not persisting.
Distribution. Endemic in C. Celebes.
CELEBES. Central: van Balgooy 3881, 4000; b.b. Cel/II-153, Cel./II-313, Cel./III-27; Johanson,
Nybom, Riebe 133; Meijer 11248; 11278.
Ecology. Primary and disturbed forest (with Jmperata, Gleichenia, Melastoma
and scattered Castanopsis) on ultrabasic soil; 100-700 m alt.. Flowers in March and
November, fruits in April and July.
Vernacular name. Peroso laki (C. Celebes)
NOTES
1. Fieldnotes. Bark and leaves with aromatic scent. Branches horizontal. Cauli-
florous; flowers yellow with strong scent. Perianth fleshy. Ripe fruits orange.
2. Around the insertion of the androecium in most flowers there are a few
minute wart-like appendages or ‘disc-lobes’, c. 0.1 mm high.
3. Vegetatively or in fruit, the present species very much resembles H. costulata,
which has a much larger distribution in Celebes and the Philippines. However, the
latter differs generally by the thinner membranous leaves, with the upper midrib
including the base glabrous, by the lateral nerves usually forming a greater angle
with the midrib, less conspicuous lenticels on the twigs, leaves drying generally
more Olivaceous; also, the fruits are generally larger with a thicker pericarp,
ae Gard. Bull. Sing. 39(1) (1986)
8-10(-15) mm thick. Furthermore, the male flowers are quite different, in H.
costulata arranged in rather dense clusters of 5-10 together.
4. As can be seen from the general key, the present species seems closely related
to H. xanthina (Borneo) and H. majuscula (Sumatra, Malaya), both species having
also a leathery perianth and an elongate androecium, the last-named species also
having similar fruits. In both H. xanthina and H. majuscula, however, the
androphore is broader and tapered and unhidden by the anthers; in H. majuscula
the pedicels are articulate at the base.
5. Sinclair included specimens of the present new species in the broad concept of
H. valida.
97. Horsfieldia penangiana Sinclair Fig. 1D(97)
Horsfieldia penangiana Sinclair, Gard. Bull. Sing. 16 (1958) 408, fig. 42; 28 (1975) 94 — Type: Penang,
Curtis 2406 (SING, n.v.; iso: BM, K).
Myristica griffithii auct. non. Hook. f.: King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 31, p.p., quoad
Curtis 2406, 2458 — Gymnacranthera farquhariana var. griffithii auct. non (Hook. f.) Warb.: Gamble,
Mat. Fl. Mal. Penins. 5, 23 (1912) 226, p.p.
Tree 4-25m. Twigs terete, in the apical portion 1.5-2.0(-4.0) mm diam., grey-
brown to brown, early glabrescent, tomentum grey-brown, composed of hairs up to
c. 0.1 mm long, twigs lower down with the bark finely striate, not flaking, lenticels
small, conspicuous or not. Leaves in 2 rows, membranous to thinly coriaceous,
elliptic-oblong to oblong, broadest at about middle, 5-12 x 2-4 cm, base attenuate,
top acute-acuminate or rounded (S 23689, see notes); upper surface drying oli-
vaceous brown to blackish brown, lower surface early glabrescent, provided with
regularly scattered dark brown to (rarely) pale brown dots (lens, < 30); midrib
above flattish to raised; nerves 8-11 pairs, above thin, flat, inconspicuous or
invisible, marginal arches indistinct or invisible; tertiary venation forming a lax
network indistinct or invisible; petioles 8-13 x 1-2 mm, early glabrescent; leaf bud
slender, 6-9 x 1-1.5 mm, densely greyish-brown pubescent with hairs up to c. 0.1
mm. Inflorescences with rather sparse tomentum, hairs c. 0.1 mm, in C&: 3 or 4
times ramified, moderately- to many- flowered, 2-7 X 1.5-4.5 cm, common pedun-
cle 2-20 mm long; bracts ovate-oblong, short-pubescent, 1.5-2.5 mm long, cadu-
cous; @ inflorescences: 2-5 cm long as judged from the infructescences. Flowers 2-,
3- or 4-valved (see notes), in C’; in loose clusters of 2-5 together, glabrous, pedicel
glabrous, at base + articulate or not (see notes). Male perianth rather variable in
shape (see notes), subglobose to ellipsoid, + circular to faintly triangular in
transverse section, 1.2-1.8 x 1.0-1.5 mm, top rounded, base rounded; pedicel c.
0.8-1.5(-2.0) mm, slender, not tapering; perianth at anthesis cleft to c. 4% to nearly
'2-way, valves c. 0.2 mm thick. Androecium variable, either broadly ellipsoid or
globose or depressed-globose (see notes), c. 0.6-0.7 x 1.0 mm, or perianth ellip-
soid, 0.7-1.5 X 0.6-1.2 mm, in transverse section (sub)circular; anthers 5-9(-10),
almost completely sessile, free apices (0-)0.1-0.2(-0.3) mm, erect or slightly in-
curved, column broad with narrow shallow hollow at apex, androphore up to 0.1
mm long. Female perianth not seen. Fruits 2-6 per infructescence, ovoid-ellipsoid,
1.1-2.0 x 0.9-1.5 cm, top (narrowly) rounded, based rounded, glabrous, without
lenticel-like tubercles, drying brown with finely wrinkled or granulate structure;
dry valves c. 1.5 mm thick; stalks 3-4 mm long, at base articulate; perianth not
persisting.
New account of Horsfieldia 4 53
Distribution. Malaya, Sumatra, Borneo (Sarawak, E. Kalimantan).
MALAYA. Penang Isl: Curtis 2406 (from 2 localities, see notes) — Pahang: Kadim & Noor KN 558;
Shab & Noor MS 1984 — Selangor: Whitmore FRI 0948.
SUMATRA. Tapanuli: b.b. 26117 — West: Mt. Sago, Maradjo 435 — Jambi Prov.: Roos & Franken
1471.
BORNEO. Sarawak (3rd Div.): S 23689 — E. Kalimantan (W. Kutai): b.b. 16872, 16878.
Ecology. Primary dryland rainforest, ridge-top forest, mountainous forest; 0-
1300 m alt. Flowers in June, fruits in August.
NOTES
1. Fieldnotes. The apparently mature fruits of S. 23689 (Sarawak) have been
recorded as dark green, near mature male flower buds as green.
2. Variation. Rather much variation is shown among the few specimens present-
ly considered under H. penangiana, especially in leaf shape and texture, and in the
shape of the male perianth and the androecium. The leaves of specimens from
Malaya and Sumatra have the tip acute-acuminate, as normally in Horsfieldia; the
leaf tip of the specimens from Borneo (S. 23689, and b.b. 16872, 16878) are blunt
or broadly rounded. The leaves of Maradjo 453, from Mt. Sago, West Sumatra, at
c. 1000 m, are conspicuously coriaceous. <
The flowers, including those of the Penang-specimens, are generally 3-(or 4-)
valved. Sinclair quotes from Penang Isl. two specimens, viz. Curtis 2458 (not seen)
and 2406, the type; of Curtis 2406 I have not seen the holotype in SING, but I have
examined two isotypes, in BM and K. Both these apparently belong to the same
species, but according to the labels, and from the details of the flowers when
dissected, they rather differ. The BM collection of Curtis 2406 is annotated as from
between the Coolie Lines and Experimental Nursery: the male perianths in bud are
rather ellipsoid, c. 1.8 X 1.5 mm, the androecium is depressed ellipsoid, c. 1.2-1.5
x 1.2 mm, and bears c. 7(-10) anthers. The K collection of Curtis 2406 consists of
two specimens, one from between the Coolie Lines and the Experimental Station,
the other from Government Hill; the latter has the male perianths globose, c. 1.4
mm diam., with the mature androecium depressed globose, c. 0.6-0.7 x 1.0 mm,
bearing c. 9 anthers.
The flowers of Roos & Franken 1471 from Djambi, Sumatra, deviate from the
type and West-Malesian Horsfieldias, by being all 2-valved. All flowers are rather
immature, but apart from the remarkable deviation, the specimen obviously be-
longs to present species; there are only 5 or 6 anthers.
Pedicels are usually inarticulate at the base, but among the male flowers in one
inflorescence quite often some more or less (not completely) articulate pedicels can
be found.
3. Fruits are only known from one collection from Borneo, $ 23689, a specimen
which deviates by its blunt leaves; its fruit-stalks are distinctly articulate at the base.
The pedicels of the male flowers in all specimens are either inarticulate or only
partly articulate at the base.
54 Gard. Bull. Sing. 39(1) (1986)
4. Horsfieldia penangiana is recognized by its slender twigs with smallish punc-
tate leaves, the very short tomentum of leaf buds and inflorescences, the usually
ellipsoid male perianth, and ellipsoid androecium which is (sub)circular in trans-
verse section. In general habit it resembles Gymnacranthera eugeniifolia. Sterile °
specimens may also recall H. ridleyana. Taxonomically it is close to Horsfieldia
glabra, especially the var. javanica; see there.
5. Sinclair regarded the species as endemic to Penang Isl.
98. Horsfieldia punctatifolia Sinclair Fig. 1D(98)
Horsfieldia punctatifolia Sinclair, Gard. Bull. Sing. 16 (1958) 413, f. 44, pl. XIB: 28 (1975) 105 — Type:
Singapore, fruits, Sinclair SFN 40211 (= Sinclair 7987) (SING, n.v.; iso: K, L, P; BK, BO, DD, E,
Ava).
Tree 7-30 m. Twigs terete, not ridged, towards the apex 2.5-4(-10) mm diam.,
dark grey-brown, early glabrescent, tomentum grey-brown, of hairs up to c. 0.1
mm long; bark finely to + coarsely striate, lower down not flaking; lenticles present
but generally inconspicuous. Leaves in 2 rows, membranous, elliptic-oblong to
oblong, broadest at about or slightly above the middle, 9-21 x 3.9 cm, base
attenuate, top acute-acuminate; upper surface drying dull olivaceous to dark
brown, lower surface glabrous, with scattered larger brown to blackish dots c. 0.05
mm diam.; midrib above flat or slightly raised; nerves 11-16 pairs, thin and flat
above, marginal arches not distinctt; tertiary venation forming a lax network, not
or hardly visible on both surfaces; petioles 10-17 x 1.5-3 mm, glabrous; leaf bud
slender, 8-12 X 1.5-2.0 mm, with dense grey-brown tomentum, of hairs up to 0.1
mm long. Inflorescences sparsely pubescent with hairs up to 0.1 mm or glabrescent,
in OC’; c. 3 times ramified, rather many-flowered, 4-10 x 2-8 cm, common peduncle
5-20 mm long; @ inflorescences c. 2 times ramified, 3-6 xX 1.5-4 cm, fewer-flowered
than in CG: bracts + oblong, c. 2-4 mm, short-pubescent, caducous. Flowers 3- or
4-valved, in C in loose clusters of 2-5, glabrous, pedicels glabrous, at base inarticu-
late. Male perianths depressed-globose, 1.4-2.0 x 1.6-2.2 mm; pedicels 1.0-2.0(-
2.5) mm, slender; perianth at anthesis cleft to 4-%, valves 0.2-0.3 mm thick.
Androecium depressed globose to depressed broadly ovoid, 0.6-0.7 x 1.0-1.5 mm,
top broadly rounded, transverse section nearly circular; anthers 7-9 (14-18 thecae),
completely sessile (free apices up to 0.1 mm), the apices of anthers concealing an
apical, usually small, narrow, hollow up to 0.2 mm deep, in Sumatran specimens
larger, up to c. 0.4 mm deep; androphore narrow, 0.1-0.2 mm long. Female
perianth ellipsoid, 2.8-3.5 x 2.5-3.0 mm, glabrous, cleft at anthesis to c. Y2-way,
valves 0.3-0.5 mm thick; ovary broadly ovoid, 1.5-2.0 xX 1.3-2.0 mm, glabrous,
stigma + irregularly 2-lipped, 0.2-0.3 mm high; pedicel 1-2 mm long. Fruits 1-3 per
infructescence, ellipsoid to broadly ellipsoid, top and base rounded, glabrous,
(4.5-)5.0-8.0 x (3.0-)3.5-4 » cm, drying blackish brown, without or with a few small
warts; dry valves 10-20 mm thick; stalks 4-6 mm long; perianth not persisting.
Distribution. Malaya, Singapore, Sumatra (North, E. Coast), Borneo (Sarawak,
Sabah, C. & E. Kalimantan).
MALAYA (Perak, Selangor, Negri Sembilan, Malacca): FRI 3345, 21674; KEP 76145; King’s Coll.
4078; Maingay 1286; Sinclair SFN 40082.
SINGAPORE. Maxwell 80-157; Murton 76; Sinclair s.n., 7987, 9365, SFN 40211.
Nn
Nn
New account of Horsfieldia 4
SUMATRA. East Coast: Bartlett 6990, 6991, 7300 — Tapanuli: b.b. 5656 — Riouw: b.b. 23216.
BORNEO. Sarawak (4th, 7th Div.): Jacob 6534; S 15917, 22673, 36580, 39341 — Brunei: (Ashton)
BRUN 317 — Sabah: San. 26176, 26276, 30849, 32259, 32278, 66655, 72214, 88407; Sinclair (& Kadim)
9278. — C. Kalimantan: Mogea & de Wilde 4384; Veldkamp 8492 — E. Kalimantan: (Kostermans) b.b.
35017; Kostermans 6713, 7414, 7461, 13623.
Ecology. Primary forest incl. hill side forest, ridge top forest, pole forest, marshy
forest, also kerangas forest; on a variety of soil types incl. grey soil, brown soil,
sandy clay, tertiary sandstone, dacite hill; 0-1100 m alt. Flowers and fruits through-
out the year.
NOTES.
1. Fieldnotes. Bark smooth but shallowly fissured to cracked, or brittle-scaly:
inner bark pinkish to red, with reddish watery exudate; cambium white; slash wood
white, yellowish, or pinkish; heartwood dark brown. Recorded as without or with
low-rounded or steep thick buttresses. Perianths yellow to bright yellow, with a
turpentine odour. Fruits yellow to red, apricot, orange or orange-brown flushed
pink; pericarp pink inside; aril red.
2. The phyllotaxis in S$ 159177 seems + tristichous in the fruit-bearing twig-
portion; in the leafy twig and in all other material seen the phyllotaxis is 2-stichous.
3. Sinclair records the fresh fruits (in SFN 40211, the type) as measuring c. 9 x 6
cm, pointed at both ends; fresh fruits collected in central Kalimantan (Mogea.
Veldkamp) measured about the same size, with pericarp 20-25 thick.
4 .Sinclair regarded H. punctatifolia as characterized by the unique dark brown
punctation of the leaves, not mentioning that several related species, including the
variable H. glabra, have similar punctate leaves. Our present species is particularly
distinguished by its deeply cleft male perianths, rather few anthers, and the large
fruits with a thick pericarp.
99. Horsfieldia macrothyrsa (Mig.) Warb. Fig. 1D(99)
Myristica macrothyrsa Miq.. P|. Jungh. 1 (1852) 172: A DC.. Prod. 14, 1 (1856) 203: Miq.. Fl. Ind. Bat.
1(2). 1 (1858) 66: Suppl. 1 (1860) 156 — Horsfieldia macrothyrsa (Miq.) Warb., Mon. Myrist. (1897)
307; Heyne. Nutt. Pl. 1 (1927) 637 — Type: Sumatra, Tapanuli, Junghuhn (559) (U; iso: BM. K. L).
Tree 4-15 m. Twigs terete, not ridged, towards apex (2-)2.5-6(-8) mm diam.,
brown to blackish brown, early glabrescent, tomentum grey-brown to dull brown.
of hairs c. 0.1 mm long, bark lower down finely striate, not flaking, lenticels usually
distinct. Leaves in 2 rows, membranous, elliptic-oblong to oblong, broadest at or
slightly above the middle, 12-28 x 4-12 cm, base attenuate, top acute-acuminate;
upper surface drying olivaceous to dull brown; lower surface early glabrescent, with
regularly spaced, pale brown to blackish, larger dots (no dashes) (lens x 30):
midrib above flat or slightly raised; nerves 9-17 pairs, above thin, flat on + raised
(Lorzing 1703, see notes), marginal arches not distinct; tertiary venation forming a
lax network, indistinct or invisible on both surfaces; petioles 12-20 « 2-4 mm,
glabrous; leaf bud densely grey-brown to dull brown pubescent by hairs c. 0.1 mm.
slender, 10-15 x 1.5-3 mm. Inflorescences rather sparsely pubescent by hairs c. 0.1
mm or less, in CG: c. 3 times ramified, rather few-flowered, 7-20 x 5-12 cm,
56 Gard. Bull. Sing. 39(1) (1986)
common peduncle 5-45 mm long; Q inflorescences (according to the infructesc-
ences): c. 2 times ramified, 3-6 cm long; bracts elliptic-oblong, 2-4 mm long,
short-pubescent, caducous. Flowers 3 (or 4)-valved, in & in loose clusters of 2-4, in
2 1-3 together, glabrous; pedicels glabrous, at base not or but faintly articulated.
Male perianth globose to (depressed-)broadly obovoid, 3-4.3 x 3-4 mm; pedicel 1-2
mm long, usually slender, well marked-off from the perianth; perianth in anthesis
cleft to c. 2-way, valves 0.2-0.4 mm thick. Androecium globose, depressed globose
or depressed broadly obovoid, 1.8-2.2 x 1.8-2.5 mm, in transverse section circular
to bluntly 3- or 4-angled; anthers 15-22, completely sessile, free apices up to 0.1 mm
long, towards apex somewhat curved into the apical cavity which is rather broad
and ().4-1.2 mm deep; androphore short and narrow, ().1-0.4 mm long, hidden by
the anthers. Female flowers not seen. Fruits 2-6 per infructescence, glabrous,
ovoid-ellipsoid, 2.2-2.5 x 1.6-1.8 cm, top rounded, base (broadly) rounded, with or
without a few small, lenticel-like tubercles, drying brown to blackish, dry valves
1.5-3 mm thick; stalk 2-5 mm long; perianth not persisting.
Distribution. Northern and Central Sumatra.
SUMATRA. Northern (incl. Tapanuli): Junghuhn (559); Lorzing 11703, ‘17195, 17221; Nasution 67
— Central (Mt. Sago): Jacobs 4667; Meijer 3474, 3680, 4029, 4572
Ecology. Lower and mid-mountainous forest, riverine forest; 400-1600 malt.
Flowers and fruits throughout the year. |
NOTES
I. Fieldnotes. A small tree, only 4 to 15 m tall. Bark fissured or peeling some-
what; sap dark red-brown. Wood white to yellowish with red veins. Flowers
greenish to yellow, aromatic. Fruits 2.5-3.5 cm long when fresh, ellipsoid, greenish
to light yellow, valves light yellow inside, aril green (almost mature, as in Lorzing
11703), seed pale yellow.
2. The present species is closely related to H. glabra which has similar fruits, and
flowers of a similar architecture. H. macrothyrsa, however, has markedly larger
male flowers, and about twice as many anthers as compared with H. glabra.
Apparently H. macrothyrsa also has a different distributional area: H. glabra is up
to now not found in Central and North Sumatra.
3. Variation. The specimens presently brought together under the name H.
macrothyrsa rather differ from each other in various aspects. The plants from Mt.
Sago (= Mt. Malintang), C. Sumatra, are relatively weaker in general habit; leaves |
and inflorescences are smaller, the male perianths c. 3-3.5 mm diam. The plants
from the Sibolangit Botanic Garden jungle (see note 4) are stout, with the leaves
large (to 28 cm long), have large male perianths (c. 4.3 « 4.0 mm), and deviate
furthermore by the rather raised nerves on the upper leaf surface (Lérzing 11703,
and others, see note 4). The type-specimen (from Tapanuli) and a collection from
Pematang Siantar (Lorzing 17195, © fls.) are rather intermediate in habit, but
have smaller male perianths (c. 3-3.5 diam.) as are the collections from Mt. Sago.
Lorzing 17195 deviates by short male pedicels which are only c. 1 mm long.
4. The specimens from the Sibolangit Botanic Garden jungle (Lorzing 11703,
17221, Nasution 67) are from specimens collected in 1926 (with fruits) and in 1937
and 1962 (fruits, © flowers), and annotated as growing wild in large numbers. As
ln
~~]
New account of Horsfieldia 4
remarked under note 3, these specimens differ from the rest of the material and
possibly represent a separate taxon.
5. Warburg (1.c. p. 308) describes the perianths of H. macrothyrsa as of c. 2mm
diam., although the type, seen by Warburg, clearly has the strikingly large globose
perianths of c. 3.5 mm diam.
6. Sinclair included the present species in H. glabra, which was accepted by him
as a very large and variable species.
100. Horsfieldia glabra (Bl.) Warb. Fig. 1D(100,100 b)
Myristica glabra B\., Bijdr. 2, 11 (1826) 576; Rumphia 1 (1837) 191, t. 64 fig. 1; Mig., Pl. Jungh. (1852)
172; Fl. Ind. Bat. 1(2), 1 (1858) 65 (excl. M. integra Wall.); Ann. Mus. Bot. Lugd.-Bat. 2, 1 (1865) 49
(excl. var. sumatrana) — Pyrrhosa glabra (B1.) Hasskarl, Cat. Pl. Hort. Bog. (1844) 174 — Horsfiel-
dia glabra (B\.) Warb., Mon. Myrist. (1897) 313, t. 21 fig. 1-2 (p.p.); Sinclair, Gard. Bull. Sing. 16
(1958) 411 (p.p.. for the basionym only); 28 (1975) 35 (p.p.): Backer & Bakh. f., Fl. Java | (1963) 138
— Syntype: Java, Blume (several sheets, L).
Myristica glabra var. grandifolia Miq., Fl. Ind. Bat. 1(2), 1 (1858) 65; Suppl. 1 (1860) 156 — Type: W.
Coast Sumatra, Teijsmann s.n. (U).
Tree 6-25 m. Twigs terete, towards apex 2-4(-8) mm diam., brown to blackish
brown, not ridged, early glabrescent, tomentum from grey-brown to rusty, of hairs
up to 0.2 mm long, lower down bark usually finely striate, not flaking, lenticels
usually conspicuous, especially towards the apex. Leaves in 2 or 3 rows, membra-
nous to thinly coriaceous (and very brittle when dry), elliptic or obovate to
oblong-lanceolate, broadest at or usually above the middle, 8-27 x 3-10.5 cm, base
short- to usually long-attenuate or gradually tapering from about the middle to the
petiole, top acute to acute-acuminate; upper surface drying olivaceous-brown to
dark-brown; lower surface early glabrescent, with regularly scattered brown to
blackish-brown or rarely pale brown, larger dots (not dashes) (lens, x 30); midrib
above flat; nerves 8-16 pairs, above thin, flat or sunken, or in var. glabra sometimes
faintly raised especially towards the base, marginal arches not distinct; tertiary
venation forming a lax network, indistinct or invisible on both surfaces; petioles
10-15 x 1.5-2.5 mm, glabrous (early glabrescent); leaf bud densely grey-brown to
rusty pubescent with hairs 0.1-0.2 mm, slender, 7-12 < 1.5- 2.5 mm, on 3-stichous
twigs slightly thicker. Inflorescences densely to sparsely pubescent with hairs up to
c. 0.1 mm long, sometimes glabrescent, in CG’: (2 or) 3 (or 4) times ramified, rather
many-flowered, 5-10 x 4-7 cm, common peduncle 4-15 mm long; @ inflorescences
1 or 2 times ramified, 2-4 x 1-3 cm; bracts elliptic-lanceolate, c. 2-5 mm long.
short-pubescent, caducous. Flowers (2-) 3- or 4-valved, in © (2-)3-5 in loose
clusters, in Q solitary or 2 or 3 together, glabrous; pedicels glabrous, at base either
not or more or less distinctly articulate, usually mixed in one inflorescence. Male
perianth globose (var. glabra) or broadly ellipsoid or obovoid (vars. javanica &
oviflora), 1.5-2.5 mm long (see further under the varieties); pedicel slender (vars.
glabra & javanica) or thickish (var. oviflora,; perianth at anthesis cleft to c. 2-7,
valves ().2-0.3 mm thick. Androecium (depressed-)globose (var. glabra) or ellipsoid
or short-obovoid (vars. javanica & oviflora), anthers 9-15, almost completely
sessile; androphore short and narrow, (0-)0.1-0.2 mm long; see further under the
varieties. Female perianth ellipsoid, 2.5-3 x 2.2-2.5 mm, glabrous, cleft at anthesis
to c. 43 to nearly '2-way, valves 0.4-0.5 mm thick; pedicel 0.5-1.5 mm long; ovary
ovoid, 1.5-2 1.2-1.5 mm, glabrous, stigma minutely 2-lobed, c. 0.1 mm high (var.
58 Gard. Bull. Sing. 39(1) (1986)
glabra) or if broad-lipped then 0.2(-0.3) mm high (var. oviflora). Fruits 2-6 per
infructescence, ovoid-ellipsoid, top rounded, base rounded to broadly rounded,
glabrous, 1.8-2.4 x 1.4-1.9 cm, drying blackish brown, without lenticel-like tuber-
cles; dry valves 1-2.5 mm thick; stalk 1-2.5 mm long; perianth not persisting.
Distribution. S. Sumatra, Mentawai Isl. north to Simeulué (Simaloer Isl.), Java;
in Java three varieties.
NOTE. In H. glabra the lower leaf surface is always coarsely punctate with dark
brown non-traumatic cork warts, a character which was regarded by Sinclair as
exclusive for the related H. punctatifolia and which it resembles vegetatively.
KEY TO THE VARIETIES
la. Male perianth globose or subglose, 1.7-2.5 mm diam., at anthesis cleft to c. 2-73; androecium
globose or depressed-globose, circular or faintly 3-angular in transverse section; pedicel + slender.
Leaves membranous to chartaceous, nerves flat or slightly raised above. Fruit 18-24 mm long. Area
as the species;\0-600 masalts) ..v.22000) -m. 4582: 20 aS A ee ee a. var. glabra
b. Male perianth broadly ellipsoid to broadly obovoid, at anthesis cleft to c. Y2-way; androecium
ellipsoid to obovoid; blunt-triangular in Section. 1.2)..i005.. 577... nuh nies a, ee ee 2
2a. Male perianth ellipsoid, c. 1.5 mm long; androecium ellipsoid; pedicel slender. Leaves membra-
nous, nerves flat. Fruit notseen. E.Jaya: sion... lioees cos hee ee ee b. var. javanica
b. Male perianth broadly ellipsoid-obovoid, 20-2.5 mm long; androecium broadly ellipsoid-obovoid;
pedicel rather short and thickish. Leaves chartaceous to subcoriaceous, nerves flat or sunken above.
Fruit 18-20mm long. W: & C. Java, 5001500 mi alé; Acca Me atin eee c. var. oviflora
a. var. glabra Fig. 1D(100)
Leaves membranous to chartaceous, up to 27 cm long; nerves above flat or
slightly raised. Male perianth globose, 1.7-2.5 mm diam., at anthesis cleft to c.
2-7/3; valves 3 or 4, c. 0.2-0.3 mm thick; pedicel relatively slender, well marked-off
from the perianth. Androecium globose or depressed-globose, 0.8-1.3(-1.5) x
1.3-1.6 mm, circular or faintly 3-angular in transverse section; anthers 10-15, almost
completely sessile, free apices up to 0.1 mm, curved over towards apex and + into
the apical cavity which is rather broad, 0.3-0.5 mm deep; column broad and solid;
androphore short, narrow, up to 0.2 mm long. Female flowers: stigma minutely
2-lobed, c. 0.1 mm long. Fruits 1.8-2.4 cm long.
Distribution. As the species.
SUMATRA (Central & South): Jacobs 8375; Korthals s.n. — Simaloer (Simeulué) & Enggano Isl.:
Achmad 205, 1342; Ltitjeharms 4259, 4420, 4422.
JAVA (mainly W. & C.): Backer 1163; Bakhuizen v.d. Brink 5444, 5516, 6780; b.b. Ja. 2585, 2617;
Blume 2206 (a/b), 2160 (B); van Borssum Waalkes 799 (Pulau Panaitan); Sinclair 10047, 10048; H.L.B,
565; Hochreutiner 2290; Herb. Legd. Bat./Houtsoorten Gedeh 243, 362, 378, 644; Junghuhn s.n., 230;
Koorders (8) 5198, 5200, 5208, 5284, 13145, 14615, 20173, 20242, 22690, 23711, 24292, 25585, 25595,
26937, 30471, 34030; Kostermans 11133, 23017, 23889; Loos s.n.; Teijsmann s.n. (1867); Warburg
11007; Wirawan 95.
Ecology. Primary and secondary forest, also in coastal forest on limestone; 0-800
m alt. Flowers and fruits throughout the year.
Vernacular names. Java: Ki-mokla lentik (Dyjasilin Sund.); Ki-lalakina, Ki-
minjak, Ki-sareni (Gedeh); Kelapa tjioen, Kelapa, tjiung, Kelapa tjan — Sumatra:
New account of Horsfieldia 4 59
Bonauw falah, Soemaralah silai delok (Simaloer Isl.); Prianggoe Epoekha (Engga-
no Isl.).
NOTES
1. Fieldnotes. Bark smooth to roughish, shallowly longitudinally fissured. Flow-
ers yellow, smelling of Peru-balsam. Fruits glossy greenish-orange, fresh valves to
c. 4 mm thick; aril bright orange.
2. Phyllotaxis is 2- or 3-stichous, sometimes mixed in one collection in specimens
from Java; all specimens from Sumatra are distichous.
3. The male perianths of the Sumatra specimens are faintly ellipsoid, rather than
strictly globose as in Java.
b. var. javanica de Wilde, var. nov. Fig. 1D(100b).
Gemmae foliorum tomento pilis 0.1-0.2 mm longis composito obtectae. Folia membranacea, subtus
punctis nigrescentibus obsita, nervis supra planis. A var. glabra perianthio masculino ellipsoideo ca. 1.5
mm longo atque androecio ellipsoideo differt. — Type: Java, Koorders 52108 (L).
Leaves membranous, 9-12 cm long, nerves above flat. Male perianth ellipsoid, c.
1.5 x 1.2-1.3 mm, in anthesis cleft to c. ¥%2-way; valves c. 0.1-0.2 mm thick;
pedicel slender. Androecium ellipsoid, above subtruncate, 1.0-1.2 x 0.7-0.8 mm,
blunt-triangular in transverse section; anthers 9-15, completely sessile, free apices
0-0.1 mm, at apex not or only slightly incurved; apical cavity small and narrow, c.
0.1-0.2 mm deep; column solid; androphore very short, narrow, up to 0.1 mm long.
Female flowers and fruits not seen.
Distribution. Java, possibly only in E. Java; the two known collections are from
Java without further locality (Koorders 52108, the type) and from E. Java (Be-
soeki, Koorders 21635), without notes on altitude or habitat.
NOTES
According to the flowers this variety seems very related to H. penangiana,
also with dotted leaves and with the perianth of similar size and shape (partly), but
with the androecium round in transverse section and with fewer anthers, 5-9(-10);
also, in H. penangiana the habit of the twigs is more slender, the leaves generally
smaller. However, H. penangiana as accepted by me is variable (especially in
flower shape) and might include the present H. glabra var. javanica when more
material of both taxa becomes available.
The specimens of the present variety were determined by Sinclair as H. glabra.
c. var. oviflora de Wilde, var. nov.
Gemmae foliorum tomento pilis 0.1-0.2 mm longis composito obtectae. Folia subcoriacea, subtus
punctis sparsis obsita, nervis supra planis vel immersis. A var. glabra perianthio masculino ellipsoideo-
obovoideo 2-2.5 mm longo atque androecio ellipsoideo-obovoideo differt — Type: C. Java, b.b. Ja, 3827
(L).
Leaves chartaceous to subcoriaceous, 8-15 cm long; nerves above flat or sunken.
Male perianth broadly obovate-ellipsoid, 2.0-2.5 x 1.7-2.3 mm, in anthesis cleft to
60 Gard. Bull. Sing. 39(1) (1986)
c. Y2-way; valves 0.2-0.3 mm thick; pedicel c. 1 mm long thickish. Andoecium
ellipsoid-obovoid, top subtruncate, 1.2-1.8 x 0.8-1.0 mm, blunt-triangular in trans-
verse section; anthers 10-15, + completely sessile, free apices 0.1-0.2 mm, at apex
little in-curved; apical cavity narrow to rather broad and deep, reaching to nearly
half-way the central column, 0.4-1.0 mm deep; androphore narrow, short, (0-)0.1(-
-).2) mm long. Female flowers: stigma broadly 2-lipped, 0.2-0.3 mm high. Fruits
rather small, 1.8-2.0 x 1.4-1.6 cm.
Distribution: W. and C. Java.
JAVA, b.b: Ja. 3837; Koorders 5197p, 5204, 278748; Kostermans s.n. (May 1968), s.n. (Jan. 1971).
Ecology. Forest at 600-1500 m. Flowers throughout the year, fruits in June.
Vernacular names. Kalak katjoeng, Woeroe timah.
NOTES |
1. Fieldnotes. Bark smooth. Flowers yellow, smelling of Peru-balsam.
2. [have the impression that the phyllotaxis of all 6 specimens seen is 3-stichous.
3. Sinclair included specimens of the present variety in H. glabras.1|. Possibly the
new variety represents a separate species, mainly because of the essentially diffe-
rent shape of perianth and androecium, but because of the rather poor material at
hand I have kept it under H. glabra.
Excluded
As explained under the description and redefinition of the genus Horsfieldia |
have excluded the species of the Horsfieldia macrocoma-complex as published by
Sinclair (1958) for Malaya and as the specimens enumerated by Sinclair (1975) for
its whole area. The species of this H. macrocoma-complex have been removed into a
new genus Endocomia, published in Blumea (1984). Under the genus
Endocomia go over 20 names (mostly combinations in Horsfieldia) formerly tre-
ated in relation to Horsfieldia. These names have presently all been included in the
Index and are referred to as ‘excl.’ (excluded). Their identity can be found through
the index with the article on Endocomia in Blumea.
Further excluded are:
Horsfieldia B\. ex A. DC., Prod. 4 (1830) 87, non Willd., 1805 = Harmsiopanax-
Warb. (Aral.).
Horsfieldia aculeata BL. ex DC. = Harmsiopanax aculeatus (Bl. ex DC.) Warb. ex
Boerl. (Aral.).
Horsfieldia peltata Benth. in Benth. & Hook. f., Gen. Pl. 1 (1862) 937 = Harm-
siopanax aculeatus (Bl. ex DC.) Warb. ex Boerl. (Aral.).
New account of Horsfieldia 4 61
Acknowledgements
During my personal visits to herbaria, which were indispensable, the generous
interest and help extended to me by the respective directors and staff of the
herbaria — British Museum of Natural History in London, the Royal Botanic
Gardens. Kew and the National Museum in Paris — is gratefully recorded here.
The various directors also provided me subsequently with loans of specimens and
types. My travel to Paris was financed by CNRS (National Centre for Scientific
Research) through the mediation of Z.W.O. (Netherlands Organization for the
Advancement of Pure Research).
Dr. R. C. van den Brink supplied the latin translations for almost all the
diagnoses of new taxa and Mr. J. Van Os prepared the beautiful drawings.
Errata
Horsfieldia Pt Gdns’ Bull. Corrections
yr; vol.: pg
1 1984; 37: 124 In ‘Table 1’, column 1, transpose the symbol
‘+’ for 1. H. iryaghedi to line 2 for H. kingii.
2 1985; 38: 87 In ‘2. Variation and resembling species’, H.
salicifolia is an error for Myristica salicifolia.
Bibliography
Armstrong, J.E. & T.K. Wilson, (1978). Floral morphology of Horsfieldia (Myris-
ticaceae). Amer. J. Bot. 65: 441-449.
. (1980). Wood anatomy of Horsfieldia (Myristicaceae). [AWA Bulletin n.s.,
Vol. 1(3): 121-129.
King, G., (1891). The species of Myristica of British India. Ann. Roy. Bot. Gard.
Calc. U1: 275-331, Plates 106-174.
Koster, J. & P. Baas, (1981). Comparative leaf anatomy of the Asiatic Myristi-
caceae. Blumea 27, 1: 115-173.
Sinclair, J., (1958). A revision of the Malayan Myristicaceae. Gard. Bull. Sing. 16:
205-472.
. (1974). The genus Horsfieldia (Myristicaceae) in and outside Malesia I: H.
sabulosa and H. whitmorei J. Sinclair spp. nov. Gard. Bull. Sing. 27: 133-141.
. (1975). The genus Horsfieldia (Myristicaceae) in and outside Malesia II.
Gard. Bull. Sing. 28: 1-181.
Warburg, 0., (1897), Monographie der Myristicaceen. Nova Acta Acad. Leop.-
Carol. 68: 1-680.
Wilde, W.J.J.O. de, (1979). New account of the genus Knema (Myristicaceae).
Blumea 25, 2: 321-478.
. (1981). Supplementary data on Malesian Knema (Myristicaceae), including
three new taxa. Blumea 27, 1: 223-234.
—______.. (1983). Horsfieldia inside and outside Malesia. Acta Bot. Neerl. 32: 117.
—_____. (1984). Endocomia, a new genus of Myristicaceae. Blumea 30:173-196.
62 Gard. Bull. Sing. 39(1) (1986)
Index of Names
Names that are new are printed in bold, those accepted by the author are in roman, and synonyms are
in italics. Numbers in bold are ordinal numbers of descriptions in this work whereas those in roman are
page numbers, and these refer to the Gardens’ Bulletin vol. 37(2).
Embelia ridleyi King & Gamble = excl. brachiata (King) Warb. = 72
Gymnacranthera
farquhariana var. griffithii auct. = 97
ibutii Holthuis = excl.
Horsfieldia B\. ex A.DC. = excl.
Horsfieldia Willd.: see p. 123
sect. Bivalves Sinclair, nom. inval. = sect.
Irya
sect. Horsfieldia = species 1; see p. 126
sect. Irya (Hook.f. & Th.) Warb. = species
6-45; see p. 127
subsect. Euirya Warb., p.p. = sect.
Pyrrhosa
subsect. Euirya Warb. = sect. Irya
subsect. Trivalves Warb. = sect.
Pyrrhosa
sect. Orthanthera Warb., p.p. = sect.
Pyrrhosa
sect. Orthanthera Warb. = sect. Horsfieldia
sect. Phyrrhosa (BI.) Warb. = species 2-5,
46-100; see p. 130
subsect. Bivalves Warb. = sect. Irya
ser. Globularia Warb. = sect. Irya
ser. Smithii Warb. = sect. Irya
subsect. Eupyrrhosa Warb. = sect.
Pyrrhosa
subsect. Papillosae Warb. = excl.
sect. Trivalves (Warb.) Sinclair, comb.
inval.
subsect. Orthanthera (Warb.) Sinclair,
comb. inval. = sect. Horsfieldia
subsect. Trivalves = sect. Pyrrhosa
aculeata B\. ex A. DC. = excl.
acuminata Merr. = 6
affinis de Wilde = 66
amklaal Kanehira = 6
ampla Markgraf = 24
ampliformis de Wilde = 25
amplomontana de Wilde = 88
amygdalina auct. = 47
amygdalina (Wall.) Warb. = 5
var. amygdalina = 5a
var. lanata de Wilde = 5b
androphora de Wilde = 87
angularis de Wilde = 26
ardisiifolia (A. DC.) Warb. = 12
aruana (B1.) de Wilde = 28
aruensis Warb. = 29a
atjehensis de Wilde = 47
australiana auct. = 32
australiana S.T. Blake = 20
bartlettii Merr. = 95
basifissa de Wilde = 31
batjanica Warb. = 7
bivalvis (Hook.f.) Merr. = 10
borneensis de Wilde = 85
var. brachiata = 72
var. laticostata Sinclair = 79
var. sumatrana (Miq.) Sinclair = 81b
bracteosa Henderson = 48
var. bracteosa = 48
var. microcarya Sinclair = 49b
canariformis (Bl.) Merr. = 7
canariodes (King) Warb. = excl.
carnosa Warb. = 69
clavata de Wilde = 22
confertiflora Merr. = 91
congestiflora A.C. Smith = 6
coriacea de Wilde = 96
corrugata Foreman = 40
costulata (Miq.) Warb. = 91
crassifolia (Hook.f. & Th.) Warb. = 68
“crassithyrsa’’ Warb. ex Koord. = 91
crux-melitensis Markgraf = 21
decalvata de Wilde = 38
disticha de Wilde = 76
endertii de Wilde = 83
erubescens Sinclair, in sched. = 32
flocculosa (King) Warb. = 59
fragillima Airy Shaw = 86
fulva (King) Warb. = 53
var. paludiocla (King) Warb. = 68
gigantifolia Elmer = 12
glabra auct. = 5, 16
glabra (BI.) Warb. = 100
var. glabra = 100a
var. javanica de Wilde = 100b.
var. oviflora de Wilde = 100c.
glabrescens Warb. = 44
globularia auct. = 29a
globularia (Bl.) Warb. = 10
var. minahassae Warb. = 10
gracilis de Wilde = 62
grandis (Hook. f.) Warb. = 56
hainanensis Merr. = 2
hellwigii (Warb.) Warb. = 44
var. brachycarpa de Wilde = 44b
var. hellwigii = 44a
var. hellwigii < var. pulverulenta = 42
var. lignosa de Wilde = 44c
var. novobritannica Sinclair = 35b
var. pulverulenta (Warb.) Sinclair = 42
hirtiflora de Wilde = 71
inflexa de Wilde = 8
iriana de Wilde = 27
irya (Gaetn.) Warb., incl. forms ceylanica,
javanica, malayana, moluccana,
siamensis, wallichii = 6
iryaghedhi (Gaertn.) Warb. = 1
karengasicola Sinclair, in sched. = 74
kingii (Hook. f.) Warb = 2
New account of Horsfieldia 4
labillardieri Warb. = 6
laevigata (BI1.) Warb. = 35
var. laevigata = 35a
var. novobritannica (Sinclair) de Wilde
= 35b
lancifolia de Wilde = 37
lauterbachii Warb. = 29a
laticostata (Sinclair) de Wilde = 79
lemanniana auct. = 8la
lemanniana (A. DC.) Warb. = 6
leptantha de Wilde = 43
leptocarpa Warb. = excl.
leptosperma, nom. = excl.
longiflora de Wilde = 3
longipedunculata H.H. Hu = excl.
macilenta de Wilde = 78
macrobotrys Merr. = 60
macrocoma (Miq.) Warb. = excl.
var. canarioides (King) Sinclair = excl.
var. macrocoma = excl.
var. rufirachis Sinclair = excl.
macrothyrsa (Miq.) Warb. = 99
majuscula (King) Warb. = 95
megacarpa Merr. = 91
merrillii Warb. = excl.
minahassae auct. = 91
minahassae (Warb.) Koord. = 10
moluccana de Wilde = 9
var. moluccana = 9a
var, petiolaris de Wilde = 9b
var. pubescencs de Wilde = 9d
var. robusta de Wilde = 9c
montana Airy Shaw = 89
motleyi Warb. = 60
nervosa de Wilde = 80
nesophila auct. = 29a
nesophila (Miq.) Warb. = 35a
novae-lauenburgiae Warb.: see 35b
novo-guineensis Warb., p.p. = 27, 36
novo-guineensis Warb., pro lectotype = 28
var. moseleyana Warb. = 39
nunu Kanehira = 6
oblongata Merr. = excl.
obscura de Wilde = 93
obscurinervia Merr. = 11
obtusa de Wilde = 75
odorata Willd. = 1
olens de Wilde = 17
oligocarpa Warb. = 82
olivaeformis Warb.; see 9
pachycarpa A.C. Smith = 41
pachyrachis de Wilde = 73
pachythyrsa Warb. (‘‘crassithyrsa’”’) = 91
palauensis Kanehira (‘‘palauense’’) = 16
“palewensis’’ auct. = 34
pallidicaula de Wilde = 49
var. macrocarya de Wilde = 49c
var. microcarya (Sinclair) de Wilde =
49b
var. pallidicaula = 49a
pandurifolia H.H. Hu = excl.
papillosa Warb. = excl.
parviflora auct. = 7
63
parviflora (Roxb.) Sinclair = 10
paucinervis Warb. = 63
peltata Benth. = excl.
penangiana Sinclair = 97
pilifera Markgraf = 36
polyantha auct. = 31
polyantha Warb. = 35a
polyspherula (Hook. f. emend. King)
Sinclair = 81
var. maxima de Wilde = 81c
var. oligocarpa (Warb.) Sinclair = 82
var. polyspherula = 8la
var. sumatrana (Miq.) de Wilde = 81b
var. tenuifolia Sinclair = 77
praetermissa Sinclair, in sched. = 41
prainii (King) Warb. = excl.
prunoides C.Y. Wu =5
psilantha de Wilde = 33
pulcherrima de Wilde = 58
pulverulenta Warb. = 42
punctata de Wilde = 90
punctatifolia Sinclair = 98
racemosa (King) Warb. = excl.
ralumensis auct. = 35b
ralunensis Warb. = 45
ramosii Merr. = 11
ramuensis Warb = 29a
reticulata Warb. = 67
ridleyana (King) Warb. = 74
rostrata Markgraf = 29d
roxburghii Warb. = 7
rufo-lanata Airy Shaw = 65
sabulosa Sinclair = 46
samarensis de Wilde = 14
schlechteri Warb. = 30
sepikensis Markgraf = 18
sessilifolia de Wilde = 55
sinclairii de Wilde = 32
smithii Warb. = 15
solomonensis A.C. Smith = 39
sparsa de Wilde = 50
spicata (Roxb.) Sinclair = 7
var. sepikensis (Markgraf) Sinclair =
18
var. spicata = 7
splendida de Wilde = 64
squamulosa de Wilde = 23
sterilis de Wilde = 70
subalpina Sinclair = 92
subsp. kinabaluensis de Wilde = 92b
subsp. subalpina = 92a
subglobosa auct. = 81b
subglobosa (Miq.) Warb. = 6
var. brachiata (King) Sinclair = 72
var. subglobosa auct. = 81b
subtilis (Miq.) Warb. = 29
var. aucta de Wilde = 29¢
var. calearea de Wilde = 29b
var. rostrata (Markgraf) Sinclair = 29d
var. schlechteri (Warb.) Sinclair = 30
var. subtilis = 29a
sucosa auct. = 50
sucosa (King) Warb. = 48
64 Gard. Bull. Sing. 39(1) (1986)
subsp. sucosa = 48a var. grandifolia Miq. = 100
subsp. bifissa = 48b var. sumatrana Miq. = 81b
superba (Hook. f. & Th.) Warb. = 54 globularia auct. = 8la
sylvestris (Houtt.) Warb. = 19 globularia Bl. = 10
var. villosa Warb. = 19 var. subglobosa (Miq.) Miq. = 6
talaudensis de Wilde = 13 (Cnema) glomerata Miq. = 1
tenuifolia (Sinclair) de Wilde = 77 glomerata Thunb. =1
tetratepalaC.Y. Wu = 2 grandis Hook. f. = 56
thorelii Lecomte = 4 griffithii auct. = 97
tomentosa Warb. = 61 hellwigii Warb. = 44 |
tonkinensis Lecomte = 5 horsfieldia auct.; p.p. = 57, 68
var. multiracemosa Lecomte = 5 horsfieldia (“‘horsfieldia”’) B1. = 1
triandra de Wilde = 51 integra Wall. = 81b
trifida A.C. Smith = excl. irya Gaertn. = 6
tristis de Wilde = 52 var. crassifolia Miq. ex Hook. f. = 68
tuberculata (K.Sch.) Warb. = 39 var. /ongifolia King = 6
var. crassivalva de Wilde = 39b var. wallichii King = 6
var. tuberculata = 39a iryaghedhi Gaertn. = 1
valida (Miq.) Warb. = 84 javanica Bl. = 6
villamilii Elmer ex Merr. = 91 kingii Hook. f. = 2
vulcanica Elmer ex Merr. = 91 kurzii King = §
wallichii (Hook. f. & Th.) Warb. = 57 labillardieri (Warb.) Boerl. = 6
warburgiana Elmer = 12 laevigata Bl. = 35
whitmorei Sinclair = 34 lemanniana A. DC. = 6
xanthina Airy Shaw = 94 macrocoma Mig. = excl.
subsp. xanthina = 94a macrothyrsa Miq. = 99
subsp. macrophy Ila = 94b majuscula King = 95
Myristica micrantha Wall. = 6
sect. Caloneura A. DC., p.p. = sect. microcarpa Willd., nom. dub. = 10
Pyrrhosa motleyi (Warb.) Boerl. = 60
sect. Eumyristica Hook. f. & Th., p.p. = nesophila auct. = 27
sect. Horsfieldia sect. Pyrrhosa nesophila Migq., p.p. = excl.
subsect. Horsfieldia (A. DC.) King = nesophila Miq. = 35a
sect. Horsfieldia notha auct. = 1
sect. Horsfieldia A. DC. = sect Horsfieldia odorata Reinw. ex de Vriese = 1
sect. /rya auct. = sect. Horsfieldia sect. oligocarpa (Warb.) Boerl. = 82
Pyrrhosa olivaeformis (Warb.) Boerl.: see 9
sect. Irya Hook. f. & Th. = sect. Irya pachythyrsa (Warb.) Boerl. = 91
sect. Pyrrhosa Bl., p.p. = sect. Horsfieldia, paludicola King = 68
sect. Irya papillosa (Warb.) Boerl. = excl.
sect. Pyrrhosa Bl. = sect. Pyrrhosa parviflora Roxb. = 10
amygdalina Wall. = 5 paucinervis (Warb.) Boerl. = 63
var. f hookeri A. DC. = excl. pendulina Hook. f. = 19
ardisiifolia A. DC. (“‘ardisiaefolia’’) = 12 pinnaeformis Zipp. ex Miq. = 19
aruana Bl. = 28 polyantha (Warb.) Boerl. = 35a
aruensis (Warb.) Boerl. = 29a polyspherula Hook. f. = 81
batjanica (Warb.) Boerl. = 7 prainii King = excl.
bivalvis Hook. f. = 10 pulverulenta (Warb.) Boerl. = 42
brachiata King = 72 racemosa King = excl.
canariformis Bl. = 7 reticulata (Warb.) Boerl. = 67
canarioides King = excl. ridleyana King = 74
carnosa (Warb.) Boerl. = 69 roxburghii (Warb.) Boerl. = 7
collettiana King = 81b rubiginosa King = 56
costulata Miq. = 91 salicifolia Willd. in Roem & Usteri = 19
crassifolia Hook. f. & Th. = 68 smithii (Warb.) Boerl. = 15
“crassithyrsa”’ = 91 spherocarpa Wall. = 6
edulis F.v.M., in sched. = 19 spicata Roxb. = 7
exaltata Wall. ex King = excl. subglobosa Miq. = 6
flocculosa King = 59 subglobosa Miq., p.p. = 68
floribunda Wall. = 5a subtilis Mig. = 29
fulva King = 53 sucosa King = 48
glabra auct. = 5 superba Hook. f. & Th. = 54
glabra Bl. = 100 ; sylvestris Houtt. = 19
New account of Horsfieldia 4
tingens Bl. = 10
tomentosa Hook. f. & Th., nom. illeg. = 61
tuberculata K. Sch. = 39
valida Miq. = 84
vrieseana Miq. = 6
wallichii Hook. f & Th. = 57
Palala
aruana Rumph. = 28
canariformis Rumph. = 7
dentaria Rumph. = 7
globularia Rumph. = 10
“kigil’’ Rumph. = 10
minima Rumph. = 10
quarta Rumph. = 7
quinta Rumph = 10
tertia Rumph. 10
tingens Rumph. = 10
? Phelima Noronha = 1
Pyrrhosa Endl., nom. illeg. = Horsfieldia Willd.
glabra (BI.) Hassk. = 100
globularia (B1.) Hassk. = 10
horsfieldii (B1\.) Hassk. = 1
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Annotated List of Seed Plants of Singapore (X)*
HSUAN KENG
Department of Botany, National University of Singapore
Index to Families
Page Page
Acanthaceae 81 Labiatae 93
Bignoniaceae 79 Lentibulariaceae 77
Boraginaceae 71 Pedaliaceae 81
Convolvulaceae 67 Scrophulariaceae 74
Gesneriaceae 78 Solanaceae ps
Hydrophyllaceae 70 Verbenaceae 87
Il. Angiospermae-Dicotyledons (cont’d)
131. CONVOLVULACEAE
Key to the genera
Pe) tumeic. teatiess, twimitig plants, yelOw OF PIAS... 2.55.20. cccgeec cece sec ec nce ce nsec ececenesveenys Cuscuta
A. Autotrophic, leafy twining or trailing plants, green
B. Style(s) absent: stigma solitary, conical, usually 5-10-rayed; corolla-lobes 2-fid: ovary 1-loculate:
PENNE NITY. 595 0. Gee d. 12. ae Lh. eas Lie Abe. cdi ea. Erycibe
B. Style(s) present
fe Fant Sees HICH CHAT OCU 10h ELUM, CLOCIOUOUS 202. 66-2525 ix ofa seria +» GV Dede cores decpShes setae Porana
C. Sepals enlarged in fruit or not, persistent
D. Pollen grains spiny
E. Fruit a thin-walled capsule. dehiscent; mostly herbaceous twiners: corolla mostly
ens WINNS PY AS less. ie rere ily WB Ot) Sele}, ass dave dows Ipomoea
E. Fruit indehiscent or leathery: woody twiners; corolla mostly with hairy bands outside
Soe iti aga ale pt ane et BE ede lb feeeipd pga gtentl Dive Dip SP -5 ea age Ses Argyreta
D. Pollen grains not spiny
F. Styles 2, free or united near the base
Ra IN MR IONS sd ghd Sok 5k sia ss es an a tae. CORURT Bonamia
G. Styles 2, free
H. Each style forked with 2 filiform stigmas; small herbs, not twining = Evolvulus
H. Each style with a kidney-shaped lobed stigma, long woody twiners ..............
P< SARE RES SPR Pea he DAW de ted oe Pattee FOUN Le gaat hones Sukie ob oUesepectageadeas Neuropeltis
F. Style 1, entire or with 2 minute branches
1. Outer sepals acute or acuminate, much longer and broader than the inner ones .....
EE ee OO ae Saree: ern ee ae eer Bey Pee Aniseia
I. Outer sepals not both distinctly longer and broader than the inner ones Merremia
Aniseia martinicensis (Jacq.) Choisy
Twining herb, the basal parts rooting, to 1.5 m long: leaves narrowly oblong,
3.5-7 cm long; flowers in clusters of 2-3; corolla funnel-shaped, white, 2-3 cm
long; capsule ovoid, 4-valved. Collected once at Kampong Ayer, Bajau (Sinclair
38917).
* Continued from Gdns’ Bull. Sing. 38: 174. 1985.
The author is indebted to Dr. Richard T. Corlett for going through the manuscript and for his
suggestions.
67
68 Gard. Bull. Sing. 39(1) (1986)
Argyreia ridleyi Prain ex Coststr. (= Lettsomia ridleyi Prain)
Woody twiner, densely hairy; leaves elliptic or ovate, 7-16 cm long; flowers
funnel-shaped, 3 cm long, white or purplish, 8-10 together in a head-like cyme.
In damp forests, Bukit Mandai, Chua Chu Kang, Nee Soon (Sinclair SF 40369).
Bonamia semidigyna (Roxb.) Hall.f.
Twiner, to 15 m long; leaves ovate or heart-shaped, 5-16 cm long; cymes
2-5-flowered; corolla funnel-shaped, 3-4 cm long, white. Collected once at Pulau
Merlimau (Sinclair 5926).
Cuscuta australis R. Br. (= C. hygrophilae Pears.)
Twining parasitic plant; stems slender, filiform, yellow or reddish, with haustor-
ia; leaves reduced to tiny scales; flowers small, in clusters. In waste ground,
formerly found in Paya Lebar and Victoria Street (Ridley 12124), parasitic on
Hygrophila quadrivalvis Nees (Acanthaceae); now extinct.
Erycibe festiva Prain
Creeper, to 20 m long; leaves oblong, 7-14 cm long; flowers greenish white,
5-lobed, less than 1 cm across, in axillary clusters. In forest, formerly found in
Gardens’ Jungle (Ridley 6043).
Eryc. griffithii Clarke
Scandent shrub; leaves elliptic to oblong, 7-14 cm long, leathery; flowers white.
In hedges and woods, Gardens’ Jungle, Bukit Timah Road, Changi (Goode-
nough 3846).
Eryc. leucoxyloides King ex Prain
Slender climber; leaves lanceolate to ovate, 1.6-5.5 cm long; flowers white,
solitary, axillary. In open thickets, Changi, Jurong, Gardens’ Jungle, (Bukit
Timah, Ridley 6897, type).
Eryc. maingayi Clarke
Slender woody climber; leaves oblong or ovate, 6-10 cm long; flowers creamy
white. On hedges, and edges of forests, Jurong, Bukit Timah, Tanglin, Mac-
Ritchie (Sinclair SF 40215).
Eryc. malaccensis Clarke
Creeper or scandent shrub; leaves elliptic to ovate, 5-16 cm long; flowers white,
in axillary cymes. In forests, Chua Chu Kang (Hullett 845), Changi.
Evolvulus alsinoides L.
Creeping or prostrate small herb, hairy; leaves lanceolate, to 2.5 cm long;
flowers 1-3 in leaf-axils, light blue to white; capsule round, splitting into 4
1-seeded parts. On sandy shores at Changi (Ridley s.n. in 1890).
Ipomoea alba L. (= I. bona-nox L.) :
Twining herb; leaves ovate-rounded, entire or 3-lobed, 6-20 cm long; corolla
trumpet-shaped, 7-15 cm long, greenish white, fragrant, opening at dusk (thus
called ‘the moon flower’). Native to tropical America, sometimes cultivated.
Ipom. aquatica Forsk. (= /. reptans Poir.)
Herb, trailing on moist soil or on mud in shallow ponds; leaves varying from
Seed Plants of Singapore X 69
linear to ovate, 3-15 cm long, the base cordate or sagittate; flowers funnel-
shaped, 2.5-5 cm long, pink or pale lilac. Pantropical, young shoots and leaves
used as a vegetable. Geylang (Teruja 2945). Vern. Kangkong. 38,
Ipom. batatas (L.) Lamk.
Herb, prostrate and ascending, with large tuberous roots; leaves broadly ovate to
orbicular, 4-14 cm long; flowers bell-shaped, 3-4 cm long, white or pale violet.
Native to the New World, the edible tubers can be found in the market. Vern.
Keledek. #2.
Ipom. cairica (L.) Sweet (= I. pulchella auct. non Roth.)
Twining herb; leaves ovate or rounded in outline, 3-10 cm long and wide,
palmately divided into 5 lanceolate to ovate segments, the basal segments often
lobed again; flowers funnel-shaped, white with purplish red tinge, 4-6 cm long.
Native to tropical Africa and Asia, cultivated and naturalized in waste places and
thickets.
Ipom. digitata L.
Large perennial twiner; stems terete, glabrous; leaves broadly orbicular in out-
line, 6-14 cm long, palmately deeply divided into 5-7 pointed lobes; flowers
funnel-shaped, 5-6 cm long, pale reddish purple. In waste ground and seashores,
sometimes cultivated. Ang Mo Kio (Ridley s.n. in 1894), Bukit Timah Road,
Chang.
Ipom. gracilis R. Br. (= J. littoralis Bl.)
Slender prostrate or twining herb; leaves ovate-cordate, variable in size (1-10 cm
long); flowers funnel-shaped, 3-4 cm long, pink or purple, inside darker near the
base. On sandy beaches and in thickets. Changi, Sungei Morai, Raffles Light-
house (Pulau Satumu) (Burkill & Kiah, HMB 467). Vern. Tarpak kuda.
Ipom. horsfalliae Hook.
Woody climber; leaves orbicular in outline, 5-20 cm long and wide, deeply
palmately lobed almost to the base into 3-5 segments, the mid-segment usually
much larger than the lateral ones; flowers salver-shaped, about 4 cm long, red or
reddish purple. Native of the West Indies, sometimes cultivated.
Ipom. illustris (Clarke) Prain
Woody twiner; stems glabrous or pubescent; leaves ovate to orbicular, some-
times ovate-oblong, 6-16 cm long; cymes axillary; corolla tubular, reddish purple
with a dark centre, rarely white, 10 cm long. Seashore of Tanjong Gul (Sinclair
s.n. 1 1950).
Ipom. pes-caprae (L.) Sweet ssp. braziliensis (L.) Ooststr.
Herb, stems long-trailing and rooting at the nodes; leaves varying, ovate, quan-
drangular to rounded, the apex rounded, truncate or shallowly 2-lobed; flowers
funnel-shaped, 3-5 cm long, reddish purple or rarely white. Common on the
sandy coast, Jurong (Ridley s.n. in 1894), Changi.
Ipom. quamoclit L. (= Quamoclit pinnata Bojer)
Twining annual; stems very slender; leaves oblong in outline, 2-10 cm long,
deeply pinnately lobed, the lobes linear, in 8-20 pairs; flowers 1-4, axillary;
corolla trumpet-shaped, deeply red (rarely white), 2-3 cm long, 5-lobed near the
top. Native to tropical America, sometimes cultivated. Fi ¥#¢ .
70 Gard. Bull. Sing. 39(1) (1986)
Ipom. triloba L.
Twining herb, 1-3 m long; leaves broadly ovate to orbicular in outline, 3-8 cm
long, often 3-lobed; corolla funnel-shaped, pink or pale red-purple, 1.8-2 cm
long. Native of tropical America, found on reclaimed land and waste places.
Ipom. tuba (Schlecht.) G. Don
Glabrous twiner; stems woody; leaves orbicular or ovate, 5-6 cm long, deeply
cordate; flowers axillary, 1 or few together; corolla white, salver-shaped, 9-12 cm
long. Pulau Samulun, near seashore (Sinclair 38844).
Merremia hederacea (Burm. f.) Hallier (= M. convolvulacea Dennst.)
Twining or prostrate herb; stems slender, mostly glabrous; leaves ovate in
outline, 2-5 cm long, the base cordate; flowers cymose, bell-shaped, 0.5-1 cm
long, yellow. In thickets and open grassland, Jurong, Kranji, Bukit Timah
(Ridley s.n. in 1894),
Merr. hirta (L.) Merr. (= M. caespitosa Hallier)
Twining or prostrate herb; stems slender, mostly hairy; leaves variable in form
(linear, oblong to ovate), 3-6 cm long, the base rounded, cordate or hastate;
flowers in cymes; corolla funnel-shaped, 1.5-2 cm long, pale yellow or white. In
open grassland, Chua Chu Kang (Ridley 2686), Kranji, Tanglin.
Merr. tridentata (L.) Hallier f.
Prostrate herb; stems angular, glabrous; leaves varying from linear to oblong, or
spathulate, 4-20 cm long, the apex mostly obtuse or emarginate (a variety, var.
hastata with acute apex), the base often 2-lobed and the lobes toothed; corolla
funnel-shaped, 1-1.2 cm long, yellow or white, with a pink eye. On sandy shores,
Changi (Ridley s.n. in 1891).
Merr. umbellata (L.) Hallier f.
Twining or prostrate herb; stems slender, soft pubescent or glabrous; leaves
ovate or oblong, variable, 4-12 cm long, the base rounded, truncate or lobed;
flowers in umbellate cymes; corolla funnel-shaped, 2-3 cm long, white or orange-
yellow. In thickets or grasslands, Sepoy Lines (Ridley 11956).
Porana volubilis Berum f.
Large woody twiner; leaves ovate, 3-9 cm long; flowers in large panicles; corolla
bell-shaped, deeply 5-lobed, less than 1 cm long, white fragrant. Native to the
neighbouring countries (Burma to Indo-China and Indonesia) but not to Malaya
and Singapore, occasionally planted in gardens.
132. HYDROPHYLLACEAE
Hydrolea zeylanica (L.) Vahl
Creeping herb with erect branches; leaves lanceolate or oblong, 2-10 cm long;
flowers solitary or in panicles; corolla wheel-shaped, 5-7 mm long, blue. Accord-
ing to Ridley, it was formerly found in Geylang, beneath coconut trees in ditches
(no specimens available), now extinct.
Seed Plants of Singapore X 71
133. BORAGINACEAE
Key to the genera
A. Erect shrubs or trees
B. Shrubs or small trees
C. Shrubs or small trees; flowers in unbranched spike-like inflorescences, or in
UMPC ICIUE GAINED. Ot TE ne FU ang Sepa, 2 Sarees) Qs tis geves. sneaks wettacan MUO Cordia
C. Shrubs; flowers solitary or in small cymes, axillary ......................0.0024. Carmona (Ehretia)
B. Herbs; inflorescences scorpioid, usually not branched ...............0.0..0 0. ccc cc ee cee eee Heliotropium
A. Scandent shrubs; inflorescence spike-like, 2-3 times branched .......................0.2.005. Tournefortia
Carmona retusa (Vahl) Masam. (= Ehretia buxifolia Roxb., E. microphylla Lam.)
Erect shrub; leaves spirally arranged, leathery, oblong-ovate, 3-5-tipped, coarse-
ly dentate, 1-6 cm long, dark green and hairy above; flowers axillary, solitary or
in small cymes; corolla white, 7.5 mm across; drupe light red, globose. Native to
Malaya, sometimes cultivated for training as bonsai or miniature plants and
naturalized.
Cordia cylindristachys R. & S.
A bushy shrub, 1-1.5 m tall; leaves elliptic or ovate, 5-10 cm long, rough and
toothed; flowers small, crowded in terminal spike-like inflorescence; corolla
bell-shaped, 4-6 mm long, white. Native to tropical America, introduced to
Singapore as a hedge plant at the end of the last century, became very common in
1940s to 60s; gradually disappearing since the introduction of a beetle (Schemati-
za cordiae).
Cord. dichotoma Forster f. (= C. obliqua Willd.)
Tree, 10-15 m tall; Leaves ovate, 5-8 cm long, thin leathery; flowers in branched
cymes; corolla white, bell-shaped, 1-1.5 cm long. A native of tropical Asia,
formerly planted in kampongs at Ponggol, Changi, Chua Chu Kang (Ridley
8059).
Cord. subcordata Lamk.
Tree, 3-10 m tall; leaves elliptic-ovate or heart-shaped, 7-20 cm long; petiole 2-8
cm long; cymes 3- to many-flowered; corolla orange-red, 2-4 cm long. Once
collected at Pulau Busing (Sinclair 39128).
Heliotropium indicum L.
Erect hairy herb, 10-20 cm tall; leaves ovate-cordate, 3-5 cm long; flowers in
terminal scorpioid cymes; corolla bell-shaped, pale violet with a yellow eye, very
small (2-4 mm long). A weed in waste ground, Kranji (Ridley s.n. in 1894).
Tournefortia tetrandra Bl. (= 7. wallichii DC.)
Scandent herbaceous shrub; leaves fleshy, ovate, 5-8 cm long: flowers in terminal
and axillary branched scorpioid cymes; corolla green, tubular, | cm long, 4-
toothed. In damp shady places, Water Catchment area, Changi (Ridley s.n. in
1890), Jurong.
eZ Gard. Bull. Sing. 39(1) (1986)
134. SOLANACEAE
Key to the genera
A. Cultivated ornamentals or vegetables, usually not producing fruits under local conditions
B. Herbs
C. Corolla funnel-shaped; stamens 51.2020 (0.4), 0.200% Bae Se ee Petunia
C. Corolla tubular, with spreading lobed limb; stamens 4 .................e cece eeeeee eee eee Browallia
B. Shrubs
D Thorny, locally not flowering; leafy branchlets sold as a vegetable .......................4. Lycium
D. Not thorny; corolla long-tubular, curved near apex ................ cece ecececceeececeeeeees Brunfelsia
A. Cultivated or native plants, normally producing fruit
E. Fruit capsular
F. Flowers solitary, terminal, large (to 25 cm long) capsule spiny ...................sceceeeeees Datura
F. Flowers in panicles or corymbose racemes; fruit unarmed ...............:e cece eeeee eens Nicotiana
E. Fruit baccate
G. Anthers forming a cone surrounding the style
H. Leaves 2-3 times pinnate; anthers with narrow sterile tip, opening by slits Lycopersicum
H. Leaves entire, lobed or pinnatifid; anthers without narrow sterile tip, opening at tip by
BHO POPE oo oie. o.oo ol os cakes ROU te ot ORO Riders ite oats yeh aaa Solanum
G. Anthers free
I. Herbs or shrubs; corolla wheel-shaped; stamens exposed ..................cceeeee eee Capsicum
I. Shrubs; corolla salver-shaped, with stamens hidden within ........................0008. Cestrum
Browallia speciosa Hook.
Shrubby, usually less than 1 m tall; leaves ovate, alternate or opposite, 4-5 cm
long; flowers axillary, white, blue or violet; corolla salver-shaped, about 2.5 cm
long. Native to tropical America, sometimes cultivated.
Brunfelsia americana L.
Shrub, 1-2 m tall; leaves oval or ovate, 8-10 cm long; flowers solitary or several
together; corolla salver-shaped, about 5 cm long, opening white and changing to
yellowish, fragrant at night (thus called ‘Lady of the Night’); the calyx less than
’s5 of the corolla tube. Native to tropical America, sometimes cultivated.
Brunf. calycina Benth.
Like the above species, but the calyx about half as long as the corolla which is
whitish or violet in colour. Native to the West Indies, sometimes cultivated.
Capsicum annuum L.
Shrub or shrubby, up to | m tall; leaves ovate, 2-10 cm long; flowers white or
tinged purple, axillary, solitary or several together; fruits various, including: (1)
the long pepper (var. acuminatum) (e.g., the chilli), with red fruits 7-15 or more
cm long, often pendulous; (2) the bell or sweet pepper (var. grossum) with
slightly inflated, globose, ovate or oblong, green or red fruits, 7-10 cm long; (3)
the cherry pepper (var. cerasiforme), with red, yellow or purplish, cherry-shaped
fruits, 1-2.5 cm across. Native to tropical America, widely cultivated. ¥R Ai.
Cestrum nocturnum L.
Shrub, 3-4 m tall, the branches drooping; leaves ovate-lanceolate, 2-6 cm long;
flowers slender, 2-2.5 cm long, clustered at the leaf-axils, more fragrant at night
than day (thus called ‘Night Jasmine’). Native to the West Indies, sometimes
cultivated in gardens.
Datura candida (Pers.) Pasq.
Shrub; leaves short-hairy, ovate or oblong, 20-30 cm long with entire margins;
flowers pendent, white, to 25 cm long (thus called ‘Angel’s Trumpet’), with 5
Seed Plants of Singapore X 73
long-pointed lobes; fruit smooth, ovoid. Native to tropical America; flowers,
seeds and leaves poisonous; occasionally planted.
Datura metel L.
Glabrous herb, 1-1.5 cm long; leaves smooth, narrowly ovate, 15-20 cm long with
wavy margins; flowers erect, pale yellow, white or reddish, single or double,
14-18 cm long; fruit a spiny capsule, globose, 2-2.5 cm across. Native of India,
sometimes planted. BPE.
Lycium chinensis Mill.
Thorny shrub, 1-2 m tall; leaves alternate or sometimes fascicled at the nodes,
lanceolate-oblong, 2-7 cm long. Native to E. Asia. Fresh leafy branches are
found in local markets and dried red berries can be obtained from Chinese
medicine shops. Vern. Chinese box-thorn. fajfG.
Lycopersicum esculentum Mill.
Weak-stemmed herb, to 1 m tall; leaves 15-35 cm long, divided into 5-9-toothed
leaflets; flowers yellow, 3-7 in a cluster; berry (‘tomato’) varies in shape and
dimensions, yellow or red, edible. Native to S. America, often cultivated for the
edible fruits. One variety, the cherry tomato, (var. cerasiforme Alef), with
smaller (1-1.5 cm across) globose, red or yellow fruit, is sometimes grown in pots
as an ornamental. The egg-shaped fruit of the ‘Tree-tomato’ (Cyphomandra
betacea Sendtn.), another native to S. America, sometimes found in the markets,
is also edible and flavoured like the tomato. It can only grow in mountain
stations. &An.
Nicotiana tabacum L.
Herb, to 2.5 m tall, sticky-hairy; leaves oblong-ovate, acute, hairy on both
surfaces; larger blades 30-60 cm long; flowers in panicled racemes; corolla-tube
3.5-6 cm long, widened in upper part, limb 2-3.5 cm across, pale red. The leaves
are the source of commercial tobacco. Native to tropical America, the tobacco
plant was at one time extensively grown in Mandai, Pulau Tekong and else-
where. (RH.
Petunia hybrida Vitmor
Erect or ascending hairy herb; leaves ovate or lanceolate, 1-10 cm long, thick;
flowers axillary, solitary; corolla funnel-shaped, 2-5 cm long, varying in colour,
sometimes variegated. It is a hybrid between two S. American species, P.
axillaris B.S. and P. interifolia S. & T.
Physalis minima L.
Annual herb, 30-50 cm high, soft hairy; leaves opposite, ovate or heart-shaped,
1-9 cm long; flowers solitary, nodding, bell-shaped, pale yellow with brown
spots; berry 1.2 cm across, enclosed in an inflated, reticulate urn-shaped calyx. A
weed, found in grassland and waste places, Pulau Ubin (Ridley 367). »|\\P§ #6 #2.
Solanum ferox L.
Herb, stellate hairy and prickly, to 1 m tall; leaves broadly ovate with a cordate
base, 5-30 cm long, shallowly pinnately lobed, prickly on the veins on both sides;
corolla white or purple, as long as or shorter than the calyx, divided; berry with
stellate hairs. Ridley mentioned that this plant is ‘‘said by Malays to have been
brought from Java’’.
74 Gard. Bull. Sing. 39(1) (1986)
Sol. mammosum L.
Shrubby, to 1 m high. pubescent and spiny; leaves ovate, 8-12 cm long, velvety,
nerves with spines, flowers in short racemes, densely hairy; corolla blue, 1.2-1.5
cm across; fruit inverted pear-shaped, 6-8 cm high, orange, with about 5 short
mamilla-like protrusions from the base. A native toS. America, wild, occasional-
ly planted for its decorative fruits.
Sol. melongena L.
A large stellately hairy herb, sometimes shrubby, to 1 m high, spiny or not;
leaves oblong to oval, 10-30 cm long, often lobed; flowers solitary or a few
together, opposite the leaves; corolla white or purple, 1-1.5 cm long; berry
globose or ovoid, greenish white, purple or black, 10-30 cm long. Native of
tropical Asia, cultivated for the edible fruits. Vern. Egg Plant. fp.
Sol. nigrum L.
Herb, 30-80 cm high; leaves ovate, 1.5-10 cm long, with wavy margins; flowers 2
to many in an extra-axillary cluster; corolla white, 0.5-1 cm across; ripe berry
black, about 0.5 cm across. A weed in gardens and waste places, variable;
Geylang (Ridley 8084). Vern. Ranti.
Sol. torvum Sw.
Shrubby, to 1.5 m high; stems prickled; leaves ovate, 5-10 cm long, lobed, hairy;
flowers in cymes; corolla white, 1-1.2 cm across; ripe berry yellow. A weed in
waste ground, Tanglin (Ridley 12383). 7K hn.
Sol. tuberosum L.
The potato is a native of tropical America, now widely cultivated in many parts
of the world. The starchy underground tubers are imported for food. FE 9 52.
Sol. wrightit Benth.
An ornamental tree, 5-10 m tall; twigs smooth or sparsely prickly; leaves ovate,
9-20 cm long, shallowly or deeply lobed, unequal at base; flowers in cymes;
corolla blue or purple, 3-5 cm across; ripe berry globose, 3-5 cm across. Native to
Peru, sometimes cultivated.
135. SCROPHULARIACEAE
Key to the genera
A.; Slender herbs, tiny, parasitic on, grass-fOots, s -: sus..++>casaupens van <opagds deeka wee onal eee Striga
A. Autotrophic herbs, not parasitic
B. Corolla not or less clearly 2-lipped
C. Flowers axillary, solitary or in pairs; corolla wheel-shaped, 4-lobed ....................5 Scoparia
C. Flowers in terminal spikes; corolla cylindric, 5S-lobed ................:ceceeeeeeenenee es Centranthera
B. Corolla cylindric or campanulate, distinctly 2-lipped
D. Calyx-lobes very unequal
E. Stamens 4, 2 or all with one reduced anther-locule ..............+050+..00 apne Adenosma
E, Stamens 4, all anther-locules perfect .4..4..5;3:...5.t/ad.es)- heya gee ee Bacopa
D. Calyx-lobes + equal
F. Fertile stamens 2
G-~ Calyx 54obed, the lobes narrow’ ..0..000 00.0.2. ,.i..210 te oe Lindernia (p.p)
G. Calyx 5-toothed «.......5500 07.2001. UNL). AD ee Microcarpaea
Seed Plants of Singapore X 75
F. Fertile stamens 4
oe eR PERUSE SOOT INNS Bg ir okie 58 Gob nies po on dw'acinloh dale dh dbase sadder noes Limnophila
H. Anther-locules contiguous
I. Corolla with a 2-lobed palate in the throat .....0....0.0 0. ccc cece cece ec eeceeeecuees Mazus
I. Corolla without appendages in the throat ...........0.00.00.00c cece Lindernia (p.p.)
Adenosma indicum (Lour.) Merr. (= A. capitatum Benth.)
Aromatic herb, 30-60 cm tall; leaves ovate, 3-5 cm long; flowers in dense
terminal heads with many leafy bracts at the base; corolla 6-7 mm long, blue,
2-lipped. In sandy fields by roadsides, Tanglin, Chua Chu Kang, Jurong (Ridley
1826), Serangoon.
Aden. inopinatum Prain
Erect herb; stems hairy; leaves ovate, 2.5-4 cm long; flowers solitary, axillary;
corolla white or purplish. In grassy spots, Tanglin, Bukit Timah (Nur 25986).
Aden. javanicum (BI.) Koord. (= A. ovatum Benth.)
Pubescent low creeping herb, 15-50 cm tall; leaves ovate, 1.5-2 cm long; flowers
axillary, sessile; corolla pale blue. On roadsides, Tanglin, Chua Chu Kang, Bukit
Timah Road (Ridley 6894).
Angelonia salicariaefolia Humb. & Bonpl.
Perennial herb, 60-70 cm tall; leaves in a spiral above, opposite and verticillate
below, narrow lanceolate, 4-5 cm long; flowers in narrow terminal clusters;
corolla short and swollen, 2-lipped, 1-1.2 cm long, blue. A cultivated ornamental
from tropical America.
Bacopa monnieri (L.) Pennel (= Herpestis monnieri (L.) Rothm.)
Small glabrous herb, 10-15 cm tall; leaves obovate-spathulate, 1-2 cm long;
flowers solitary, axillary; corolla 2-lipped, 1 cm long, white. In wet places. Bukit
Timah, Tampinis, Rochore, Serangoon (Ridley s.n. in 1891).
Centranthera tranquebarica (Spreng.) Merr. (= C. humifusa Wall. ex Benth.)
Erect annual herb, 10-15 cm tall, glabrous; leaves linear, 1-1.5 cm long; flowers
axillary; corolla tubular, curved, 5-lobed; stamens 4, in 2 pairs. In grassy banks
and roadsides, Bukit Timah Road (Ridley s.n. in 1898).
Limnophila laxa Benth.
Small annual herb; stamens about 30 cm long, ascending; leaves opposite, linear
or elliptic; flowers axillary, solitary or few in a cluster; corolla 2-lipped, less than
1 cm long. In damp places, Seletar (Hullett 588, not seen).
Limn. sessiliflora B\. (= Ambullia sessiliflora Baill. ex Wettst.)
Aquatic slender herb; leaves of two forms, the submerged ones 3-6 in a whorl,
finely pinnatisect, the aerial ones opposite, narrowly spathulate, toothed, 2-2.5
cm long; flowers solitary, axillary, on aerial branches; corolla pink, 1-1.2 mm
long, 2-lipped; stamens 4, in 2 pairs. Introduced water plant; common in aquaria,
native to northern Malaya and S. Asia.
Limn. villosa BI.
Small aromatic herb, pubescent, 10-15 cm long; leaves lanceolate to elliptic,
crenate, 1-1.5 cm long, sessile; flowers solitary, axillary; corolla violet, 5 mm
long. In ditches and damp spots, Tanglin, Chua Chu Kang, Bukit Timah (Goode-
nough s.n. in 1891).
76 Gard. Bull. Sing. 39(1) (1986)
Lindernia anagallis (Burm.f.) Penn. (= Vandellia pedunculata Benth.)
Annual herb, suberect to prostrate, 30-50 cm tall, rooting at the nodes; leaves
shortly petioled or subsessile, varying from linear to broadly ovate; flowers
solitary, axillary; corolla 2-lipped, 7-10 mm long, white to purple; stamens 4, in 2
pairs; capsule cylindric, to 1.3 cm long. In grasslands; Geylang (Ridley s.n. in
1896).
Lind. antipoda (L.) Alston (= Bonnaya veronicaefolia Spreng.)
Slender herb, 10-25 cm tall, tufted, sometimes creeping; leaves ovate, 1.5 cm
long; flowers axillary, solitary; corolla 2-lipped, 5-10 mm long, violet or white,
with yellow spots. In wet places, Ang Mo Kio, Chua Chu Kang (Ridley 3948).
Lind. ciliata Penn. (= Bonnaya brachiata L. & O.)
Slender erect herb, 5-15 cm tall; leaves ablong, 1-2 cm long, deeply serrate,
sessile; flowers in elongate racemes; corolla 2-lipped, white, 1 cm long; stamens
in 2 pairs, only the upper pair functional. In damp spots by the stream, Tanjong
Katong (Ridley s.n. in 1906).
Lind. crustacea (L) F.v.M. (= Vandellia crustacea Benth.)
Small herb, erect or decumbent and rooting at the nodes; leaves lanceolate or
ovate, 1-2 cm long; flowers axillary; corolla 2-lipped, violet with dark purple
blotches. In waste ground, often a garden weed; Tanglin, Bukit Timah (Ridley
2700). Vern. Kerak nasi.
Lind. ruellioides Penn. (= Bonnaya reptans Spreng.)
Stoloniferous herb, tufted, 1 cm high; leaves oblong-obovate, 1-1.5 cm long,
serrate; flowers in racemes; corolla 1.5 cm long, violet-white. In damp grassy
spots by streams; Chua Chu Kang (Ridley 2940), Bukit Panjang, Tanglin.
Lind. sessiliflora (Bth.) Wettst. (= Vandellia sessiliflora Bth. )
Annual herb, to 10 cm tall; stem creeping and rooting at the nodes; leaves
broadly ovate to suborbicular, 1.5-2.5 cm long, subsessile; flowers axillary,
solitary or in pairs; corolla white; capsule cylindric, 1.2 cm long. A weed
sometimes found in gardens and wasteground.
Lind. viscosa (Hornem.) Merr. (= Vandellia hirsuta Bth.)
Small herb, 8-10 cm high, hairy; leaves oblong to ovate, 2-5 cm long, crenate;
flowers small, in a lax terminal raceme; corolla 2-lipped, 3-4 mm long, pale lilac.
In sandy spots, Tanglin, Botanic Gardens (Purseglove 4029).
Mazus rugosus Lour.
Small creeping herb; Leaves obovate, 1.5-2 cm long, crenate; flowers in a
terminal raceme; corolla 2-lipped, 4-5 mm long, pale violet, white with yellow
spots in the centre of the lip. A weed on waste ground, Botanic Gardens (Ridley
s.n. in 1889).
Microcarpaea minima (Koen.) Meer. (= M. muscosa R. Br.)
Very slender herb, 3-10 cm tall, tufted; leaves opposite, oblong, 2-3 mm long;
flowers solitary, axillary; corolla cylindric, 5-lobed (1 linear oblong, 4 short,
acute, all fringed with white-hairs); fertile stamens 4. Edge of damp spots or
ponds, ‘Reservoir’ (Ridley 12513).
Seed Plants of Singapore X 77
Russelia equisetiformis Champ. & Schlecht. (= R. juncea Zucc.)
Shrubby, 30 cm to 1 m tall; branches 4-angled, green, drooping; leaves oe
reduced if present, linear-lanceolate or ovate; flowers in a simple terminal
panicle, nodding; corolla tubular, 2-2.5 cm long, 5-lobed, bright red, A native of
Mexico, planted for its red fire-cracker-like flowers. Vern. Coral Plant.
Scoparia dulcis L.
Herb, 30-60 cm tall; leaves opposite or verticillate, lanceolate, 3-5 cm long,
serrate; flowers axillary, solitary or in pairs; corolla pale purple, 4-lobed, with
long white hairs in the throat; stamens 4. Native to tropical America, naturalized
in wasteland.
Striga asiatica (L.) O. Ktze. (S. hirsuta Benth.)
Tiny stiff erect herb, semi-parasitic on grass roots, 8-15 cm high; leaves linear,
opposite below and alternate above, 1-1.5 cm long; flowers axillary, solitary;
corolla 2-lipped, 1-1.2 cm long, yellow or pink. In grassy places and on roadsides,
Kranji, (Ridley s.n. in 1894). Vern. Rumput siku-siku.
Torenia polygonoides Benth.
Small creeping herb, 10-30 cm long; leaves ovate 1-1.2 cm long, serrate; flowers
axillary; calyx tubular, 3-winged; corolla 2-lipped, upper lip reddish brown, the
lower white. Common in grassland and on banks, Tanglin (Ridley s.n. in 1980).
coal! 136. LENTIBULARIACEAE
Utricularia bifida Linn.
Terrestrial slender herb; stolons few; leaves linear-spathulate, 1-1.5 cm long, on
the stolons; flowering stalk 5-15 cm high; flowers yellow, 6-8 mm long. In damp
sandy spots and marshes; MacRitchie Reservoir, Chua Chu Kang (Ridley s.n.
1894), Bukit Timah.
Utric. caerulea L. (= U. albina Ridl.)
Terrestrial slender herb; flowering stems 8-15 cm tall; leaves rosulate, linear-
spathulate, to 7 mm long; flowering stalk to 30 cm long; flowers white or pink,
subcapitate; corolla 0.4-1 cm long. In grassy sandy spots, Changi (Ridley 1474),
MacRitchie Reservoir, Telok Kurau.
Utric. exoleta R. Br.
Floating plant; stolon filiform; leaves slender, capillary, with bladders; traps
obliquely ovoid, 1-1.5 mm long; flowering stalks 5-15 cm tall; flowers yellow, 4-8
mm long. In ponds and ditches; Botanic Gardens’ lake, Ang Mo Kio (Ridley
s.n. in 1889).
Utric. punctata Wall. ex A.DC. (= U. fluitans Ridl.)
Aquatic; stolons filiform, to 20 cm long; leaves finely dissected, 2-6 cm long;
flowering stalks 8-30 cm long; flowers violet or white; corolla 6-10 mm long. In
swamps, Changi Road (Ridley 5642).
Utric. uliginosa Vahl (= U. griffithii Wight) .
Terrestrial slender herb; stolons 30-40 cm long; leaves linear or linear-spathulate,
1.5-2(-4) cm long; flowering stalks to 30 cm long; flowers bright purplish blue;
78 Gard. Bull. Sing. 39(1) (1986)
corolla 3-18 mm long. In wet sandy places or in shallow water; Changi (Ridley
s.n. in 1889), Bedok, Tampines.
137. GESNERIACEAE
Key to the genera
A. Native plants
By ‘Creeping epiphytes; fertile stamens 4; fruit linea 3.5.) ...9..24J...5) 634; deena Aeschynanthus
B. Erect herbs or shrubby; fertile stamens 2
C. Fruit ovoid, fleshy, in@ehiscent .............00030<00+0ceseuweeeden seen ieee ee Cyrtandra
C. Fruit linear, dry, dehiscent. .. ....4.24-rere-61: paappaye. 2). eee Didymocarpus
A. Cultivated pot plants
Db. Fertilestamens 2; coroila-tube short «02005. -02.. dh. a4s-deags 002g gee ae Saintpaulia
D. Fertile stamens 4
E. Herbs with stoloniferous stems; corolla-tube trumpet-shaped ......................000000: Episcia
E. Herbs with very short stems; corolla bell-shaped,./....2.4.4%00-4. 4th. 2k oeeea aoe Sinningia
Aeschynanthus parviflora R. Br.
Creeping epiphyte on trees; leaves leathery, lanceolate-ovate, 4-5 cm long;
flowers 5 cm long, curved, deep red. In mangroves; Kranji, Sungei Tengah, Tuas
(Goodenough s.n. in 1890).
Aeschy. purpurascens Hassk.
Slender epiphyte, on trees or branches; leaves fleshy, lanceolate to ovate, 8-10
cm long; flowers light green, tipped purplish brown, 2-3.5 cm long. In forests,
Chua Chu Kang (Ridley s.n. in 1892).
Aeschy. radicans Jack
Creeping on rocks and epiphytic on tree trunks; leaves ovate, 1.5-3 cm long,
hairy; flowers axillary, solitary or in pairs, bright red. In forests, Bukit Timah
(Md Nur 24637).
Aeschy. wallichii R. Br.
Shrubby, slender and glabrous; leaves lanceolate or elliptic, 5-7 cm long; flowers
axillary, several together, 1.5-2.5 cm long, dark red. In dense forests; Bukit
Mandai (Goodenough s.n. in 1890), Kranji.
Cyrtandra pendula BI.
Herb; stems creeping and ascending; leaves alternate, ovate, 10-20 cm long,
reddish hairy beneath; cymes axillary; corolla funnel-shaped, creamy white or
pale yellow, 3-4 cm long, with purple spots. In forests, Bukit Timah (Ridley s.n.
in 1890).
Didymocarpus perdita Rid.
Shrubby, 10 m long; leaves elliptic, peltate, 6-7 cm long, crenulate; peduncles
slender, hairy. Endemic to Singapore, only 2 specimens were collected, in
ravines, Seletar (Ridley s.n. in 1889, type).
Didym. platypus Clarke
Shrubby; stem woody; leaves oblanceolate, 20-30 cm long, sessile, serrate,
sparsely hairy; flowers in axillary cymes; flowers white with a yellow mouth, 2-3
cm long. In forests, Bukit Timah (Hullett 454), Seletar.
Seed Plants of Singapore X 1%
Episcia fulgida (Lind.) Hook. f.
Perennial herb; stems stoloniferous; leaves opposite, elliptic or ovate, greenish
with brown lines and blotches, 5-7 cm long; flowers solitary, axillary, corolla
salver-shaped, bright red, lobes 5, rounded. Native to Colombia, cultivated
commonly as a pot plant. Another species, E. cupreata (Hook.) Hanst., a native
of Nicaragua, with hairy reddish green leaves and green veins and red flowers, is
also planted.
Saintpaulia ionantha Wendl.
Perennial herb, hairy; leaves succulent, oblong or rounded, 3-4 cm long, long-
stalked; flowers 1-6 on a long peduncle; corolla short-tubed, 2-lipped, 2-2.5 cm
across, violet or in various other colours. Native to tropical Africa (‘African
Violet’); many forms, some with double flowers. JE}\ =.
Sinningia speciosa (Lodd.) Hiern.
Hairy herb; stems short or creeping underground; leaves oblong or ovate, hairy,
8-15 cm long, long-stalked; flowers bell-shaped, 10-15 cm long, variously col-
oured (violet, purple, red or white), with 5 round lobes. Native to Brazil,
sometimes cultivated as a pot plant, horticulturally known as ‘Gloxinia’ #445
138. BIGNONIACEAE
Key to the genera
A. Woody climber or twiners
B. Leaves simple or 2-foliate with a long tendril; flowers reddish violet ......................... Saritaea
B. eaves pinnate, leaflets 3-15; flowers creatiy white }..262 j2.0..2. <0. icc. cc0e gc ee cem eens seen Pandorea
A. Erect shrubs or trees
C. Leaves simple, in whorls or 3-4 each; flowers bright yellow ......................0000eeeee Deplanchea
C. Leaves compound
D. Leaves palmately compound, leaflets 1-5; flowers yellow or pinkish ................... Tabebuia
D. Leaves pinnately compound
E. Leaves once pinnately compound
F. Shrubs, rarely small trees
G. Native shrubs (or small trees) by tidal rivers; corolla trumpet-shaped, white ........
oh IES LE Re Bt eT ON oe he te Se re Dolichandrone
G. Introduced shrubs, cultivated in gardens
Hi (Corolla bell-shaned bright yellow, osc ied.5 2622-5 oh sas Gees ak dws Tecoma
H. Corolla narrow trumpet-shaped, orange red
F. Tall trees
I. Capsules long-linear, twisted or curved; corolla pinkish or dull yellow, fringed ......
DSi eee ue aie SURG es Sis OF Coe aia ola alan om Sen ke oe nae e Ges wakc anew vers sab asesetaaseces Stereospermum
I. Capsules oblong, not twisted
J. Calyx campanulate, 5-lobed; corolla pale yellow .......................55. Pajanelia
PA any ek Sette KG. COPMMa OTaMBe TE... oi... 5... cas enene sees ences .... Spathodea
E. Leaves 2-3 times pinnate
K. Leaflets large (4-12 cm long) and few; corolla trumpet-shaped, pinkish white, capsule
oR Se dpe EAS Ey PS Sas Se) Pee Se ee Radermachera
K. Leaflets small (0.5-1 cm long) and numerous; corolla bell-shaped, violet, capsule
a a Bk eC a oa ny d vile wnt pweyup bok wansaepee Jacaranda
Deplanchea bancana (Scheff.) Steen.
Tree, about 30 m tall; leaves simple, 3-4 in a whorl, obovate to elliptic, 9-30 cm
long, glandulate at base; flowers in terminal panicles; corolla tubular, 3-3.5 cm
long, bright yellow; stamens 4, in 2 pairs; capsule elliptic, 10-14 cm long. In
forest, very rare; Kranji (Goodenough s.n. in 1889), Bukit Timah, Seletar.
80 Gard. Bull. Sing. 39(1) (1986)
Dolichandrone spathacea (Linn. f.) K. Schum.
Shrub or tree; leaves pinnate; leaflets 3-11, lanceolate or ovate, 6-20 cm long;
flowers 2-10 in terminal raceme, opening in the early morning and closing at
noon; corolla white, trumpet-shaped, 16-20 cm long; stamens 4; capsule linear,
curved and compressed, 25-60 cm long. Formerly common along tidal rivers,
Ponggol, Bajau, Pulau Ubin, P. Tekong, Tanjong Gul (Burkill HMB 2791).
Jacaranda filicifolia (Anders.) D: Don
Small tree; leaves opposite, bipinnate; leaflets numerous, elliptic, oblique, 0.5-1
cm long; flowers in terminal panicle; corolla bell-shaped, violet with white-
blotched throat; stamens 4, with a long hairy staminode; fruit ovate, 4-7 cm long.
A native of S. America, sometimes planted.
Pajanelia longifolia (Willd.) K. Schum. (= P. multijuga DC.)
Tree, little branched, 20-25 m tall; leaves pinnate, 40-120 cm long; leaflets 8-12
pairs, obliquely ovate, 10-12 cm long; flowers in terminal thyrses; corolla pale
yellowish, dark purple inside, bell-shaped, 5-7.5 cm long, thick and hairy;
capsule flat, obovate, 30-45 cm long. Formerly recorded from the coast of Kranji
(? Cantley s.n. in 1782), now extinct.
Pandorea pandorana (Andr.) Steen.
Climber; leaves opposite, pinnate; leaflets 3-13, varying from linear to orbicular,
2-8 cm long; flowers in cymes; corolla funnel-shaped, white or creamy, 2.5 cm
long. Native to E. Malesia, sometimes cultivated.
Radermachera gigantea (BI.) Mig. (= R. lobbii Migq.)
Shrub or tree, leaves mostly twicely pinnate, 12-35 cm long; leaflets often elliptic,
4-12 cm long; flowers in terminal panicles; corolla trumpet-shaped, 5-6 cm long,
white tinted pink, with yellow streaks in the throat; capsule linear, 15-60 cm long.
Tanglin, Bukit Timah, Bukit Mandai and Reservoir Jungle.
Saritaea magnifica (Steen.) Dugand (= Arrabidaea magnifica Steenis)
Climbing shrub; leaves opposite; lower leaves simple, obovate, 8-12 cm long;
higher ones 2-foliate with a long terminal tendril; flowers bell-shaped, 6-8 cm
long, reddish violet. Native of Colombia, cultivated or escaped.
Spathodea campanulata Beauv.
Tree; leaves opposite or in whorls of three, pinnate; leaflets 7-17, ovate-oblong,
4-12 cm long; flowers in terminal racemes; calyx closed in bud, spathe-like in
flowers, containing watery fluid inside, corolla orange-red, obliquely bell-shaped
(thus ‘African Tulip Tree’), 11-13 cm long, broadly 5-lobed; stamens 4; capsule
erect, oblong-lanceolate, 15-20 cm long. Native of tropical Africa, more or less
naturalized. iki.
Stereospermum fimbriatum (Wall. ex G. Don) DC.
Deciduous tall tree; leaves pinnate, opposite, 30-75 cm long; leaflets usually 7,
rarely 5 or 9, ovate-oblong, 8-16 cm long; flowers in large spreading clusters on
the bare twigs; corolla pale pinkish, narrowly funnel-shaped, 6-7 cm long, the
lobes finely long-fringed; capsule 4-angled, 35-60 cm long, twisted (thus ‘Snake
Tree’). Native of Burma and.Malaya, sometimes planted by roadsides.
Ster. personatum (Hassk.) Chatterjee (= Ster. chelonoides A.P. DC.)
Deciduous tree; leaves pinnate, opposite, 20-50 cm long; leaflets 7-13, elliptic-
oblong, 5-15 cm long; flowers in terminal panicles; corolla dull yellow, trumpet-
Seed Plants of Singapore X 81
shaped, 2-3 cm long, with dark red stripes inside; capsule long-linear, to 45 cm
long, 4-ribbed, curved. Rare. Tanglin, Bukit Timah (Ridley 3642).
Tabebuia chrysantha Nichols
Small spreading tree; leaves opposite, palmate; leaflets 5, elliptic pointed. Cen-
tral one the largest (6-25 cm long); flowers in cymes, on old branches, yellow,
funnel-shaped, 9-12 cm long. Native to Brazil, recently cultivated as a roadside
tree; other species including the pinkish flowered T. pallida Miers (leaflets
obtuse, capsules smaller, less than 15 cm long from the West Indies) and T. rosea
DC. (leaflets acute, capsules larger, over 22 cm long, from Central America).
Tecoma stans (L.) HBK. (= Stenolobium stans Seem.)
Erect shrub, 2-5 m tall; leaves opposite, pinnate; leaflets 7-11, ovate-lanceolate,
4-12 cm long; flowers in terminal racemes; corolla bell-shaped, bright yellow
(thus ‘Yellow Bell’), 3-4 cm long; capsule linear, 10-18 cm long. Native to South
America, planted as ornamental in gardens. ###/E.
Tecomaria capensis (Thunb.) Spach
Straggling shrub; leaves opposite, pinnate, leaflets 5-9, ovate, acute, 1-3.5 cm
long; flowers in terminal racemes; corolla orange-red, narrowly funnel-shaped,
3-4 cm long; capsule linear, 6-8 cm long. Native of S. Africa, often planted in
gardens.
139. PEDALIACEAE
Sesamum orientale L. (= S. indicum L.)
Erect herb, covered with long and short (mucilaginous) hairs; lower leaves
opposite, upper ones spirally alternate, oblong-lanceolate, 4-10 cm long; flowers
axillary, solitary; corolla white (pale-seeded form) or violet (black-seeded form),
obliquely and narrowly bell-shaped, 2.5-3 cm long; capsule erect, 4-angled.
Native of Africa; formerly cultivated for its seeds (the ‘Sesame seeds’), now an
escape occasionally found on wasteland. AA fii , ABS.
140. ACANTHACEAE
I. Synoptic key to the genera
1. Subshrubs, in mangroves or on muddy shores; corolla 2-lipped, with the upper lip obsolete, lower
RR TT 2 ce EEE RS a a ee oe Acanthus
1. Not as above
2. Scandent herbs or shrubs, rarely erect; calyx small, ring-shaped or 10-15-toothed ...... Thunbergia
2.Otherwise
3. Corolla-lobes in bud twisted to the left
4. Ovules 3-12 in each locule; capsule usually 6- or more-seeded
oe A OE |e ot en ee ey Ore ee ee eet Cee Hemigraphis, Ruellia
ne i ediennside sou etp eas <peveumonasaiens Hygrophila
4. Ovules 2 in each locule; capsule 4- or fewer-seeded ................. 6c ccc eee ee ee ees Strobilanthes
3. Corolla-lobes in bud imbricate
6. Ovules 3-10 in each locule; capsule usually 6- or more-seeded
6. Ovules 2 or | in each locule; capsule 4- or fewer-seeded
7. Corolla nearly regular subequally 5-lobed ......................cceeeeeeeees Asystasia, Barleria,
tin ERE A IS Sa, SC ee ee Crossandra, Eranthemum
JathAst a cvbsg dues ass Andrographis
82 Gard. Bull. Sing. 39(1) (1986)
II. Key to some ornamental genera and species*
1. Corolla nearly regular, limb subsequently 5-lobed
2. Leaves variegated or coloured
3. Creeping herb; leaves ovate or heart-shaped, purplish, veins sunken; flowers white, in spikes
COROREE, LEER SAA Ga Bia od EAS te ee ood REE Sa ed Sad ae Hemigraphis alternata
3. Shrubs; leaves often mottled with coloured veins
4.. Flowers yellow, 4-5.cmJong; bracts red, broad J. . .n.cess) erento dae ae Sanchezia
4. Flowers white with purplish dots smaller bracts green, narrow ......... Pseuderanthemum
2. Leaves generally green above
5. Bracts of flowers very conspicuous, often colourful,
6. Bracts red; corollayyellow’?.:.5.... Sat a A. eee Sanchezia
6... Bracts whitish with green veins; corolla, blue.....24.5.< -.4n0.dyesenee Eranthemum nervosum
5. Bracts of flowers inconspicuous or less so, usually green,
7. Corolla + cylindric
8. Herbs or small shrubs; flowers variously coloured; stamens 4 ..................45 Ruellia
8. Shrubs; flowers white, purplish-spotted; fertile stamens 2 ......... Pseuderanthemum
7. Corolla funnel- or bell-shaped.
9. Bracts edged with long-pointed teeth or thorns; small shrubs; flowers violet-blue,
White OF VEHOW ........2. 5h ceetes «tne pitieltin yes-eaabs mee get ta oleae ake ee Barleria
9. Bracts not as above
10. Woody or herbaceous climbers, rarely erect shrub; calyx ring-shaped or 10-15-
toothed; bracts fairly large;.capsule rounded se... 2.7.05. Tee Thunbergia
10. Herbaceous climbers; calyx 5-toothed; bracts small; capsule pointed-topped ....
se Epa tiaulaSd ae.a/aic Aine lean gases A «0 giana cab Satan Semis Me Ota as ae Asystasia
1. Corolla distinctly 2-lipped
11. Leaves variegated
12. Creeping herbs; leaves green with red or white veins; flowers yellowish ................ Fittonia
12. Shrubs; leaves green or red-purple, mottled with yellow; flowers red-purple Graptophyllum
11. Leaves generally green
13. Bracts of flowers very conspicuous, often colourful
14. Spikes erect
15. Bracts orange-red; corolla light to purple-red .................. Aphelandra sinclairiana
15. Bracts white with green veins; corolla white and lilac .................. Justicia betonica
[4. Spikes curving; bractstbrick-ted>o.5 s25ad..5- seater termine nee eae Beloperone guttata
13. Bracts of flowers inconspicuous; flowers in compact clusters ................cecceee econ Jacobinia
Acanthus ebracteatus Vahl
Undershrub, 50-80 cm tall; leaves stiff, oblong, toothed, lobed and spiny, 10-16
cm long; flowers in terminal spikes, 8-10 cm long; bract ovate, 5-6 mm long,
without bracteoles; calyx-lobes ovate, free near to base; corolla 2-lipped, upper
lip obsolete, lower lip elliptic, 3-lobed, 2-2.5 cm long, white, often with bluish tip
and yellow central keel; stamens 4, pinkish. Common in tidal rivers; Jurong,
Geylang; used in local medicine. Vern. Jerujuh.
Acan. volubilis Wall.
Shrubby, twining with long internodes, unarmed; leaves lanceolate to elliptic,
7-15 cm long; spikes 6-12 cm long, narrow; bracts ovate, 5-6 cm long; corolla
white, 2-2.5 cm long. In tidal mud among grass; Kranji, Tampines.
Andrographis paniculata (Burm. f.) Nees
Herb, 40-90 cm tall, very bitter; flowers in terminal and axillary racemes,
paniculate; bracts small; calyx small, S-lobed; corolla tube straight and
narrow, 5-6 mm long, limb 2-lipped, upper lip 7-8 mm long, white with a yellow
top, lower lip 6 mm long, broad, 3-lobed, white with violet blotches; stamens 2.
Native to India and northern Malaya, cultivated as a medicinal plant, locally
naturalized.
* Based on M.C Neal, In Gardens of Hawaii pp. 777-779, (1965); rewritten.
Seed Plants of Singapore X 83
Asystasia gangetica (L.) T. Anders. (= A. coromandeliana Nees)
Ascending or trailing herb, sometimes climbing, to 1 m long; leaves ovate or
heart-shaped, 3-7.5 cm long; flowers 6-10 in spike-like inflorescence, often
arranged on one side; bracts small; calyx deeply 5-lobed, 7-9 mm long; corolla
3-3.5 cm long, light violet or white, bell-shaped, narrow at base, widened in the
upper half and at the top, segments 5, ovate, subequal; stamens 4. Native to
India and Sri Lanka, cultivated and more or less naturalized.
Asyst. nemorum Nees (= A. intrusa Bl.)
Ascending or trailing herb; leaves oblong or ovate, 3.5-10 cm long; flowers
mostly solitary; calyx (4.5-7 mm long) and corolla (2-3 cm long) smaller than the
above species. In hedges and open places, Tanglin, Fort Canning.
Barleria cristata L.
Shrub, 1-1.5 m tall; branches downy; leaves elliptic, pointed at both ends, 4-8 cm
long, hairy; flowers sessile, axillary, 1-3 together, each subtended by 2 ovate,
spiny-edged green (or white) bracts; calyx 4-lobed; corolla trumpet-shaped, 3-4.5
cm long, 5-lobed, light violet (or white); stamens 4. Native of India, sometimes
planted.
Barleria lupulina Lind.
Shrub, like the above species, but with 2 sharp spines in leaf axils; leaves
linear-lanceolate, 5-10 cm long; flowers yellow, 4-5 cm long. Native of Mauritius,
occasionally planted.
Beloperone guttata Brand. (altern. name: Calliaspidia guttata (Brand.) Bremek.)
Small shrub, to 3 m tall; leaves ovate, tip shortly pointed, 2-6 cm long, margin
entire; flowers in terminal and axillary spikes, 5-15 cm long, often slightly
curved; bracts 4-ranked, ovate-cordate, pubescent, reddish purple, 1.5-2 cm
long; calyx small, 5-lobed; corolla 2-lipped, 1.5-2.5 cm long, white, with 2 rows of
purplish spots on the 3-lobed lower lip; stamens 2. Native of Mexico, planted in
gardens.
Calophanoides quadrifaria Ridl. (= Justicia quadrifaria Wall.)
Shrubby, branched, 60-70 cm tall, glabrous; leaves narrowly lanceolate to ellip-
tic, 4-10 cm long; flowers in dense, axillary clusters; bracts small; calyx 5-lobed;
corolla small, 5-7 mm long; stamens 2. In sandy spots, Changi (Ridley s.n. in
1889).
Crossandra infundibuliformis (L.) Nees (= C. undulaefolia Salisb.)
Shrubby to 1 m tall; leaves 4 in a false whorl, oblong-ovate, 3-12 cm long, with
wavy margins; flowers in terminal, peduncled, dense spikes (later pushed aside
by new branches), 4-angled; bracts ovate oblong, 1.5 cm long; calyx 5-lobed;
corolla tube 1.5-2 cm long, limb 5-lobed, 2.5-4 cm wide, bright orange. Stamens
4. Native to India and Sri Lanka, cultivated.
Endopogon ridleyi Clarke
Branched herb, 30 cm tall; leaves opposite, elliptic, unequal-sided, tip acumin-
ate, 8-15 cm long; flowers in pairs or spikes; bracts elliptic, 1.5 cm long, green;
corolla 5-7 mm long, white. According to Ridley, it was abundant along the
stream at Stagmount (Ridley 11255), but entirely destroyed in 1910. Chua Chu
Kang, Pulau Damar.
84 Gard. Bull. Sing. 39(1) (1986)
Eranthemum nervosum (Vahl) R. Br. (= E. pulchellum Andr.)
Shrub, 1-2 m high, glabrous; leaves broadly ovate, pointed at both ends, 10-15
cm long; side veins oblique and sunken, shallowly toothed; flowers in spikes, 3-8
cm long; bracts elliptic to broadly ovate, 1.5-2.5 cm long, whitish with conspi-
cuous green venation; corolla narrowly tubular, 2-2.5 cm long, limb 5-lobed,
lobes subequal; fertile stamens 2. Native to SE. Asia, cultivated.
Fittonia verschaffeltii Coem.
Creeping herb, rooting at the nodes; leaves subsessile, oval or rounded, base
heart-shaped, 5-12.5 cm long, dark green above with a reticulation of red veins;
flowers small, 2-lipped, yellow, in a spike of 6-12 cm long. One variety, var.
argyroneura Nichols (or according to other authors, a separate species), has
smaller leaves with a network of white veins. Native of Peru, planted in pots.
Gendarussa vulgaris Nees (= Justicia gendarussa Burm. f.)
Erect shrub, branches thickened at the nodes; flowers nearly sessile, in terminal
and axillary spikes; 3-12 cm long; bracts narrow; calyx small, 5-8 mm long,
narrowly 5-lobed; corolla 2-lipped, 1.5-2 cm long, white, the lips violet-blotched
or dotted; stamens 2. Native home unknown, formerly commonly cultivated in
kampongs.
Graptophyllum pictum (L.) Griff.
Shrub, 2-3 m high; leaves broadly ovate, pointed at both ends, 10-15 cm long,
green or red-purplish with irregular yellowish blotches along the midrib; flowers
in cymes forming a terminal panicle, 3-12 cm long; bracts small; calyx 5-lobed;
corolla funnel-shaped, 2-3 cm long, 2-lipped, crimson purple; stamens 4, only 2
fertile. Native of eastern Malesia, cultivated.
Hemigraphis alternata (Burm. f.) T. Anders. (= H. colorata (Bl.) Hall. (f.)
Creeping herb, purplish, rooting at the nodes; leaves ovate to heart-shaped,
2.5-8 cm long, purplish, veins sunken, blunt-toothed; flowers in narrow spikes,
2-10 cm long, pedunculate; bracts dark purple, 1-1.5 cm long, slightly exceeding
the calyx; corolla funnel-shaped, 1.5-2 cm long, white, purple-lined, limb 5-
lobed, subequal. Native to eastern Malesia, cultivated for ornament and for
medicine.
Hygrophila erecta (Burm. f.) Hochr. (= H. quadrivalvis Nees)
Herb, 1-1.2 m tall; stem 4-angled; leaves oblong-obovate, tip round, 3-5 cm long;
flowers few, in axillary clusters; calyx shallowly 5-lobed; corolla funnel-shaped,
1-1.2 cm long, 2-lipped, the tube and the upper lip white, lower lip violet edged
and purple-dotted; stamens 4. In wet spots; Bukit Timah, Ang Mo Kio.
Hygr. meianthos Clarke
Ascending herb, 30 cm tall; leaves obovate, tip obtuse, 1.5-3 cm long; flowers
few, in axillary clusters; calyx white hairy; corolla 1 cm long, white, edged violet.
In wet grassland, rare; Botanic Gardens. (Probably merely a variety of the above
species).
Hygr. phlomoides Nees
Herb, 1 m tall; branches soft hairy; leaves obovate, tip blunt, 2.5-10 cm long;
flowers many, in axillary clusters, bract lanceolate; corolla violet, lower lip
darker. In wet places, edge of Garden’s lake, Changi (Ridley s.n. in 1891), Water
Catchment Areas.
Seed Plants of Singapore X 8
(Nn
Hygr. spinosa T. Anders.
Herb, 0.5-.15 m tall, with axillary spines (2.5 cm long) leaves linear to lanceolate.
7-14 cm long, sessile: flowers in large clusters; bracts lanceolate; calyx lobed
nearly to the base; corolla 2 cm long, pale purple. Native to India, an accidental
weed.
Jacobinia carnea (Lindl.) Nichols (altern. name, Cyrtanthera carnea Bremek.)
Shrub, 0.5-2 m tall: leaves ovate-oblong, 12-25 cm long; panicles subsessile.
erect, 12-20 cm long; bracts narrow, 1-2.5 cm long; calyx deeply 5-lobed; corolla
tubular, deeply 2-lipped, 6-7 cm long, pink. Native of Brazil, cultivated.
Jacob. coccinea (Aubl.) Hiern. (altern. name, Pachystachys coccinea Nees)
Shrub, 1-2 m tall; leaves oblong or elliptic, 12-27 cm long, glabrous; flowers in
dense terminal spikes; bracts ovate, green, hairy, 2-3 cm long; corolla tubulate.
laterally compressed, 2-3.5 cm long; 2-lipped, bright red, pubescent. Native to
French Guyana, cultivated and escaped.
Justicia betonica L. (= Nicoteba betonica (L.) Lindau)
Shrubby, 1-2 m tall: leaves narrowly ovate, pointed, 5-7 cm long; flowers in
terminal spikes, 8-10 cm long, often branched; bracts conspicuous, ovate, white
with green nerves, 3 bracts subtending one flower; corolla 1-1.5 cm long, white or
lilac. Native of Mexico, planted.
Justicia vasculosa Wall.
Straggling herb, 20-30 cm tall; leaves opposite; subequal or very unequal, lan-
ceolate to ovate, 10-12 cm long; flowers in terminal and axillary spikes, 5-10 cm
long, bracts not larger than calyx; calyx deeply 5-lobed, lobes lanceolate,
hairy; corolla 1.5 cm long, yellow, spotted with pink in the mouth, pubescent. In
forests; Bukit Panjang, Bukit Mandai (Ridley s.n. in 1889), Chua Chu Kang.
Peristrophe acuminata Nees
Herb, | m tall; leaves lanceolate, 6-7 cm long, narrowed at both ends; flowers in
terminal and axillary short cymes, briefly peduncled; bracts linear, 1-1.5 cm long:
corolla tubular, 1.5 cm long; tube white, limb 2-lipped, purple with a creamy
blotch and many purple spots in the mouth. In waste ground and roadsides:
Tanglin, Chua Chu Kang, Botanic Gardens (Purseglove 4041).
Peris. roxburghiana (Schult.) Bremek. (= P. tinctoria Nees)
Herb, 1-1.2 m tall; leaves elliptic-ovate, 5-8 cm long, narrowed at both ends;
peduncles 2-3 cm long, pubescent; bracts ovate, 2-4 cm long; corolla 3-4 cm long.
tube white, limb bright purple with a curved creamy blotch in the mouth. In
shade spots, usually near cultivated land; Changi, Blakang Mati (Ridley s.n. in
1892).
Pseuderanthemum kingii (Clarke) Rid.
Shrubby, slender, 30-40 cm tall; leaves elliptic, 5-7 cm long, acuminate; flowers
in pairs on terminal spikes, 6-7 cm long, bracts linear, small; calyx-lobes linear,
nearly free to the base; corolla cylindric, 5-lobed, 2-3.5 cm long, white, not
2-lipped. In forests; Changi, Seletar (Ridley 3776).
86 Gard. Bull. Sing. 39(1) (1986)
Pseud. reticulatum Radlk.
Shrub, | m tall; leaves leathery, ovate or rounded, 3-10 cm long, yellow and pale
green with or without yellow bands and blotches; corolla 2-3.5 cm long, the lobes
acute, white with purple spots. native of Melanesia, planted as an ornamental.
Ruellia amoena Nees (Altern. name, Stephanophysum longifolium Poir.)
Erect herb; leaves opposite, ovate-oblong, 6-15 cm long, acute, shallowly tooth-
ed; flowers in axillary long-peduncled (2-10 cm long) cymes; calyx small, 0.7-1
cm long; corolla 3-3.5 cm long, bright red, round-lobed. Native of Brazil,
cultivated as an ornamental.
Ruel. repens L. (altern. name, Dipteracanthus repens Hassk.)
Herb, erect or ascending, 20-70 cm tall; leaves narrowly lanceolate, 3-10 cm long;
flower solitary, axillary, bracteoles leaf-like, ovate, 1-2 cm long; corolla bell-
shaped, violet, with 5 rounded lobes. In grassland, Botanic Gardens (Hullett s.n.
in 1884).
Ruel. tuberosa L.
Spreading or ascending herb, roots tuberous; leaves opposite, oblong or ovate,
6-18 cm long; flowers in axillary cymes, 1-many-flowered; bracts narrow; calyx
2-3 cm long; narrowly 5-lobed; corolla trumpet-shaped, 4-6 cm long; limb violet,
sometimes pink or white, 3-5 cm across. Native to W. Indies, planted.
Sanchezia nobilis Hook. f.
Shrub, 1-2 m tall; branches 4-angled, glabrous; leaves oblong, 10-25 cm long (one
variety often with a white midrib), petioles winged; flowers in terminal spikes,
often branched, bracts ovate, reddish, 3-5 cm long; corolla 4-5 cm long, bright
yellow. Native of tropical America, cultivated.
Staurogyne griffithiana O. Ktze.
Creeping or ascending herb, 10-20 cm long; leaves ovate or broadly elliptic, both
ends rounded; flowers in terminal, simple or branched racemes, 5-12 cm long;
bracts broader than calyx; calyx-lobes linear, one larger than the others; corolla
cylindric, 1-1.5 cm long, white. In forests; Bukit Timah (Hullett 613), Kranji,
Sungei Buluh.
Staur. setigera O. Ktze.
Ascending herb, 1-25 cm long, pubescent; leaves lanceolate to ovate, 4-6 cm
long; flowers in terminal dense racemes, 5-10 cm long; corolla 1.5 cm long, white
with pink spots in the mouth. In forests; Bukit Timah (Ridley s.n. in 1890), Chua
Chu Kang, Jurong.
Strobilanthes dyerianus Mast. (altern. name, Perilepta dyeriana Bremek.)
Herb, to 1 m tall; leaves opposite, of the same pair very unequal, larger ones
10-20 cm long, smaller ones much reduced; flowers in terminal and axillary
spikes; bracts about the same size as calyx; calyx 2-lipped; corolla bluish violet,
2.5-3 cm long. Native of Burma, formerly planted.
Thunbergia affinis S. Moore
Erect shrub, 1-2 m tall; leaves ovate or rhomboid, 4-11 cm long; flowers axillary,
solitary or in pairs; bracteoles persistent; corolla dark violet, tube 4-5 cm long,
limb 5-6 cm across. Native to tropical E. Africa, sometimes cultivated.
Seed Plants of Singapore X 87
Thunb. alata Boj. ex Sims
Slender climber; leaves ovate or cordate, 3-5 or more cm long; petioles winged;
flowers axillary, solitary; calyx with 11-14 teeth; corolla 3.5-5 cm across, orange-
yellow, with (or without) a dark purple mouth, (thus called ‘Black-eyed-susan’),
rarely white. Native of tropical Africa; in waste places.
Thunb. erecta T. Anders
Erect shrub, 1-2 m tall; leaves ovate-rhomboid, narrowed above, 2-9 cm long;
flowers axillary, solitary or in pairs; bracteoles deciduous; corolla dark violet or
white, tube 0.5 cm long; limb 3.5-5 cm across. Native of tropical W. Africa;
sometimes cultivated.
Thunb. fragrans Roxb.
Slender climber, glabrous; leaves oblong or elliptic, 5-7 cm long; base rounded or
hastate; flowers axillary, solitary or in pairs; calyx with 12 teeth; corolla 4-5 cm
across, white. Native to India, Java and Australia; an escape from gardens.
Thunb. grandiflora Roxb.
Woody climber, pubescent; leaves cordate, 5-7 nerved at base, 7-18 cm long,
shallowly to deeply palmate-lobed; flowers raceme-like, terminal; corolla-tube
3-5 cm long; limb blue-violet or rarely white. Native to SE. Asia; planted as an
ornamental, sometimes on overhead bridges, more or less naturalized here.
Thunb. laurifolia Lindl.
Woody climber, glabrous; leaves ovate, lower leaves broad at base and often
toothed, upper ones narrower and entire, 3-5 nerved at base, 7-12 cm long;
flowers in terminal racemes; calyx a mere rim; corolla-tube 3-4 cm long; limb
dark blue. Native to SE. Asia, sometimes cultivated.
141. VERBENACEAE
Key to the genera
1. Inflorescences + head-like
2. Woody
3. Shrubs, erect or scrambling; inflorescence flat-topped, not surrounded by large bracts ........
a A SR i ee Re Oe ie 6 sn Ww gnc «SASH AME c's oo pcp csscc sy gna ae Lantana
3. Woody climbers; inflorescence surrounded by large, colourful bracts
IRE Se SENN AIR fo So Fe OcisVsifai eileen ty shivb awe owecesvsntonessnesevnsuveones Congea
Aimee GO COLMA CYNMOTIC JS-GAOIE oak. 5.5 «ois desc w awl eceweneswhesdedededen Sphenodesma
2. Herbaceous
a oe OI SDR Ie ce new 0 5 cig ee ores s onyunae ips tease d-dinn Verbena
ee I RNR Bs RE ra i, fon ass ewgin tins ogc caress sacs bcecescemmereeessnaes Phyla
1. Inflorescence not head-like
6. Inflorescence spike- or raceme-like
7. Inflorescence spike-like, simple or branched; flowers sessile or nearly so
8. Trees, on muddy shores or in mangroves, with peg-like breathing roots = Avicennia
8. Not as above
Pe ME, WUMRTMPNCAEY,. 25 aA ss os cso sc ees bu bncpecvewcescovadestacenby cavers Citharexylum
PEE (ESCM WOON 88s FSGS, cei ersten hee ceed Stachytarpheta
7. Inflorescence raceme-like; flowers stalked; creeping or climbing shrubs
10. Leaves thin and smooth; calyx of 5 green teeth; fruit a yellow berry .......... Duranta
10. Leaves thick and downy, sandpaper-like; calyx of 5 large, star-shaped, colourful
SMES SEAREE GEV OMMINRNE deo, che iv cece ccps vaserc ens anise caves cpesusbeatnes cas sevansedeateeres Petrea
88 Gard. Bull. Sing. 39(1) (1986)
6. Inflorescence cymose-paniculate
11. Corolla regular or nearly so, not 2-lipped; stamens exerted
12. Trees; leaves large, 30-GDicm lome:. 2... 5. 5. £45: ue arch cienacrsaveds oa baane ae Tectona
12. Shrubs
13. Cymes axillary, shorter than the leaves; calyx tubular, small; flowers 4-merous .......
sec ne neces cee clelad URN Ue UO es hs Rita STE ICL dA de SOMA Alek ae fe Callicarpa
13. Cymes axillary and terminal, often in terminal panicles; flowers 5-merous
14. ° Calyx of flower bell-shaped ...........4.c.-:100s0000-.qeapeeee eee ae Clerodendrum
14- (Calyx of Mower saticer-shaped _..............---2..428- eB org ae Pe Holmskioldia
11. Corolla distinctly 2-lipped; stamens included
15. Leaves simple, entire or toothed
16.)\Flowers large, corolla 2-2:S:em long. <7, 2... 0.0095... Bi ee Gmelina
16. Flowers smaller, corolla less than 0.5 cm long .....:......2333.e2)92-ebe eee Premna
15. Leaves palmately compound, with 3-5 leaflets (or sometimes reduced to one leaflet in V.
trifolia)
17. Petioles and petiolules strongly swollen towards the apex ...... Teijsmanniodendron
V7. Not asabove. ..5. 00h. 0c. een th et a ah ee eee Vitex
Avicennia alba BI.
Tree, to 20 m tall; leaves simple, opposite, lanceolate, 8-12 cm long, pointed,
lower surface greyish white; flowers small, in branched spikes; corolla tubular,
orange, 4-5 mm long, 4-lobed; fruit lanceolate-elliptic, leathery, 1-1.5 cm long,
pointed; seedlings semi-viviparous (i.e., fruit dehiscing by 2 valves, the emerging
seedling consisting of (1) 2 large, green, subequal cotyledons conduplicately
arranged, (2) a fully developed plumule and (3) a stout radicle with tangling
hairs). Common in mangrove and along tidal river. Tuas (Ridley 6312). Vern.
Api api.
Avic. lanata Ridley
Young branches and lower leaf-surfaces densely covered with brown hairs;
leaves ovate or elliptic, 8-10 cm long; fruit broadly ovoid, 1.5-1.8 cm long. In
mangroves on sandy soils, Pandan Nature Reserve (Jsmail 54). Vern. Api api.
Avic. officinalis Bl.
Leaves oblong-elliptic, 8-10 cm long, round-tipped; flowers in clusters; fruit
ovoid, slightly beaked, 3-3.5 cm long, velvety. Common in mangrove and along
tidal rivers. Changi, River Valley Road (Burkill 3792), Pulau Ubin. A fourth
species, called A. intermedia Griff. [? = A. marina (Forsk.) Vierch.] is somewhat
intermediate between A. alba (with the lower leaf surface glaucous) and A.
officinalis (with round-tipped elliptic leaves) but differing from both in its 4-
angled stem. Tuas, Pulau Sudong (Womersley 61).
Callicarpa longifolia Lam.
Straggling shrub; 2-3 m tall; leaves lanceolate, 7-15 cm long, stellately hairy
beneath; flowers many in cymes; corolla tubular, liliac. Formerly common in
hedges in Tanglin, Bukit Timah (Ridley 2786) and Changi, now rare.
Citharexylum spinosum L. (= C. quadrangulare Jacq.)
A shrub (in country of origin, a small tree); branches 4-angled; leaves opposite or
in 3s, oblong or oval, 15-20 cm long, smooth, pointed; flowers in narrow
clustered spikes, 10-20 cm long; corolla tubular, white 5-lobed, curved, less than
1 cm long. Native to the West Indies, known as ‘Fiddlewood’, sometimes
cultivated in gardens.
Seed Plants of Singapore X 89
Clerodendrum deflexum Wall.
Small shrub, 1 m tall; leaves opposite, lanceolate or elliptic, 15-30 cm long:
flowers in nodding heads, with red bracts: corolla tubular, white, 2-2.5 cm long:
drupe purple to black. Common in woods; Gardens’ Jungle, Tanglin, Chua Chu
Kang (Goodenough 2796), Tampinis.
Cler. indicum (L.) Ktze (= C. siphonanthus R. Br.)
Shrub; leaves narrowly oblong, 10-12 cm long, in whorls; corolla white, night-
blooming, fragrant; fruit purplish with red calyx. Native of SE. Asia, cultivated
for its fragrant flowers.
Cler. inerme (L.) Gaertn.
Small shrub, about 1 m tall; leaves elliptic, narrowed at both ends; 4-8 cm long;
flowers in axillary small cymes; corolla cylindric, white, 2-2.5 cm long; stamens
exsert, dark purple. Common near the sea in tidal mud; Rochore, Changi
(Hullett 625), Seletar, Pulau Ubin.
Cler. laevifolium Bl. (= C. disparifolium B1.)
Shrub or small slender tree, 5-7 m tall; leaves oblong or lanceolate-elliptic, 5-15
cm long; flowers in slender cymes; corolla pale yellow; fruit black with red calyx.
Common in woods; Tanglin, Gardens’ Jungle, Bukit Timah, Changi. Vern.
Guriam padang.
Cler. nutans Jack (= C. penduliflorum Wall.)
Shrub, less than 1 m tall; leaves ovate-lanceolate, pointed, 12-20 cm long:
panicles pendent; corolla funnel-shaped, white, 1-1.5 cm long. Native of N.
Malaya, occasionally cultivated.
Cler. paniculatum L.
Shrub, 3-4 m tall; leaves heart-shaped, shallowly 3-5 lobed, 15-20 cm long:
flowers in large terminal panicles; corolla scarlet, 2-3 cm long. Native of Con-
tinental Asia, cultivated for the showy inflorescences.
Cler. philippinum Schauer. (= C. fragrans Willd.)
Erect shrub, with root-suckers; leaves broadly ovate, hairy, 8-20 cm long; flowers
in dense corymbs; corolla white, 2-3 cm long. Native to Java, cultivated as
ornamental. There is a double-flowered variety, var. pleniflorum Schauer.
Cler. phyllomega Steud. var. myrmecophilum (Ridl.) Moldenke (= C. myrme-
cophilum Ridl.)
Slender shrub, usually unbranched; stem and petioles hollow and tenanted by
ants; leaves oblong or lanceolate, 15-30 cm long; panicles large, terminal; corolla
cylindric, 1-1.5 cm long, 5-lobed, orange-red; stamens long exsert, dark pink. In
muddy wet spots in forests, formerly found at Chua Chu Kang (Ridley 6700) and
on Pulau Damar.
Cler. thomsonae Balf.t.
Climber; leaves ovate to heart-shaped, 8-12 cm long: calyx white; corolla red
(thus ‘Bleeding Heart’). Native of W. Africa, often cultivated.
90 Gard. Bull. Sing. 39(1) (1986)
Cler. villosum BI.
Shrubs, 2-3 m tall, hairy; leaves ovate-cordate, 10-20 cm long; flowers in terminal
panicles; corolla white, the tube less than 1 cm long; fruit black, with enlarged
white calyx. In waste land, Chua Chu Kang (Goodenough 2795).
Congea velutina Wight
A climber, all parts covered with greyish woolly hairs; leaves ovate-elliptic,
thick, shortly pointed, 3-15 cm long; panicles much branched, formed by 5-7-
small-flowered heads; flowers pink, inconspicuous, the 3-4 bracts subtending the
flower-heads spathula-shaped, 2-3.5 cm long, lilac. Native to India and Malaya,
cultivated as ornamental.
Duranta repens L.
Creeping shrub with slender, drooping branches; leaves opposite or in whorls,
ovate, 3-8 cm long; racemes terminal and axillary, forming panicles; corolla
tubular, light blue or white, less than 1 cm long; globose, drupe bright (thus
‘Golden Dewdrop’), less than 1 cm across. Native of tropical America, culti-
vated or escaped. 4 #8 {é.
Gmelina asiatica L.
Small bushy tree or climber, with axillary spines; leaves simple, opposite, obo-
vate or rhomboid, 2-4 cm long, often 3-lobed, glaucous beneath; flowers in
cymose-racemes on a terminal panicle; corolla obliquely funnel-shaped, yellow,
3-4 cm long; drupe pear-shaped, 2-3 cm across. On waste grounds; Geylang.
Gmel. elliptica. J. E. Sm. (= G. villosa Roxb.)
Spiny shrub: leaves ovate, acute, 2.5-7 cm long, densely hairy underneath;
corolla yellow; drupe subglobose, 2-2.5 cm across, yellow. Formerly collected at
Chua Chu Kang and Geylang. Vern. Bulang.
Gmel. philippinensis Cham. (= G. hystrix Schultes)
Scandent shrub, spiny; leaves elliptic, blunt, 8-10 cm long; flowers in pendent
clusters, 6-8 cm long, with dark red, ovate bracts; corolla yellow. Tanglin,
Gardens’ Jungle. Fruit used in local medicine.
Holmskioldia sanguinea Retz.
Shrub, less than 3 m tall; leaves opposite, ovate, pointed, 2.5-8 cm long; flowers
in small clusters, both axillary and on branch tips; calyx red or orange, saucer-
shaped; corolla funnel-shaped, curved, 5-lobed, 2-2.5 cm long, red (thus ‘Cup-
and-saucer Plant’). Native of Himalayan region, occasionally planted.
Lantana camara L. (incl. L. aculeata L.)
A shrub, branches prickly (var. aculeata Mold.) or unarmed; leaves opposite,
rarely 3 in a whorl, ovate, 5-12 cm long, pungent-scented; flowers in head-like
dense spikes; corolla salver-shaped, curved, 1-1.2 cm long, orange, pink, red, or
variegated; drupe bluish. Native to tropical America, originally a garden plant,
later growing wild in waste places, several cultivated varieties with an array of
flower colours. FER Ff, )
Peronema canescens Jack
Small tree, to 14 m tall, with 4-angled branches; leaves opposite, pinnate, 30-90
cm long; leaflets 4-10 pairs, with or without a terminal leaflet, sessile, 10-30 cm
Seed Plants of Singapore X 9]
long; terminal panicles 30-60 cm long; corolla small (0.3 cm wide), 2-lipped,
greenish white; capsule round, 3 mm broad. In secondary forests, formerly found
in Bukit Mandai, now occasionally cultivated.
Petrea volubilis L.
Woody climber; leaves opposite, oval, pointed, 5-15 cm long, thick and downy;
racemes hanging from branch tips; calyx 5-lobed, accrescent and persistent,
spreading, reaching 3-4 cm across, bluish; corolla funnel-shaped, violet. Native
to tropical America often planted in gardens.
Phyla nodiflora (L) Greene (= Lippia nodiflora A. Rich.)
Creeping perennial herb, rooting from the nodes, densely covered with greyish
soft hairs; leaves opposite, obovate, 2-5 cm long; dense spikes erect, axillary,
with prominent bracts; corolla small (2-3 mm long), white and then lilac with a
dark eye. In dry sandy waste places.
Premna corymbosa Rottl. & Willd. (= P. angustior Ridl., P. integrifolia L.)
Low shrub, sometimes creeping; leaves simple, oblong-ovate, 3-10 cm long:
panicles corymbose, terminal; corolla small (2-3 mm wide), white, 2-lipped;
drupe ovoid, black. On sandy places near the sea. Changi (Ridley 2785), Loyang,
Chua Chu Kang.
Prem. foetida Reinw. ex BI.
Large shrub, to 6 m tall, hairy; leaves ovate, acute, 8-15 cm long; corymbs 8-12
cm long and wide; corolla greenish white. In damp low-lying ground, leaves
served as a pot herb; Tanglin, Thomson Road (Hullett 395).
Prem. parasitica Bl. (= P. trichostoma Miq.)
Large climbing shrub, glabrous; leaves oblong-obovate to nearly orbicular,
nerves 5-6 pairs; corymbs 10-12 cm across. In woods; Chua Chu Kang (Ridley
6828).
Prem. punctulata Clarke
Scandent shrub; leaves elliptic or broadly oblong, rigid, nerves 4-5 pairs, sparsely
hairy, 10-15 cm long; corymbs pubescent, branched. Once found in the former
Economic Gardens in 1915 (Ridley) where it has since disappeared.
Prem. ridleyi K. & G.
Climbing shrub; leaves ovate, or oblong, blunt, subleathery, 4-8 cm long;
corymbs 5-12 cm across. Climbing on trees in woods, only one plant was found in
the Gardens’ Jungle in 1896 (Ridley 6826, photo).
Sphenodesma pentandra Jack
Climbing shrub, pubescent; leaves lanceolate or ovate, acute, 5-15 cm long;
raceme-like inflorescences consisting of many pairs of heads; heads small, 5-7-
flowered, surrounded by 6 large, oblong, 2-3 cm long bracts; corolla bluish. On
edge of woods; Seletar, Changi. Vern. Akar sulang.
Stachytarpheta indica (L.) Vahl
Erect herb, to 1 m tall; leaves opposite, bright green, elliptic oblong, 4-11 cm
long, margin toothed, the teeth of very different sizes, secondary veins inconspi-
92 Gard. Bull. Sing. 39(1) (1986)
cuous beneath; spikes 15-40 cm long; corolla bright violet, sometimes pale violet
or white. Common on seashores.
Stachy. jamaicensis (L.) Vahl
Herb or shrubby, 2 m tall, glabrous; leaves dark green, ovate or oval, 3-8 cm
long, toothed, the teeth more or less of same size, secondary veins prominent
beneath; terminal spikes 10-20 cm long; corolla tubular, blue, less than 1 cm
long. Native to tropical America, a common weed on waste ground.
Stachy. mutabilis (Jack.) Vahl
Differs from S. jamaicensis in being hairy throughout, in having a thicker spike,
and larger, crimson to pink flowers. Also from tropical America; an ornamental
and a garden escape.
Tectona grandis L.f.
Large deciduous tree to 50 m tall; leaves opposite or in threes, oval, 30-60 cm
long; panicles large, terminal, 30-90 cm long; flowers small; calyx bell-shaped,
much enlarged in fruit; corolla funnel-shaped, white or reddish, 1 cm across;
drupe globose, 1-1.5 cm across. Native of monsoon forests of E. India to
Thailand and Java, valued for its durable wood (teak), occasionally planted.
Teijsmanniodendron coriaceum (Clarke) Kosterm. (= Vitex coriacea Clarke)
Tree; leaves 3-foliate; leaflets leathery, oblong to elliptic, 7-12 cm long, stalks
swollen at the base; flowers in paniculate cymes; corolla violet, 2-lipped; drupe
globose to oblong, 1-seeded. In forests; Bukit Timah, Gardens’ Jungle.
Teijsm. pteropodum (Miq.) Bakh. (= Vitex pteropoda Miq.)
Small tree; leaves palmately compound, petiole 10-15 cm long, broadly winged;
leaflets 5-7, elliptic to oblong, 10-50 cm long; panicles terminal; corolla purplish;
drupe ovoid. In swampy forests; formerly found at Chua Chu Kang, Pulau
Damar.
Verbena hybrida Voss.
Annual herb; leaves oblong, blunt-toothed, 4-8 cm long; flowers in broad heads;
corolla red, pink or yellow or variegated, often with a white eye. A hybrid of V.
peruviana Druce with other S. American species, sometimes cultivated.
Verb. tenuisecta Briq. (= V. tenera Spreng.)
Perennial herb; stems creeping; leaves 1.5-2.5 cm long, twice pinnatifid, seg-
ments linear; flowers in dense heads; corolla violet, blue or white. An ornamen-
tal, native to S. America.
Vitex negundo L.
Shrub; leaves palmate; leaflets 3-5, elliptic, the mid-leaflet distinctly stalked;
flowers in terminal panicles; corolla pale to deep blue, with a yellow curved mark
on lower lip. A medicinal plant for various ailments; in wasteland, probably
introduced. #F#ij.
Vit. pinnata L. (= V. pubescens Vahl)
Hairy tree to 10 m tall; leaves palmately compound; leaflets usually 5, almost
sessile, the mid-leaflet the largest, about 7.5-20 cm long; panicles terminal;
Seed Plants of Singapore X 93
corolla violet blue; fruit globose, black, flattened. Common in open wasteland
and in secondary forests.
Vitex trifolia L.
White hairy shrub; branches drooping; leaves 3-foliate, rarely simple (var. re-
pens); leaflets 3-7.5 cm long, mid-leaflet almost sessile; corolla pale blue; fruit
oblong. In gardens and villages near the sea; Pasir Panjang, Pulau Ubin.
Vit. vestita Wall. ex Schau.
Small tree; branches pubescent; leaves 3-foliate; leaflets elliptic, long-acuminate,
8-18 cm long, mid-leaflet the largest; corolla cylindric, 1-1.5 cm long, 2-lipped,
yellow. In inland forests; Bukit Timah, Jurong.
142. LABIATAE
Key to the genera
1. Flowers mostly in dense axillary (sometimes also terminal) clusters
2. Leaves finely.dissected intoi/lobes.and narrow. sepments 5... .0h;,..0.5- hn de el ee teeees. Leonurus
2. Leaves entire or toothed, not dissected
3. Calyx 6-10-toothed
Br RCT oe Cred PONS, OFAlipe OF FEC «ste ha te he Se nn echoes vik da enn £0 hae oa 4 SR Cat Leonotis
ae eeemile less than iS Cit tome Pouiite Gr mish 2 282, eee oe eee eeeceseseeses Leucas
3. Calyx 4-5-toothed
5. Flowers in dense ball-like heads or in clusters; corolla 2-lipped, the upper lip bent down-
SETS La a ae aE SSO Ee a gS he eee ee pe Hyptis
5. Flowers in axillary clusters; corolla subequally 4-lobed; cultivated ...................... Mentha
1. Flowers generally in terminal spike- or raceme-like inflorescences, simple or branched
6. Spike-like inflorescence densely packed with very tiny (less than 3 mm long), bluish flowers ........
Suan on a Rae ea Ry: PED an ee, 1 ROP ws Lay eR SSR Se sek ag ah ces daehin).. oo BoB ijm Lew sge svn dies Pogostemon
6. Spike- or raceme-like inflorescence clearly in whorls; flowers larger (over 5 mm long, usually
much longer)
Pere aay GEA WE PAB, OAOUCE: WEES 6.609.182 e.5 oee co res wcs odes ocln ssa le scch vases geentneess Anisomeles
7. Calyx irregular, generally 2-lipped; generally cultivated
8. Stamens 2, anther-cells widely separated on a long slender connective ................... Salvia
8. Stamens 4, anther-cells parallel, not separated,
9. Stamens coiled in bud, projecting 1.5-2 cm beyond the corolla ................ Orthosiphon
9. Stamens not as above
10. Lower corolla-lip nearly flat; plants strongly aromatic .....................00000 Ocimum
10. Lower corolla-lip boat-shaped; plant not or only faintly aromatic Plectranthus (incl.
Coleus)
Anisomeles indica (L.) OK. (= A. ovata R. Br.)
Herb or shrubby, to 1.5 m high, more or less hairy; leaves ovate, 4.5-6 cm long;
flowers in dense false whorls in a spurious spike; corolla tubular, 2-lipped, 7-8
mm long, white or violet. In open and waste places, formerly found at Tanglin
(Ridley 2692), now probably extinct.
Hyptis brevipes Poit.
Erect herb, to 1 m high; leaves narrowly lanceolate to oblong-ovate, 4-8 cm long;
flowers in spurious heads, 1-1.2 cm across in fruit; corolla white, 4-5 mm long,
2-lipped. Native to S. America, more or less established.
Hypt. capitata Jacq.
Annual herb, 1-1.5 cm high; leaves lanceolate or rhomboid elliptic, 6-12 cm long;
flowers in axillary globular spurious heads, 1.5-2 cm across in fruit; corolla 5-6
94 Gard. Bull. Sing. 39(1) (1986)
mm long, white. Native to S. America, a weed in waste places.
Hypt. suaveolens (L.) Poit.
Branched herb, to 1.5 m high, strongly aromatic; leaves ovate to broadly obo-
vate, 3-5 cm long, pubescent; flowers in 2-5 flowered cymes, on one side of a
spurious raceme; corolla blue, tubular, 6-8 mm long. Introduced from S. Amer-
ica, One time very common in waste grounds.
Leonotis nepetaefolia (L.) R. Br.
Erect herb, to 2 m tall; leaves oblong to ovate, 4-12 cm long, toothed; flowers in
dense axillary spurious whorls; corolla bright orange or red, 2-2.5 cm long.
Native of tropical Africa, occasionally escaped from cultivation, a weed.
Leonurus sibiricus L.
Herb, to 1.5 m tall, leaves linear to ovate, palmate-pinnately dissected, 4-7 cm
long, flowers in dense axillary spurious whorls; corolla white, about 1 cm long, 2
lipped. A native of E. Asia; cultivated as an ornamental or for medicinal
purposes, occasionally found in wasteland. Vern. Seranting, 4 BEE.
Leucas lavandulifolia J. E. Sm. (= L Jinifolia Spreng.)
Annual herb, less than 1 m tall; leaves linear, to lanceolate, 4-6 cm long; flowers
in terminal and axillary leafy false whorls, often congested towards apex and
forming clusters; calxy oblique, 10-toothed; corolla tubular, 1 cm long. 2-lipped.
In open places, rare. Mt Faber (Ridley 3888).
Leucas zeylanica (L.) R. Br.
Annual, to 60 cm high, hairy; leaves lanceolate, 4-5.5 cm long; flowers in
terminal spurious whorls, usually 6-8 whorls forming a globular head, 1.5-2 cm
across; corolla white, tubular, 8-10 mm long, 2-lipped. In open grassland. Pasir
Panjang (Mat. s.n. in 1894). Vern. Ketumbit.
Mentha arvensis L.
Aromatic perennial, often prostrate, variously. pubescent; leaves ovate, 2-4.5 cm
long, weakly toothed. A native of the northern hemisphere, a small-leaved form
sometimes cultivated in pots, no flowers have been observed. Another related
species, M. spicata L., an erect or ascending herb of 0.5-1 m tall, with dense
axillary clusters of small pinkish flowers, of the northern temperate countries, 1s
occasionally planted. Vern. Pokok kepari, 7& faj.
Ocimum americanum L. (= O. canum Sims)
Aromatic herb, erect, 30-50 cm high; leaves lanceolate to elliptic, 2.5-5 cm long;
false whorls in spurious racemes, usually branched, 7-15 cm long; pedicels much
shorter than calyx, 2 lower calyx-teeth slightly longer than the upper one; corolla
white, 4-6 mm long. In settled areas, sometimes cultivated.
Ocim. basilicum L.
Like O. americanum, except flowers are larger (corolla 7-12 mm long, white or
pinkish). Often cultivated. Vern, Basil, Selaseh.
Ocim. tenuiflorum L.
Like O. americanum, except the two lower calyx-teeth equalling the upper tooth
and less aromatic. A pantropical weed occasionally found in waste places or in
settled areas.
Seed Plants of Singapore X 95
Orthosiphon aristatus (Bl.) Miq. (= O. stamineus Benth.)
Slender ascending herb. 30-60 cm high: leaves ovate or rhomboid, 3-9 cm long:
terminal spurious racemes 10-15 cm long: corolla pinkish or white. tubular. 10-12
mm long. 2-lipped: stamens coiled in bud, projecting about 2 cm beyond the
corolla throat. Native to SE. Asia, sometimes cultivated as an ornamental or
medicinal plant. Vern. Kumis kuching, 34% B.
Plectranthus scutellarioides (L.) R. Br. (= Coleus atropurpureus Benth... C. blumei
Benth. )
Branched herb, erect or ascending: leaves very variable in size, shape and
colouring, generally ovate in outline. 4-7 cm long, toothed; usually in 6-flower-
ed cymes, forming spurious racemes or panicles; corolla boat-shaped, 8-13 mm
long, bright purple. Formerly occurred in open places (Jurong, Ridley s.n. in
1890), now in cultivation only.
Plectr. rotundifolius (Poir.) Spreng. (= Coleus tuberosus Benth., C. parviflorus
Benth.)
Herb, pubescent; lower stem creeping; older roots often swollen into dark brown
tubers, 2-4 cm long; leaves thick, juicy, ovate or suborbicular, 2-5 cm long.
Native of India, sometimes cultivated for the edible roots.
Pogostemon auricularius (L.) Hassk. (= Dysophylla auricularia (L.) Benth.)
Annual herb, 30-70 cm high, pubescent; leaves narrowly ovate, 4-6 cm long:
flowers in dense spurious terminal spikes, hairy: corolla very small, 2-2.5 mm
long. Formerly common in open wet spots, Tanglin, Changi (Hullett 375); now
probably extinct. Vern. Ekor kuching.
Salvia coccinea Juss. ex Murr.
Slender herb, to 1 m high, hairy or not; leaves ovate or deltoid-ovate, 2.5-3.5 cm
long; false racemes terminal, branched or not: flowers 6-10 in a false whorl:
corolla bright scarlet, 2-2.5 cm long, the tube straight, 2-lipped. Native of
tropical America, sometimes cultivated from imported seeds. —347.
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The Evaluation of the Effectiveness of Water Absorbent Polymers in
Reducing the Irrigation Needs of Potted Plants
T.W. FOONG & C.N. YANG
Botanic Gardens, Singapore
Abstract
The available-water-enhancing effects of the water-absorbent polymers, Agrosoke, Soiltex G1 and
Terra-Sorb, on a typical local sandy-clay topsoil were evaluated. using Mussaenda Dona Luz, Tecomaria
capensis, and Hibiscus rosa-chinensis as indicator plants. Results did not indicate significant beneficial
effects at the three rates of polymers used. Instead, the practicability of using sludge to augment the
water-holding capacity of the topsoil was validated.
Introduction
Synthetic polymeric soil conditioners are increasingly being used as modifiers of
physical soil attributes such as water-holding capacity and permeability, with
promising results (Azzam, 1985; Callebaut et al, 1979: De Boodt,, 1979 and Gunn,
1984). Many of the synthetic soil conditioners are polyacrylamide-based, having the
following basic structure.
CH, Preacher Cl CH,
C=O C=O Like
NH, Mu, NH,
An interesting property of these polymeric soil conditioners is their ability to hold
many times their own weight in plant available water (Flannery & Busscher, 1982:
Gehring & Lewis III, 1980 and Hamilton & Lowe, 1982). Moreover, rehydration of
spent conditioners is possible after the available water is depleted by plant uptake
and evaporation. Consequently, they have been used in potting media, especially
those of open and sandy nature, to enhance their water-holding capacity thereby
reducing the watering frequency (Flannery & Busscher, 1982 and Gehring & Lewis
III, 1980).
In the Republic of Singapore, watering is a horticultural operation which is not
yet fully mechanized. In some cases, watering of potted plants in the nurseries is
accomplished by manual labour. Therefore, if plant growth media could be physi-
cally altered by polymeric additives to hold more water, labour expended on
watering and water itself, a scarce resource, can be saved.
Materials And Methods
The effects of three polyacrylamide-based polymeric conditioners viz: Agrosoke
(Chemical Discoveries (UK) Limited), Soiltex Gl (Interlates Limited, UK) and
Terra-Sorb (Industrial Services International, Inc.), on the water-holding capaci-
ties of a local topsoil (a sandy clay loam of composition 53% sand, 24% silt and
23% clay; pH 5.5) were investigated.
97
9§ Gard. Bull. Sing. 39(1) (1986)
The indicator plants selected were Mussaenda Dona Luz, Tecomaria capensis
and Hibiscus rosa-sinensis, all known to be susceptible to water stress. Uniform
plants, especially in terms of leaf coverage, of the above species were chosen and
inplanted into a standard amount of topsoil alone, topsoil with treated and pulve-
rized sludge in the ratio 3:1 v/v (standard departmental mix for planting holes),
topsoil with 3 rates of each conditioner (Table 1) or a synthetic topsoil of formula-
tion 1/3 Rengam subsoil: 1/3 treated and pulverized sludge: 1/3 granite dust (v/v/v;
pH 6.2) (Teoh & Chua, 1975; Wells, 1977). Clay pots of dimensions 23 cm (top
internal diameter) X 16cm (height) x 18 cm (bottom internal diameter) were used
and each was packed with 4.5 kg of substrate. The treatments are summarized in
Table 1.
Table 1
Summary of treatments
Treatment Specification
>
Control: topsoil without additive
Topsoil: Agrosoke (2000: 1/w/w)
Topsoil: Agrosoke (1000:1 w/w) (Recommended rate)
Topsoil: Agrosoke (1000:2 w/w)
Topsoil: Soiltex G 1 (2000:1 w/w)
Topsoil: Soiltex G 1 (1000:1 w/w) (Recommended rate)
Topsoil: Soiltex G 1 (1000:2 w/w)
Topsoil: Treated sludge (3:1 v/v)
1/3 Rengam subsoil: 1/3 Treated sludge: 1/3 granite dust (v/v/v; synthetic topsoil)
Topsoil: Terra-sorb (2000:1 w/w)
Topsoil: Terra-sorb (1000:1 w/w) (Recommended rate)
Topsoil: Terra-sorb (1000:2 w/w)
Pan a Ss © Ge Ml Cae we
The conditioners were mixed with the topsoil in the deactivated state and later
activated by repeated flooding after the plants were established in the mixes.
All treatments and control were replicated 6 times for each plant species used,
giving a total of 72 pots per plant species. After implantation, plants were arranged
in randomized complete blocks and allowed to establish for a week under a shed of
transparent polythene cover. This provided shelter from the rain while leaving
ample room for air circulation and heat transmission.
To initiate the trial, pots were soaked thoroughly and allowed to drain for half an
hour to achieve field capacity. Immediately, a 10 g sample of substrate was
collected from a depth of 8 cm from each pot for the determination of moisture
content at field capacity by drying to constant weight at 105°C. For the first 4 days,
sampling was done twice daily, once at 10 a.m. and the other at 4 p.m. Thereafter,
sampling was carried out once daily in the afternoon for moisture determination
until the permanent wilting point was reached. The number of days taken to reach
the permanent wilting point was recorded (DTW). The difference between the
moisture content at field capacity and that at the permanent wilting point yielded
the index for plant-available water.
Water Absorbent Polymers and Irrigation Needs 99
Results And Discussions
Results on total water contents, available water contents and days-to-wilting
(DTW) are presented in Tables 2, 3, 4 and 5. Data were analysed with the Duncan
Multiple Range Test for significance.
Data in Table 2 indicated that the departmental planting mix comprising 3
volumes of topsoil to 1 volume of treated sludge was significantly superior to all
other treatments in terms of total water-holding capacity. Total water-holding
capacity was similar with or without adding polymers to topsoil (cf. Control (A)
and treatments with conditioners in Table 2). Increasing the rate of conditioner in
certain cases did not result in a higher total water-holding capacity (cf. Treatments
(B) and (C), and Treatments (E), (F) and (G) in Table 2). It appeared that the
three rates of conditioners applied did not augment significantly the water-holding
capacity of the topsoil, a sandy clay loam. Also, according to Flannery & Buscher
(1982), the full capacity of polymeric soil conditioners could only be effected after
long periods of saturation.
In the case with Tecomaria capensis as the indicator plant (Table 3), DTW was
not affected by treatments. The available water contents did not relate to the rates
of the soil conditioners added. In fact, the highest rate of soiltex Gl appeared to be
the poorest in maintaining available water for the trial plant. In terms of available
water, Treatments H (the departmental planting mix) and A (Control) yielded
results better than Treatment G but comparable to those of other treatments.
In the case with Hisbiscus rosa-sinensis (Table 4), no significant treatment-differ-
ence on available water was detected. However, the effect of Treatment H on
DTW was significantly better than a majority of other treatments. DIW was
similar in the absence (A, control) and presence (Treatments B, C, D, E, F, G, J,
K and L, Table 4) of polymers in the topsoil.
Table 2
Effects of various treatments on the total water-holding capacities
Treatment A B i D Fs F G H I J K L
Total water 21.58 22.55 21.04 22.07 24.33 23.19 18.73 27.19 18.55 18.26 19.04 20.03
(%) bg be cdefg bf ob a oe 2 ¢ cfg cefg
Treatments A-L have reference in Table 1.
Values in each row if not followed by the same letter are significantly different as judged by the DMR
test at P<0.01.
Table 3
Effects of various treatments on the number of days taken to reach the permanent wilting point (DTW)
of Tecomaria capensis and the available water contents
Treatment A B Cc D E F G H I J K L
DTW 16.0a 14.7a 14.5a 15.0a 15.5a 16.0a 16.0a 16.8a 13.8a 14.0a 12.5a 15.8a
Available 18.4 20.4 163 16.0 18.9 20.3 12.0 16.9 18.0 16.2 16.7 19.1
water (%) ad a bed bd ad ac b ad ad bd ad ad
Treatments A-L have reference in Table 1.
Values in each row if not followed by the same letter are significantly different as judged by the DMR
test at P<0.01.
DTW = Days to Wilting.
100 Gard. Bull. Sing. 39(1) (1986)
Table 4
Effects of various treatments on the number of days taken to reach the permanent wilting point (DTW)
of Hibiscus rosa-sinensis and the available water contents
Treatment A B iD D E F G H I J K b
DTW 10-0, 1.9.7 1037, .1155. 4 10:2 TOS. a4 2 tS tee Te ee 8.0
ape S.a0C . witGe ca abc . ce bd d abe™ “ac abe. ae
Available 18.2 17:1). 18.3 °©20:2 19:9. .19.3) 121.0%) 22}4>14930 AG ae
water (%) a a a a a a a a a a a a
Treatments A-L have reference in Table 1.
Values in each row if not followed by the same letter are significantly different as judged by the DMR
testt at P<0.01.
DTW = Days to Wilting.
Table 5
Effects of various treatments on the number of days taken to reach the permanent wilting point (DTW)
of Mussaenda Dona Luz and the available water contents
Treatment A B . D E E G H I J K L
DTW 5.7a -5.8a” 3:5a°* 35a’ 5.8a 52a" Sd S.9a “4eSar aa
Available 15.9 17.6 (15:5 25.59 e220 1e tot 103 16:8) 16.) there iGy se
water (%) a a a ac a a be a a ac a a
Treatments A-L have reference in Table 1.
Values in each row if not followed by the same letter are significantly different as judged by the DMR
test at P<0.01.
DTW = Days to Wilting.
Mussaenda Dona Luz appeared to be the most susceptible to water stress among
the three indicator plants selected. It reacted to water stress by shedding its leaves.
The first sign of leaf drop was therefore construed to be the permanent wilting
point. Results obtained with this trial plant showed no significant treatment effect
on DTW (Table 5) and only marginal effects on available water. As with Tecomaria
capensis, there was no significant correspondence between the rate of conditioner
applied and the water-holding capacity. Treatments H and A were significantly
different from Treatment G but similar to all other treatments in terms of available
water.
Conclusion
The present trial did not show any significant enhancing effect of polymers on the
water-holding capacity of a typical topsoil. Control A, irrespective of the indicator
plant used, showed similar effects in total water-holding capacity, available water
and DTW to those of treatments with conditioners. Our results indicated that the
standard planting mix of the Parks and Recreation Department compared more
than favourably to topsoil treated with various rates of the polymeric soil condition-
ers, in the three attributes measured.
However, our results do not rule out the possible usefulness of these polymers in
sand or more sandy soils, for which such products are more intended for, and in
Water Absorbent Polymers and Irrigation Needs 101
planting where a larger volume of substrate is involved. In fact, Gehring & Lewis
III (1980) have found significant relationships between the wilting time and the
container size used in their study of a polymeric soil conditioner.
References
Azzam, R.A.I. (1985). Tailoring polymeric gels for soil reclamation and hydropo-
nics. Commun. In Soil Sci. Plant Anal., 16, 1123-1128.
Callebaut, F., Gabriels, D. & M. De Boodt (1979). The effect of polymer
structure on soil physico-chemical properties and soil water evaporation. Journal
of Chemical Technology & Biotechnology, 29, 723-729.
De Boodt, M. (1979). Soil conditioning for better soil management. Outlook on
Agriculture, 10, 63-70.
Flannery, R.L. & W.J. Busscher (1982). Use of a synthetic polymer in potting soils
to improve water-holding capacity. Commun. Jn Soil Sci. & Plant Anal. 13,
103-111.
Gehring, J.M. & A.J. Lewis III (1980). Effect of hydrogel on wilting and moisture
stress of bedding plants. J. Amer. Soc. Hort. Sci. 105, 511-513.
Gunn, S. (1984). Like a sponge. GC & HTJ, August 10, 19, 21 & 24.
Hamilton, J.L. & R.H. Lowe (1982). Use of a water-absorbent polymer in
tobacco seedling production and transplanting. Tobacco Science, XXXVI, 17-20.
Page, E.R. (1982). Soil conditioners. Scientific Horticulture, 33, 1-13.
Teoh, T.S. & S.E. Chua (1975). The performance of several ornamental shrubs in
different potting mixtures of subsoil, sludge and granite dust. Singapore J. Pri.
Ind., 3, 21-34.
Wells, N. (1977). The role of soils in the utilization of sludge in Singapore. N. Z.
Soil Bureau Scientific Report 30. Department of Scientific & Industrial Research,
New Zealand.
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5
a JUN 23 1987 ISSN 0374-7859
THE GARDENS’ BULLETIN
\
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x VOL. 39 (Part 2) | December 1986
oO Oooo
CONTENTS
N
. PAGES
\
CORNER, E.J.H.: |
The Agaric Genus Panellus Karst. (including Dictyopanus Pat.) in Malaysia ................ 103-147
BIDIN, Aziz & JAMAN, RAZALI:
A New Species of Platycerium trom Peninsular Malaysia ..................0.0.:0ceeeeeeeeeeee eens 149-15]
y
, x HOLTTUM, R.E::
3 STUDIES IN THE FERN-GENERA ALLIED TO Tectaria Cav. VI: A conspectus of genera in the
Old World regarded as related to Tectaria, with descriptions of two genera ................. 153-167
(J TINDALE. Mary D.:
a \ A New Genus and Three New Species of Pteridophytes from North Eastern Queensland 169-175
BAY
: ¢ KOSTERMANS., A.J.G.H.:
‘ i epee een eraee C semcemel (GelaStFACCAC) 2. so. .c 2. Uiwndevcds uc deecesnseceeseceece-edeveeeseess 177-191
iy \
: \ STONE. BENJAMIN C.:
7) The Genus Pandanus (Pandanaceae) on Christmas Island, Indian Ocean ................... 193-202
aX
f ) WONG, Yew Kwan:
m The Use of Tifgreen and Tifdwarf Bermuda Grasses in two Singapore Golf Courses ..... 203-214
Published by the Botanic Gardens
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Ministry of National Development
Cluny Road, Singapore 1025.
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THE GARDENS’ BULLETIN
SINGAPORE
Wat. 39 (Part2) 1 December 1986
CONTENTS
PAGES
CORNER. E.J.11.:
The Agaric Genus Panellus Karst. (including Dictyopanus Pat.) in Malaysia ................ 103-147
BIDIN, Aziz & JAMAN, RAZALI:
Peavew Specics o1 Piatycersum from Peninsular Malaysia ...cov.: 0.00. ee ces. een le eee eee ak neces 149-151
HOETTUM:. R-E *
STUDIES IN THE FERN-GENERA ALLIED TO Tectaria Cav. VI: A conspectus of genera in the
Old World regarded as related to Tectaria, with descriptions of two genera ................. 153-167
TINDALE., Mary D.::
A New Genus and Three New Species of Pteridophytes from North Eastern Queensland 169-175
KOSTERMANS. A.J.G.H.:
nomen vali amscime 4-~ (CClas(racede yi)... 1 ide Wau cedece «leila deca deka codes chevansiees 177-191
STONE, BENJAMIN C::
The Genus Pandanus (Pandanaceae) on Christmas Island, Indian Ocean ................... 193-202
WONG, Yew Kwan:
The Use of Tifgreen and Tifdwarf Bermuda Grasses in two Singapore Golf Courses ..... 203-214
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The Agaric Genus Panellus Karst.
(including Dictyopanus Pat.) in Malaysia
E.J.H. CORNER
91 Hinton Way, Great Shelford, Cambridge CB2 SAH, England
EFFECTIVE PUBLICATION DATE: 24 JAN. 1987
Abstract
Panellus, with Dictyopanus as a synonym, has 27 species in Malesia; 11 species (all new) are lamellate;
13 species (8 new) are poroid. Two new subgenera are made for species with more or less centric pileus
and well-developed stem, namely Megalopanellus (1 species) and Mesopanellus (2 species). P. longin-
quus (Berk.) Singer is recorded tentatively for Queensland.
New taxa in subgen. Panellus — lamellate species, P. alutaceus, P. ambiguus, P. bambusarum, P.
brunneomaculatus, P. dichotomus and var. pinnatus, P. exiguus, P. fuscatus, P. intermedius and var.
stenosporus, P. parvulus, P. pendens, and P. sublevatus; as poroid species, P. albifavolus, P. bambusifa-
volus, P. brunneifavolus, P. hispidifavolus, P. megalosporus, P. microsporus, P. pauciporus, and P.
sublamelliformis; subgen. Megalopanellus, P. magnus; subgen. Mesopanellus, P. glutinosus, P pyru-
liferus.
Contents
Ss RE ae I Log ee A, a re p. 103
aN aD ote es Se ies oe ca eee Mahe eed Sas acetic eceecceasacehsewerses p. 104
NR to ae e's cog ahac's asp igh ee enioe s so00 eh io osereamescedversaeeee p. 104
ite ee OI SNCCICS OFF GTICHIUES I DAANCSIA 68202228 2o eo. ede cadence cones eee sceceseencececeees p. 105
a ee RCE ESE TATE CNMI Dn cds = ade cade eevee cea gegsocacs Sens cnscdeeserecerbeccansde p. 106
A ag hos a See St ne oe ae p. 109
2 Esse DJS Ce SS EBA A pe a ote oes Sh eee ee ee ee ee p. 110
nee Oa MEMOIR SRD MRED stag eis BL CRASS Co eaa sees a bs Vs Ad hte, sWowercpblnnn ovine deccgeeeons be’ p. 110
i LESS oS Ree 5 ESS UN (eC RS ar eA Eee <5 Sy en ae Cae A Ol Pt ee eee ee p. 112
ee es Ee AN SPMENSUMIREHICT A gc: dew eid 9 - Seer nce nabs dooce ks We nce cna secee sino teceseocares p. 113
Inert A BA ETO a cee acts 3 hace Cpt 0 Soca rieaugs baw icine vous e-sbevegrstracnadancuaess p. 114
UI IER le oe ee at IE les asic ima ns ane ns np annals 8p p. 128
Sa TOME BS BRINE ace ce Ss ER as aiding Suk aces are eunpene seas nies yes p. 139
Sa Fas Te PR RIND oe ho Sa 22 To calc aco wet T= mnie soma es dee aaa nawsesceedceesesesenes p. 143
SI Nia SE ei BE eS I oe a ac wipe Aim Coons fon dhe ea eed be enecetaceseenacces p. 146
I es te Oe Sou, enact Monee PIA Dane yoragid ccc deeoneccnpaucesctscsesncwedensucse p. 147
Introduction
Some twenty years ago, I studied in detail the collections of this genus which I
had made in the Malay Peninsula, Borneo, New Guinea, and the Solomon Islands.
I offer these notes and drawings for others to pursue the quest. The Malacca fungus
flora enlarges the concept of the genus very considerably, and there are certainly
more species to be found. In a few cases I have introduced drawings of exotic
material because they convey better the construction than such as I have of the
Malesian.
Received on 9 October 1985
103
104 Gard. Bull. Sing. 39(2) (1986)
Panellus is like Pleurotus but with more or less amyloid spores, commonly with
acerose basidioles in the hymenium (as a marasmioid feature), and commonly with
granular-encrusted dichophysial hyphae on the edges of gills or pores, and on the
pileus and stem. In consequence of this last feature, the pileus is usually opaque,
not striate. The fruit- bodies are generally rather small to minute, rather tough or in
parts gelatinous, and on the whole without bright colours. They are white to dull
ochraceous, brownish, or somewhat rufous, but there is an exception in the golden
yellow P. aureofactus Horak of India.
The best introduction is to study illustrations of the north temperate P. stypticus,
which is the type-species and has much in common with the Malesian P. dichoto-
mus. They have the opaque incrustation which renders almost impossible the
microscopic study of the fruit-body until sections are treated with dilute potash
which fortunately, dissolves all or most of the granular matter fairly quickly. With
this picture in mind and with the amyloid test for the spores, it is easy to recognise
most lamellate collections. The species grow on dead wood and various plant-
remains, and are not terricolous. A chalky white edge to the gill may be a sure sign.
However, I have added from the Malesian flora two new subgenera, Megalo-
panellus and Mesopanellus, in which the stem is not lateral or the pileus sessile, as
usual in Pleurotus, but central or excentric so that one begins to think of an
ordinary agaric as Collybia, Marasmius, or Tricholoma. The decurrent gills, amy-
loid spores, acerose basidioles, dichophysial processes from the superficial hyphae,
and the pallid colour of the pileus become diagnostic. In fact, Megalopanellus
begins to disclose with its large fruit-bodies the ancestry of the genus.
Dictyopanus Pat.
In the course of collecting, one will find that the underside of the fruit-body, with
obvious resemblance to Panellus, is not lamellate but closely poroid with the
characteristic chalky opacity (Fig. 2). These polyporoid species were the object of
the genus Dictyopanus, for which it would serve a useful purpose if generic names
were taken as ideas and not, as now, for type-species. It has been realised for some
time that no sharp line can be drawn between the lamellate and the poroid species;
there are intermediates partly poroid and partly lamellate or with the gills closely
connected by poroid reticulations in the interstices. It happens that the type-species
of Dictyopanus, namely D. pusillus, has just such an intermediate variety in the
American tropics; it is var. pseudorhipidium Singer, of which var. sublamellatus
Corner may be a synonym (Fig. 3). Accordingly, Dictyopanus has been reduced to
Panellus by Burdsall and Miller (1975) and referred to subgen. Panellus. Presum-
ably all species of Dictyopanus must be transferred to Panellus whether or not they
have lamellate indication; otherwise, they are without a genus unless it is Favolas-
chia.
Favolaschia (Pat.) P. Henn.
This genus of long uncertainty, dating from 1895, has been revised by Singer
(1945, 1975) and by Dennis (1952, 1970). The extraordinary fact that emerges is
that their definitions reveal no distinction from Dictyopanus. Perusal of the species
now attributed to Favolaschia shows that some are panelloid as Dictyopanus and
others are mycenoid, including the type-species. Singer (1969) made the new family
Favolaschiaceae to be allied with the thelephoraceous Aleurodiscaceae, in which
he is followed by Jiilich (1981): Alliance with Panellaceae was ruled out; yet, there
is no fundamental difference between the descriptions of the two families; they
read almost exactly alike. Clearly, there is still some misunderstanding. Thus, D.
gloeocystidiatus Corner (1954), which I here transfer to Panellus, was left out of
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Fig. 1. Panellus sp. (aff. P. pteridophytorum Singer) from Brazil, to show the lamellate form with
distant gills; x 12, the spores x 1200.
consideration because it upset any supposed distinction. I do not regard Favolas-
chia as a suitable deposit for poroid Panellus.
Lamellate and poroid species of Panellus in Malesia
Dictyopanus having been lost, I refer to the many species with gills, not pores, as
the lamellate, and to the many with pores, not gills, as the poroid. The question
arises whether the species of one group can be related to any of the other. The
105
106 Gard. Bull. Sing. 39(2)(1986)
more or less lamellate variety of P. pusillus has not been found in Malesia, but
there is the new species P. intermedius of Borneo with deeply poroid-reticulate gills
as an intermediate (Fig. 14), and I add the poroid P. sublamelliformis of the
Solomon Islands, the pores of which become lamelliform peripherally. I have
searched for instances and found only the close relation between the lamellate P.
parvulus and the poroid P. sublamelliformis, both growing on bamboo remains in
the Solomon Islands. I listed the species of both categories according to spore-size
as the most immediate means of assessing affinity, and found four other possible
pairs of lamellate and poroid species. When, however, other details of construction
were taken into account, it became obvious that the pairs could not be closely
related in spite of their similarity in spore. These points of construction showed that
there were remarkable anatomical parallels between the lamellate and the poroid.
Thus, for the species with acerose basidioles, there are three sets of parallels, as
follows:
1. Cheilocystidia and pileocystidia absent.
Lamellate — P. brunneomaculatus, P. sublevatus
Poroid — P. copelandii, P. pauciporus
1. Cheilocystidia present.
2. Cheilocystidia and/or pileocystidia gloeocystidial, subcylindric
Lamellate — P. dichotomus, P. parvulus
Poroid — P. bambusifavolus, P. gloeocystidiatus, P. megalosporus, P. orien-
talis
2. Cheilocystidia not gloeocystidial, more or less clavate.
Lamellate — P. bambusarum, P. exiguus, P. intermedius
Poroid — P. brunneifavolus, P. luminescens, P. microsporus, P. pusillus, P.
sublamellatus.
It looks, therefore, as if the poroid may have arisen along three separate lines of
evolution, and that Dictyopanus must be regarded merely as an unnatural form-
genus.
I have recorded 8 species as growing on bamboo remains, one from Borneo and
the others from the Solomon Islands, thus:
Lamellate — P. ambiguus, P. bambusarum, P. parvulus, P. sublevatus (Borneo).
Poroid — P. bambusifavolus, P. brunneifavolus, P. pauciporus, P. sublamelli-
formis.
Of these, one could associate on spore-size the lamellate P. parvulus with poroid P.
sublamelliformis, and the lamellate P. ambiguus with the graminicolous and poroid
P. copelandii. Singer has described several bambusicolous species from South
America none of which seems to fit with the oriental.
Surface structure of the fruit-body
In subgen. Panellus, to which most Malesian species belong, the general con-
struction of the surface of the pileus, stem, and gill-edge is shown in Figs. 4 and 5.
Cystidia protrude into a superficial felt of narrow hyphae the ends of which ramify
into lobules, spicules, or diverticula, and the ends are more or less heavily en-
crusted with the granular matter that dissolves in dilute potash. The construction
seems to be a perfection of Singer’s Rameales-structure. The narrow, ramified, and
diverticulate hyphae have been called acanthophyses, dendrophyses, dichophyses,
and astereostromelloid according to the appearance of one or other of these
branching hyphal ends with die-away growth. The branchings and lobings, howev-
er, are generally so varied and intricate that there are many transitions, even in the
Same species, from one ideal form to another. I prefer to call them simply
Fig. 2. Panellus luminescens, showing the poroid form of the genus, previously called Dictyopanus; X
15. (from Mycologia 42, 1950, 424).
Fig. 3. The sublamellate variety of Panellus pusillus; x 10. (from Trans. Brit. Mycol. Soc., 37, 1954,
260).
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Fig. 4. Panellus gloeocystidiatus. Diagram of the structure of the pileus, with the gloeocystidia shown
in black. (from Trans. Brit. Mycol. Soc. 37, 1954, 257).
dichophysial hyphae. They appear as sterile hymenial hyphae given over to exces-
sive branching with decreasing width until mere lobules or spicules are produced.
They do not occur in the fertile hymenium except occasionally as an invasion from
the edge of gill, pore, or pileus.
This surface structure develops as a palisade of cystidia on the primordial parts.
The palisade becomes disrupted as the internal tissues expand and the dichophysial
hyphae grow out between the cystidia to overtop them; they may extend up to 150
before developing the terminal mat of encrusted diverticula. In some cases,
especially when the cystidia are gloeocystidial and subcylindric, they lengthen into
the intricate felt of hyphae, even exceeding themselves and becoming septate
(without clamps) at the apex as the cytoplasm is withdrawn. In other cases,
especially with more clavate ones, they may themselves produce an intricate tangle
of coralloid and spiculiform dichophysial hyphae.
There are the following variations in surface structure:
1, The cystidia may have dense oleaginous contents and can be called gloeocysti-
dia. In some cases the contents become vacuolate and the original gloeocysti-
dial nature is not evident in the mature tissue.
2, As mentioned, the cystidia may themselves produce dichophysial hyphae and,
in so doing, may also become unrecognisable in the mature tissue.
3, The mat of dichophysial hyphae may not develop, particularly on the pileus or
the edge of gill or pore. Examples are the lamellate P. alutaceus, P. fuscatus,
P. pendens, and the poroid P. albifavolus, P. hispidifavolus, and P. micro-
sporus.
4, The surface of the pileus may be more or less gelatinous. Examples are the
lamellate P. exiguus and P. longinquus and the poroid P.
Singer gives several examples from tropical America.
108
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4
f;
(au
BY,
Fig. 5. Panellus gloeocystidiatus. Details of the surface of the pileus with gloeocystidia and dichophy-
sial hyphae (simplified), with basidia and acerose basidioles, x 1000. (from Trans. Brit. Mycol.
Soc. 37, 1954, 257).
This is a degradation series in which the original and detailed construction
disappears. As in other cases with a hymeniiderm, or palisade of clavate cells, such
as Amauroderma, Boletus, Entoloma, or pluteus, it presupposes a trichoderm or
cortical pile of inflated and septate ends. The extensive hymeniiderm is shown by
subgen. Mesopanellus (Figs. 24, 25) and the primitive trichoderm by subgen.
Megalopanellus (Fig. 28). Both subgenera have the more primitive characters of
centric pileus and inflating hyphae.
Hyphae of the fruit-body
The hyphae are monomitic, clamped, inamyloid, and usually colourless, and
have short cells up to 250 yw long, in a few cases longer but not of exceptional length.
There are, however, such differences in the degree of inflation, the thickening of
the hyphal walls, and their tendency to become gelatinous that more precise
characterisation is impossible (Fig. 6). In subgen. Panellus the greatest inflation
109
110 Gard. Bull. Sing. 39(2) (1986)
occurs in P. parvulus with thin-walled hyphae -20 uw wide. P. intermedius and P.
sublevatus have some hyphae -16 w wide. In others, the hyphae have slightly
inflated cells 8-14 wide in the stem and base of the pileus; commonly these cells
are inflated unequally in different parts, perhaps because of the lateral pressure
from the hyphae. Failure to support the rather dense fruit-body by turgidity is
compensated by thickening of the hyphal walls, even up to 5 uw thick, with corres-
pondingly reduced lumen. As a variation, the whole tissue becomes toughly gelati-
nous with agglutinated and uninflated hyphae, for instance P. dichotomus.
The gradual loss of inflation implies the derivation of Panellus from some typical
agaric ancestry which is presented by P. magnus of subgen. Megalopanus. This
species shows the tendency to dichophysial hyphae arising almost like binding
hyphae in the inner tissue (Fig. 29). In contrast, the poroid species have more or
less uninflated hyphae in the pileus in accordance with their subcircular pores.
Basidioles — pleurocystidia
The hymenium of most species carries, as well as basidia, narrow fusiform cells
3-5 w wide, often with acute, or even, mucronate tip. They are regarded as sterile
cells or basidioles though it seems that it has still to be shown that they are not
merely immature basidia which, on charging, would become clavate (Fig. 7).
Acerose basidioles occur in several genera of agarics, such as Collybia and Maras-
mius, but it is not clear if they have generic distinction. In some species which agree
in other respects with Panellus, basidioles seem not to occur. It is impossible to
prove a negative, in this case their total absence, and I have not used the feature as
a primary character.
Pleurocystidia differ from basidioles in their larger size which follows from their
early differentiation in the developing hymenium; they are usually the first mature
cells to be formed and have, in consequence, a deeper origin in the subhymenium,
even from the first out-turned branches of the tramal hyphae. Basidioles are the
last formed cells and appear in numbers as branches from the more superficial
subhymenial hyphae. Pleurocystidia have been described for some South American
species, as P. cystidiosus Singer and P. dumontii Singer, but seem to be lacking
from other species of subgen. Panellus unless, perhaps, in the Malesian P. bambu-
sarum. The structures called pleurocystidia by Burdsall and Miller (1975) seem to
be basidioles, but the temperate P. serotinus, in their subgen. Mitellus has abun-
dant pleurocystidia. Species of this subgenus, the distinction of which is not clear,
have not been found in Malesia.
Spores and basidia
The impression, gathered from the literature, is that Panellus (including Dic-
tyopanus) has rather small, often cylindric to allantoid, spores. This contrasts with
the rather large, ellipsoid to subglobose, spores in many. species of Favolaschia.
Any such distinction does not hold, however, in the eastern tropics. In 1963
Kobayasi described D. orientalis with subglobose spores 8 X 7 w. I add several
lamellate and poroid species with moderately large ellipsoid or subglobose spores,
among which the poroid P. megalosporus is outstanding with subglobose spores up
to 18 X 16 uw, produced by enormous basidia averaging 90 x 17.5 w. In contrast, the
smallest spores in Malesian species are the minute allantoid spores, averaging 3.5 x
0.8 uw, of the poroid P. microsporus (Fig. 8). Panellus is revealed as a genus with
great latitude in spores and basidia and one that must have come from an agaric
source with large spores on large basidia which qualify for Singer’s Hygrophorus
kind; that is with basidium-length at least 5-6 times the width of the spore. No
: ‘
x ” a -) v= *
: 2
Fig. 6. Panellus sp. (aff. P. pteridophytorum Singer), showing the variation in inflation of the hyphae
of the fruit-body and in the thickening of the walls; x 500.
lamellate Panellus is known with spores as large as those of P. megalosporus,
though such a one may be found. I note that the minute allantoid spores of P.
microsporus should place it in subgen. Mitellus but, in all other respects it belongs
in subgen. Panellus. I note, too, that the spores of the Malesian P. albifavolus, P.
ambiguus, and P. brunneomaculatus are hardly amyloid, if at all.
With this range in size of spore and basidium I constructed sporographs and
basidiographs but found, as with Polyporus (Corner 1981), that no clear locus was
obtainable. The reason seemed to be that the variation in size shown mainly by the
poroid species, was effected by compression of the basidium-unit in the tubes.
Thus, the ratio of spore-width to basidium-width varied from 0.21 in the highly
compressed and allantoid spores of P. microsporus through all gradations to 0.8 in
P. megalosporus and ().87 in P. orientalis. However, the usual trends were clear, as
follows:
1, Spore-volume increased in direct proportion to basidium-volume.
2, Spore-width increased with spore-length.
3, Basidium-width increased with basidium-length, the smallest structures being
the basidioles.
11]
DT a We Ey
yd )~
TLC
2
Tyit
>
os Fn
SSS
we 2 as! , 1
Bd ft my
eg cee” :
/
Ol St
Pau
ans
‘ 2 7
Zn ae *
BRON th
Fy. < abi see »> e
al yr ex
ak’ SXF Vins ae
Hue ard 9 MS 2 Aan eee Oy te
Ce SNe hig sit re yy Liv poe
Bi 3 oS Ai ve es ere, vt :
Fig. 7. Panellus orientalis, showing the pore-edge with gloeocystidia and dichophysial hyphae; x 600.
4, The large spores and large basidia of P. megalosporus were consistent with an
average sporograph-locus and an average basidium-locus, the value of which,
for the basidium-locus, was //w = 2.9 x 0.025/, where / is the basidium-length
and w its width.
Taking figures that I have given for other genera with large spores and basidia
(Corner 1983), //w varies as 0.069/ in Heimiella, 0.02] in Oudemansiella, and 0.0131
in Amauroderma and Ganoderma. These values contrast with 0.08/ for the narrow
clavarioid basidium.
Phosphorescence
Several species of subgen. Panellus are known to be phosphorescent with a
greenish light emitted from the gills or pores and the mycelium; others are not
phosphorescent, but the majority remains to be tested. The phosphorescent are P.
stypticus (in North America), P.-/uminescens, P. gloeocystidiatus, and the Japanese
Dictyopanus foliicolus Kobayasi. P. orientalis, according to Kobayasi, and P.
megalosporus are not phosphorescent. In the case of P. stypticus, North American
112
Fig. 8. Basidia and spores of Panellus, to show the variation in size and shape; X c. 700. a, P.
megalosporus; b, P. brunneifavolus; c, P. luminescens (large spores); d, P. orientalis; e, P.
luminescens (small spores); f, P. gloeocystidiatus; g, P. microsporus.
specimens are phosphorescent but not the European. When the mycelia of the two
forms were mated, it was found that phosphorescence appeared as a dominant
character (Macrae 1937).
Possibly the origin of Panellus is to be related with that of the phosphorescent
species of Pleurotus with large spores (Corner 1981).
Panellus Karst.
Hattsv, Bidr, Finl. Nat. Folk 32 (1879) xiv; Singer, Agaricales Mod. Tax. (1975) 339.
Dictyopanus Pat., Ess. Tax. Hym (1900) 137; Singer. Agaricales Mod. Tax. (1975) 338.
Pileus mesopodal, pleuropodal, or sessile, medium-sized to very small, mostly
white, bistre, ochraceous, to cinnamon and rufous brown but golden yellow, olive
green, or violaceous in exotic species, mostly opaque, gymnocarpic with incurved
margin. Gills adnate to decurrent, or with small to minute pores; edges of gills or
pores usually chalky pruinose. Flesh thin, soft, tough or more or less gelatinous.
Spores white, smooth, more or less amyloid, subglobose to ellipsoid, cylindric or
allantoid, large to minute. Basidia mostly 4-spored; basidioles subacerose, but
lacking in some species; subhymenium usually narrow, with interwoven hyphae;
hymenium not thickening. Cheilocystidia usually present, thin-walled, with or
without oleaginous contents, in some species emitting diverticula, forming a sterile
edge to gill or pore, generally becoming overgrown with encrusted dichophysial
hyphae. Pleurocystidia generally absent except in some exotic species. Hyphae
monomitic, usually clamped, inflating or not, with thin or thickened walls, in some
113
114 Gard. Bull. Sing. 39(2) (1986)
species more or less gelatinous, rather short-celled, not sarcodimitic, not amyloid.
Surface of pileus and stem usually with a palisade of cystidia becoming disrupted
and overgrown by dichophysial hyphae encrusted with granular matter soluble in
dilute potash in some cases practically smooth, with or without a thin gelatinous
pellicle.
On dead wood, stems, and leaves, Cosmopolitan, c. 55 species, 27 species in
Malesia and the Solomon Islands.
This is the best that I can do to extract a generic character. The genus has what
are called pleurotoid and marasmioid fruit-bodies distinguished by the amyloid
spores and, usually, by the excrescent and encrusted dichophysial hyphae. No
poroid forms are mesopodal.
1. Pileus mesopodal or excentric, with well developed stem. Gills not reticulately veined. Hyphae
generally not granular encrusted, inflating
2. Pileus 2-9 cm wide, fuscous fuliginous furfuraceous squamulose. Basidioles lacking. Pileus with a
disrupted trichoderm without dichophysial hyphae. Flesh rather tough, not gelatinous
wag nabatine ya@iunsates aan Cayaneene eee pk eo Ek +5 ga ICR. Sean Aa ae ee subgen. Megalopanellus p. 143
2. Pileus 1-2 cm wide, not furfuraceous squamulose. Basidioles acerose, copious. Cheilocystidia
clavate, with oleaginous contents. Surface of pileus hymentiform and with dichophysial hyphae
edave tenia valsinn ww bewibyeinn abv vdenagicenicde que cae Rete Se ema eee subgen. Mesopanellus p. 139
1. Pileus lateral, dorsifixed, or sessile, usually with short stem. Pileus 1-40 mm wide. Basidioles present
or not. Hyphae often granular encrusted, in some cases more or less gelatinous. Surface of pileus
without a trichoderm, generally with dichophysial hyphae ........................ subgen. Panellus p. 114
3. Lamellate, with or without poroid reticulate interstices ..................068. Lamellate species p. 114
Bu) POLO... ehel as cee ha os wah b Ve SIG a er le Poroid species p. 128
Subgen. Panellus
Singer, Agaricales Mod. Tax. (1975); Burdsall and Miller (1975). 24 species in Malesia and the Solomon
Islands (11 lamellate, 13 poroid).
Key to the lamellate species of subgen. Panellus in Malesia and the
Solomon Islands
1. Hymenium without acerose basidioles. Surface of pileus and gill-edge not granular encrusted
GROUP A
1. Pileus sessile, dorsifixed, -15 mm wide. Gills distant, | mm wide. Spores 4.3-5 x 3.5-4 uw, deeply
amyloid. Surface of pileus with appressed oleaginous hyphae in a single layer. Cystidia none.
BOtne® « ..35.¢50000:0diacns «250m ye goutlne mye aiene tela oeiuglaticle ak ta me oa ieee oe a P. pendens p. 126
1. Pleuropodal, Spores larger, pale amyloid. Pileus -30 mm wide, surface with different structure.
2. Spores 7.5-10 x 3.3-4 w m. Cystidia none. Hyphae -21 mw wide. Surface of pileus with a thin
gelatinous layer, pale bistre ochraceous. Gills -5 mm wide. Queensland ...... P. longinquus p. 124
2. Spores 5-6 4.7-5.7 yw, subglobose. Cheilocystidia clavate to ventricose. Hyphae -13 mu wide.
Surface of pileus dry, with a disrupted palisade of clavate cells with pale fuscous sap, buff-white
and fuscous brownish scurfy., BOTWEO éaiseacs«+vs5apa. opine caer A P. fuscatus p. 121
GROUP B
1. Pileus 1-3 mm wide. Gill-primaries 3-5, often pore-like if without short secondaries
2. Dorsifixed, pale fawn. Spores 8.5-10.7 x 3.7-4.5 w. Cheilocystidia cylindric to clavate. Surface of
pileus subgelatinous. On fern petioles. Malayars vs. 4.120, 00rd. 0a. 0heaoeea P. exiguus p. 120
2. Pleuropodal, white, scurfy-pruinose. Spores 7-8 Xx 5-6 w. Gill-edge and surface of pileus with
dichophysial hyphae. On dead bamboo. Borneo ..............0..ceeeeeneneeceesenees P. sublevatus p. 127
(SPOCeS 45-6. 3-37 pbs, nase sega «covledaneeicd sv tipnsod keine aker eae P. bambusarum p. 117)
The genus Panellus Karst. in Malaysia 115
1. Larger, with more gills
3. Gills with deeply poroid-reticulate interstices. Pileus-10 mm wide, white to pale fawn. Flesh not
gelatinous. Cheilocystidia clavate with a few apical processes. Pileocystidia none. On sticks.
Borneo
4. Spores 8-10 x 4.5-5.7 « Cheilocystidia 5-11 « wide. Hyphae -16 uw wide .. P. intermedius p. 123
4. Spores 7-9 X 3.5-4.5 uw. Cheilocystidia -14 « wide. Hyphae -9 uw wide ... var. stenosporus p. 124
3. Gills not conspicuously reticulate in the interstices, mostly smooth.
5. Surface of pileus and gill-edge not granular encrusted, without dichophysial hyphae. Cheilocy-
stidia involved in brown resinous matter. Spores 4-6 x 2.5-3 yw. Pileus -10 mm wide, sessile,
tiie el GR NSE COTINE Yc acrey <ey Bee ee 9 ao NEL Pala das fied aeeabine sce ees P. alutaceus p. 115
5. Surface of pileus and gill-edge granular encrusted, with dichophysial hyphae
6. Without distinct cheilocystidia or pileocystidia. Pileus -4 cm wide, pleuropodal or excentric,
with stem 3-12 x 2-4 mm. Gills very crowded. Spores 5-6.5 x 2-2.5 uw. On wood, Solomen
EAE apes ie ah SR PS See ERE aa a P. brunneomaculatus p. 118
6. With cystidia
7. Cystidia clavate, not gloeocystidial. Spores 4.5-6 x 3-3.7 w. Pileus -6 mm wide, subses-
sile, white to brownish. On dead bamboo. Solomon Islands ....... P. bambusarum p. 117
7. Cheilocystidia and pileocystidia gloecystidial
8. Cystidia 3-7 uw wide, subcylindric. Flesh subgelatinous. Pileus -3 cm wide, brownish
ochraceous to subrufous. Spores 2.7-3 X 1.7-2 w. On wood ...... P. dichotomus p. 118
SP eis Mec Aatcaly GICMOLOIMNOUS 2. so Fs cede Pe tae seeds wees var. dichotomus p. 118
Sa UMMRAN MIS och gi AR ote eR Ree a ER rg rigg Sw nbn S <p var. pinnatus p. 120
8. Cystidia clavate, especially the pileocystidia. Flesh not gelatinous. Pileus -10 mm
wide, white to pale ochraceous. Spores 6-8 X 4-5.5 uw. On dead bamboo. Solomon
Islands.
10. Cheilocystidia 5-8 « wide. Pileocystidia 12-15 w wide, with copious dichophysial
hnypnac. at the base ob the pueus fc ices2 iss Pee eee sk és dense P. parvulus p. 125
10. Cheilocystidia -16 4 wide, mixed narrow branched hyphae set with short proces-
ses. Surface of pileus set with narrow, simple or sparingly branched, excrescent
hyphae, not dichophysial. Spores not or scarcely amyloid ..... P. ambiguus p. 116
Panellus alutaceus sp. nov. Fig. 9
Ex integro alutaceus, sessilis, horizontalis. Pileus -10 mm in radio, siccus, basim versus subvillosus.
Lamellae confertae. Caro subcoriacea, haud gelatinosa.
Sporae 4-6 x 2.5-3 uw, amyloideae. Basidiola acerosa inconspicua. Cheilocystidia 20-35 x 4-7 pn,
lanceolata vel subventricosa, saepe breviter appendiculata vel capitata, haud incrustata, in resina
brunnea immersa. Pleurocystidia nulla. Hyphae -18 w irregulariter inflatae, fibulatae, tunicis 1-2 mu
crassis, haud incrustatae. Superficies pilei hyphis angustis 2-5 wu latis appressis ut in lamina c. 20 uw crassa
instructa.
Ad lignum emortuum in silva. Borneo, mt. Kinabalu, 20 Jul. 1961, Corner s.n. typus, in herb. Corner.
Fig. 9. Panellus alutaceus. Spores, X 2000; cheilocystidia and hyphae of the pileus, x S00.
116 Gard. Bull. Sing. 39(2) (1986)
Sessile, horizontal, pale biscuit colour. Pileus 1 cm radius, subvillous towards the
base, dry. Gills arising from the base, c. 14 primaries 1 mm wide, 3-4 ranks, not
dichotomous, somewhat crowded, the edge entire and darker. Flesh thin, floccoso-
firm to subcoriaceous, no gelatinous layer.
Spores 4-6 X 2.5-3 uw, white, smooth, pip-shaped, amyloid. Basidia 18-25 x 4 p;
sterigmata 4; acerose basidioles present but inconspicuous. Cheilocystidia 20-35 x
4-7 w, lanceolate to subventricose, often with a short cylindric appendage, capitate
or not, thin-walled, not encrusted but involved in resinous brown matter, forming a
sterile gill-edge. Pleurocystidia none. Hyphae monomitic, clamped, irregularly
inflating -18 uw wide, often of different widths in different parts of a cell, the walls
eventually thickening 1-2 mw closely interwoven and longitudinal in the pileus
descending and subparallel in the gills; not granular encrusted. Surface of pileus
covered by a thin layer c. 20 mw thick of narrow, radiating, mostly thin-walled
hyphae 2-5 uw wide, without pileocystidia but with some excrescent hyphal ends at
the base of the pileus. |
On dead wood in the forest. Borneo.
Mt. Kinabalu, east ridge 3200 m alt., 20 July 1961, Corner s.n.
Panellus ambiguus sp. nov.
Albus, dein in centro pilei pallide ochraceus, sessilis vel subpleuropodalis. Pileus —10 mm latus,
convexus dein planus, striatus. Stipes ut pulvinula lateralis vel nullus. Lamellae adnatae, subdistantes,
primariis 6-12, haud reticulatae, acie pruinosula. Caro mollis, haud gelatinosa.
Sporae 6-8.5 x 4-5.7 wu, haud vel vix amyloideae, Basidiola acerosa. Cheilocystidia trimorpha; 35-50
x 11-16 mw, clavata, intus oleaginosa, copiosa sed aliis superantibus; -55 x 7-12 uw, subclavata vel
subfusiformia obtusa, intus oleaginosa; -50 x 3-5 yw, subcylindrica ramosa, diverticulis copiosis, simplici-
bus spiciformibus vel furcatis, -11 mw longis praedita, leniter incrustata. Pileocystidia ut cheilocystidia
clavata, hyphis 1-1.8 « latis sparsim vel vix ramosis superantibus, haud dichophysialibus. Hyphae 3-14
latae, fibulatae, nec gelatinosae nec incrustatae.
Ad folia caulesque bambusarum in silva. Insulae Solomonenses, Guadalcanal RSS 566 typus, in herb.
Corner.
White, the pileus pale cream ochraceous in the centre, sessile or sub-
pleuropodal. Pileus -10 mm wide, convex then plane, reniform, smooth, striate;
margin subpruinose. Stem as a slight lateral cushion or none. Gills adnate, subdis-
tant, thin, dry, 6-12 primaries -1 mm wide, 3-4 ranks, not poroid or veined, the
edge minutely pruinoso-scurfy. Flesh thin, soft, not gelatinous.
Spores 6-8.5 xX 4-5.7 jw, white, smooth, pip-shaped, with rather oleaginous
contents, not or slightly amyloid. Basidia 24-33 x 6-6.5 wz; sterigmata 4, 4-5 uw long;
acerose basidioles numerous; subhymenium narrow, not corticate. Cheilocystidia
of three kinds, thin-walled, not amyloid or dextrinoid; 1, 35-50 x 11-16 pw, clavate,
with oleaginous-vacuolate contents, very abundant as a sterile edge but overgrown
by the others, smooth; 2, -55 x 7-12 w, narrowly clavate to subfusiform with obtuse
apex, with oleaginous-vacuolate contents, scattered, generally projecting beyond
the first kind; 3, as emergent hyphal ends -50 x 3-5 w, branched and more or less
copiously set along their length with simple or forked, often spicate, processes 2-11
uw long, (not clavate with apical processes), often thinly encrusted, very abundant
along the gill-edge and more or less covering the other cystidia. Pleurocystidia
none, but occasionally the acerose basidioles somewhat enlarged. Hyphae
monomitic, clamped, more or less inflating in stem, pileus and gills with cells 40-200
x 3-14 w and mostly tapered at the ends, not gelatinous, not granular encrusted,
more or less longitudinal, rather compact. Surface of pileus covered with filiform,
simple or sparingly branched, more or less erect hyphae 1-1.8 wide, not gelati-
nous, with scattered and more or less decumbent pileocystidia like the larger
The genus Panellus Karst. in Malaysia 117
clavate cheilocystidia (of first kind), with abundant spores embedded in the finely
villous surface.
On dead bamboo in the forest, Solomon Islands.
Guadalcanal, Gallego, Hidden Valley, 5 July 1965, RSS 566.
This was collected as Marasmius, which the practically inamyloid spores suggest,
but it is difficult to see any satisfactory distinction from Panellus in view of the
variety in the genus, and it is certainly close to P. parvulus.
Panellus bambusarum sp. nov. Fig. 10
Albus, dein pallide brunneus. Pileus -6 mm latus, lateralis subsessilis conchiformis dein planus. Stipes
-1 X 0.3-0.5 mm. Lamellae adnatae subdistantes, primariae 4-6, vel subconfertae primariae 6-11, acie
pruinosa. Caro mollis, haud gelatinosa, in pileo c. 100 uw crassa.
Sporae 4.5-6 X 3-3.7 uw, ellipsoideae amyloideae. Basidiola acerosa. Cheilocystidia -28 x 4-7 pu,
subclavata, hyphis dichophysialibus incrustatis superantibus. Pleurocystidia -18 x 4-6 w, ventricosa, in
apicem gracilem prolongata, sparsa. Pileosystidia -50 x 4-7 uw, copiosa. Hyphae fibulatae (ut videtur),
vix inflatae 1.5-11 mw latae. Superficies pilei stipitisque hyphis dichophysialibus superantibus, tenuiter
incrustatis.
Ad caules emortuos bambusarum. Insulae Solomonenses, Guadalcanal, RSS 1653 typus, in herb.
Corner.
——S—Oe
GE OAWAL SO) AA) FP
Vos: Naso <
Wow
ese fas wey 4 ine Si
ye
Fig. 10. Panellus bambusarum. Fruit-body, x 3; spores, X 2000; pleurocystidia and surface of pileus,
Pileus 3-6 mm wide, lateral, subsessile, conchate then flattened, white then pale
brownish drab-white, substriate. Stem 0.3-1 x 0.3-0.5 mm, lateral, relatively thick.
Gills adnate, subdistant, 4-6 primaries 0.5 mm wide, 2 ranks, but rather crowded
with 6-11 primaries and 3-4 ranks in RSS 1654, not veined or poroid, white then
pale fawn drab, edge whitish pruinose. Flesh c. 100 uw thick at the base of the pileus,
soft, not gelatinous. eS
Spores 4.5-6 X 3-3.7 w, white, smooth, ellipsoid, pale bluish amyloid. Basidia
18-22 x 5 yw; sterigmata 4; subacerose basidioles present. Cheilocystidia -28 x 4.5-7
uw, subclavate, with thin or firm walls, overgrown by a mass of profusely diverticu-
late hyphal ends, with bodies 2-4 w wide, -50 uw long overall, with thin incrustation
soluble in potash. Pleurocystidia -38 x 4-6 w, ventricose with a slender subacute
appendage, thin-walled, not encrusted, scattered. Hyphae monomitic, clamped
(apparently, but difficult to be sure), scarcely inflating, thin-walled: in the stem
longitudinal with cells 25-200 x 1.5-11 yw; in the pileus 1.5-8 wide, radiating; in
the gill-trama 1.5-6 « wide, descending, thin-walled. Surface of stem and pileus as
the gill-edge but with more copious subclavate thin-walled cells -50 x 4-7 w, (not
gloeocystidial), thinly encrusted.
On dead bamboo stems, rather scattered, Solomon Islands.
Guadalcanal. Popomanasiu, 1800 m alt., RSS 1653 & RSS 1654.
118 Gard. Bull. Sing. 39(2) (1986)
Panellus brunneomaculatus sp. nov.
Albus, dein stipite pileoque rufobrunneo-maculatus. Pileus -4 cm latus, convexus dein planus,
villosulus opacus. Stipes 3-12 * 2-4 mm, excentricus vel lateralis, villosulus. Lamellae adnexae angustae
confertissimae, acie albofurfuracea. Caro firma, haud gelatinosa.
Sporae 5-6.5 x 2-2.5 uw, cylindricae, amyloideae vel vix. Basidiola acerosa copiosa. Cheilocystidia vix
evoluta, lamellae acie hyphis dichophysialibus incrustatis copiosis. Pleurocystidia, caulocystidia et
pileocystidia nulla. Hyphae fibulatae, 2-13 y latae, ad basim stipitis crasse tunicatae. Superficies stipitis
hyphis 2-5 yw latis excrescentibus, tunicis tenuibus, plus minus incrustatis praedita, Superficies pilei
hyphis dichophysialibus 1-2.5 mw latis copiosis incrustatis praedita.
Ad truncum emortuum in silva. Insulae Solomonenses, Kolombangara, RSS 1/06 typus, in herb.
Corner.
White, then the stem and pileus suffused and spotted dull rufous brown. Pileus -4
cm wide, convex then plane, smooth, opaque, drying finely subvillous. Stem 3-12 x
2-4 mm, excentric to lateral, short, subcylindric, solid, finely subvillous. Gills
adnexed, very crowded, narrow, 26-31 primaries -1.3 mm wide, 4-5 ranks edge
minutely chalky pruinose. Flesh 2-3 mm thick in the centre of the pileus, firm but
watery. Smell none.
Spores 5-6.5 X 2-2.5 uw, white, smooth, rod-like, very pale bluish amyloid or not
at all. Basidia 19-24 x 4 w; sterigmata 4; acerose basidioles 3-4 uw wide, copious;
subhymenium very narrow. Cheilocystidia none, or as scattered enlarged basidioles
-4() X 6 pw, the gill-edge becoming overgrown with narrow, thin-walled, lobulating
dichophysial hyphae 1-2 mw wide, thinly encrusted. Pleurocystidia none. Hyphae
monomitic, clamped, subcylindric, rather short-celled, the cells 40-250 x 2-13 yw,
with firm walls (subdiffluent in potash); towards the base of the stem the hyphal
walls 1-2 w thick; in the pileus longitudinal and compact; in the gill-trama more or
less descending. Surface of stem with excrescent devious hyphae 2-5 mw wide,
thin-walled, septate, more or less thinly encrusted in places with granules and short
acicular crystals insoluble in dilute potash, forming a thin villous layer 100-200 pu
thick, not as a regular pile and without caulocystidia. Surface of pileus with
appressed radiating hyphae becoming covered with a subvillous layer of narrow,
branching and lobulating, dichophysial hyphae 1-2.5 uw wide, granular encrusted as
on the stem but more profusely branched, without pileocystidia.
On a dead standing trunk in the forest. Solomon Islands.
Kolombangara, low alt., 27 Aug. 1965, RSS 1106.
This has the appearance of a less reduced lamellate Panellus but is peculiar in the
non-dichophysial covering of the stem, in contrast with the pileus and gill-edges,
and the insoluble incrustation. In the almost inamyloid spores, it might be consi-
dered a species of Marasmius, under which name it was collected. Nevertheless, the
species appears to be very close to that described from Argentina with practically
no stem and with shorter spores as P. stenocystis Singer (Sydowia Beih, 7, 1973,
32):
Panellus dichotomus sp. nov. Fig. 11
Pileus -32 mm latus, pleuropodalis, convexus dein applanatus reniformis, minute rimosus, subfurfur-
aceus, cremeo-albus dein cervino-ochraceus vel subrufus. Stipes 1-3 x 2-5 mm, crassus lateralis brun-
neolus. Lamellae confertae, angustae, dichotomae ter quinquiens, cremeo-albae, acie cretaceopruinosa.
Caro tenax, subgelatinosa, albida, sicco cornea.
Sporae 2.7-3 x 1.7-2 w, ellipsoideae, amyloideae. Basidiola subacerosa, copiosa. Cheilocystidia 20-60
x 3-7 w, ut gloeocystidia, cylindrica vel subfusiformia, saepe ex apice diverticulata, hyphis dichophy-
sialibus incrustatis superantibus. Hyphae fibulatae, vix inflatae, tunicis tenuiter subgelatinosis. Super-
ficies pilei stipitisque ut lamellae acie, gloeocystidiis numerosis, hyphis dichophysialibus superantibus.
Ad truncos ramulosque delapsos in silva. Peninsula Malayana, Borneo, New Guinea; RSNB 5362
typus, in herb. Corner.
Fig. 11. Panellus dichotomus. Cheilocystidia, x 1000.
Pileus -21 mm radius, 32 mm wide, pleuropodal, convex then more or less
flattened and reniform, finely cracked and rather scurfy, cream-white then light
fawn, fawn ochraceous, or subrufous; margin incurved at first, white. Stem 1-3 x
2-5 mm, short, thick, lateral, cracked and scurfy, brownish. Gills narrow, crowded,
30-50 primaries -1 mm wide, dichotomous 3-5 times, sometimes with a free short
gill near the margin of pileus, white to pale cream with chalky edge. Flesh -2.5 mm
thick at the base of the pileus, tough, pliant, subgelatinous, drying very hard and
horny, pallid white.
Spores 2.7-3 X 1.7-2 uw, white, smooth, ellipsoid, faintly amyloid. Basidia 14-17
x 3.5-4.5 pw; sterigmata 4; basidioles subacerose, abundant; with more or less
amyloid mucilage round the basidia. Cheilocystidia 20-60 x 3-7 w, subcylindric to
subfusiform with oleaginous gloeocystidial contents, often developing one or more
apical processes with short lobes -7 X 1.5 yw, investing the cystidia, and with
excrescent hyphae 1-2.5 uw wide, with short intricate branches and lobings, even
coralloid, little incrusted with granular matter soluble in potash, as sterile gill-edge.
Pleurocystidia none. Hyphae monomitic, clamped, scarcely inflated, with toughly
subgelatinous walls, inseparable, more or less radiating, descending in the gills.
Surface of pileus and stem developed as the gill-edge as a more or less continuous,
but loose, layer -100 wu thick of cystidia like the cheilocystidia with gloeocystidial
contents, sending out processes 2-4 uw wide, mainly apical, developing into the
lobed and spicate dendrophyses or dichophyses, accompanied by similar processes
excrescent from the narrow superficial longitudinal hyphae, not mucilaginous but
encrusted with brownish granular matter soluble in potash.
On wood, sticks, and fallen branches in the forest. Malay Peninsula, Borneo,
New Guinea; lowland and montane.
119
120 Gard. Bull. Sing. 39(2) (1986)
MALAY PENINSULA. Pahang, Cameron Highlands 1300 m alt., 1 Oct. 1966, Corner s.n. —
BORNEO. Mt. Kinabalu 1300 m alt., Liwagu River, 3 Sept. 1961, RSNB 2715; Bembangan River 1700
m alt., 19 Feb. 1964, RSNB 5362; Mesilau 1700 m alt., 11 Feb. 1964, RSNB 5315. — NEW GUINEA.
Papua, Woitapi c. 2000 m. alt., 23 Aug. 1960, Corner s.n.
Superficially this closely resembles the temperate P. stypticus but differs in the
dichotomous gills with chalky-encrusted edge and not poroid-reticulate, the smaller
spores, the subgelatinous tissue, and the very intricate lobing of the dichophysial
hyphae on stem and pileus. The following var. pinnatus has pinnate gills and,
though intermediates have not been found, it does not seem that it can be separated
specifically from var. dichotomus.
var. pinnatus var. nov.
Differt lamellis haud dichotomis sed ordinibus 3-4(-5) instructis, cinnamomeocervinis, acie flavidula.
Peninsula Malayana, Pahang, Tembeling, 8 Nov. 1930, Corner s.n. typus, in herb. Corner.
Pileus -20 mm wide, 15 mm radius, pleuropodal, reniform, spathulate or flabelli-
form, wholly cinnamon fawn, drying paler and closely rugulose and minutely
cracked. Stem lateral, concolorous. Gills not dichotomous but pinnate with 3-4(-5)
ranks, cinnamon fawn with light yellowish cinnamon scurfy edge, drying dark
brown. Flesh subgelatinous, tough.
Spores 2.7-3 X 1.8 w, ellipsoid, aguttate, very pale amyloid. Microscopically as
var. dichotomus but the gill-edge and surface of pileus and stem with abundant
microscopic brown resinous masses dissolving in Melzer’s iodine into brown oily
masses, giving the dark colour to the dried gill-edge.
On dead fallen branches in the forest. Malay Peninsula.
Pahang, Tembeling, 8 and 10 Nov. 1930, Corner s.n.
Panellus exiguus sp. nov. Figure 12
Pileus -2 mm latus, dorsifixus cupulatus dein fere planus, laevis, sublobatus, pallide cervinus. Stipes
-1.5 x 0.5 mm, dorso-lateralis, in pileo subito expansus, concolor, ad basim disco membranaceo affixus.
Lamellae 3-5 alveoliformes, haud reticulatae, concolores. Caro superficie tenuiter gelatinosa.
Sporae 8.5-10.7 X 3.7-4.5 wz, subcylindricae amyloideae. Basidiola acerosa copiosa. Cheilocystidia -30_
x 5-8 mw, cylindrica vel clavata, hyphis dichophysialibus superantibus. Pileocystidia 20-38 x 5-7 w, in
pellicula gelatinosa immersa. Caulocystidia nulla? Hyphae fibulatae, vix inflatae, 1.5-8 latae, tunicis -2
be Crassis.
Ad petiolos filicium in silva. Peninsula Malayana, Pahang, Cameron Highlands, 4 Oct. 1966, Corner
s.n. typus, in herb. Corner.
Very small, pale fawn. Pileus -2 mm wide, dorsifixed, cupular then almost plane
and horizontal, slightly lobate at the margin, smooth. Stem 0.5-1.5 x 0.5 mm,’
subcylindric, enlarged suddenly into the pileus, with a small membranous disc at
the base. Gills 3-5 primaries -0.4 mm wide, meeting more or less centrally under the
pileus, mostly with a small secondary gill, interstices smooth, the underside of the
pileus appearing alveolate. Flesh 0.2-0.4 mm thick at the base of the pileus, firm,
with a thin gelatinous layer (30-40 uw thick) on stem and pileus.
Spores 8.5-10.7 X 3.7-4.5 jw, white, smooth, subcylindric, obtuse, thin-walled,
pale violaceous amyloid, very abundant. Basidia 23-30 x 7-7.5 uw; sterigmata 4, 5-7
fu long; basidioles acerose to mucronate, abundant; subhymenium narrow, not
corticate. Cheilocystidia -30 x 5-8 yw, cylindric to clavate, thin-walled, smooth,
contents hyaline to vacuolate (not gloeocystidial), eventually with some excrescent,
sparsely branched or lobulate hyphae 1-2 wide inserted among them. Pleurocysti-
dia none. Hyphae monomitic, clamped, scarcely inflated, the cells -230 x 1.5-8 uw
the walls 0.5-2 mw thick, firm, often shortly lobulate, very closely interwoven.
Surface of pileus and stem with a thin gelatinous layer 30-40 wu thick on the pileus,
The genus Panellus Karst. in Malaysia 121
thinner on the stem, composed of 1-1.5 « hyphae laxly branched, not or scarcely
lobulate, with gelatinous walls. Pileocystidia 20-38 x 5-7 yw, as sterile basidia with
somewhat gloeocystidial contents, more or less divergent into the gelatinous layer
from the firm tissue of the flesh, not lobulate, developing from the margin of the
pileus. Caulocystidia apparently none.
On dead fern petioles in the forest. Malay Peninsula.
Pahang, Cameron Highlands 1600 m alt., 4 Oct. 1966, Corner s.n.
Fig. 12. Panellus exiguus. Fruit-body, x 10; spores x 2000; basidia, cheilocystidia, and surface of
pileus, < 500.
Panellus fuscatus sp. nov. Fig. 13
Pileus -3 cm latus, pleuropodalis, reniformis vel flabelliformis, pallide albido-alutaceus, ultimo
fuscibrunneo-pruinosus. Stipes -25 x 3 mm, albus, basi abruptus, pruinoso-furfuraceus. Lamellae
decurrentes confertae angustae albidae, acie obtusa pruinosa, Caro tenax.
Sporae 5-6(-6.5) X 4.7-5.7 , subglobosae amyloideae. Basidiola acerosa nulla. Cheilocystidia 23-55
x 5-20 w, clavata vel ventricosa, etiam subcylindrica, tenuiter tunicata, haud incrustata. Pleurocystidia
nulla. Pileocystidia 20-80 x 5-20 w, clavata vel ventricosa, succo fuscidulo intus. Caulocystidia similia
sed incolorata, haud incrustata. Hyphae fibulatae, inflatae; in stipite -13 yw latae, tunicis tenuibus vel
crassiusculis; in pileo -20 py latae.
Ad lignum in silva. Borneo, mt. Kinabalu, RSNB 5479 typus, in herb. Corner.
Pileus -3 cm wide, pleuropodal, reniform to flabelliform, horizontal, pallid buff
white, finely fuscous brownish scurfy pruinose, not striate. Stem -25 x 3 mm,
lateral, with slightly dilated apex, solid, fibrous, white and white scurfy pruinose
upwards, base abrupt. Gills decurrent, crowded, narrow, c. 40 primaries -2 mm
wide, 2-3 ranks, some dichotomous, white, the obtuse edge pruinose. Flesh 1.5 mm
thick at the base of the pileus, rather tough, especially in the stem, white. Smell
none.
Spores 5-6(-6.5) X 4.7-5.7 uw, white, smooth, ovoid-subglobose, thin-walled,
guttulate opalescent, rather pale bluish amyloid, (bluish black in the mass). Basidia
Ne Gard. Bull. Sing. 39(2) (1986)
27-30 X 6.5-7 w, guttulate-opalescent, brown in Melzer’s iodine; sterigmata 4, 4 uw
long; acerose basidioles none; subhymenium 12-16 mw thick, composed of sub-
agglutinated hyphae 1.5-3 ww wide, interwoven. Cheilocystidia 23-55 x 5-20 yp,
clavate to ventricose or waisted, varying subcylindric, obtuse, thin-walled, colour-
less, not encrusted, as a broad sterile edge to the gill. Pleurocystidia none. Hyphae
monomitic, clamped, inflating, not encrusted, not amyloid; in the stem parallel,
longitudinal, cylindric, the cells 45-600 x 2-13 w, with broad septa and often slightly
thickened walls, mainly fibrillose but subagglutinated in places; in the pileus with
shorter and more inflated cells especially over the gills, the cells 20-120 x 3-20 p,
longitudinal interwoven, fairly compact, thin-walled, often unequally inflated or
with undulate walls; in the gill-trama, as in the pileus, descending; oleiferous
hyphae none. Surface of stem with 3-5 w subagglutinated hyphae and scattered tufts
of cystidia similar to those on the pileus but mostly smaller and colourless, not
encrusted. Surface of pileus with a disrupted palisade or short pile of more or less
clavate to subventricose, obtuse cells 20-80 x 5-20 w, or with 1-3 celled hyphal ends
with similarly inflated cells, with thin or slightly thickened walls, smooth, with pale
fuscous sap, arising from appressed longitudinal hyphae, 3-8 uw wide, at the surface
of the flesh.
On fallen wood in the forest. Borneo.
Mt. Kinabalu, Bembangan River 1800 m alt., 26 Feb. 1964, RSNB 5479.
This species lacks the acerose basidioles, the dichophysial hyphae, and the
granular incrustation of the hyphal walls, but the spores are rather intensely
Fig. 13. Panellus fuscatus. Fruit-body, x 1; spores, 2000; basidia, cheilocystidia, surface of pileus
near the base, x 500.
The genus Panellus Karst. in Malaysia 123
amyloid. Compare P. magnus (p. 143) with more or less centric pilei, narrow
spores, and brown colour in the hyphal walls; both occur in the same forest on mt.
Kinabalu.
Panellus intermedius sp. nov. Fig. 14
Ex integro pallide cervinus, pleuropodalis, lamellato-poroideus. Pileus -15 mm latus, reniformis vel
flabelliformis, opacus, siccus. Stipes -3 x 1-1.5 mm. Lamellae adnatae vel decurrentes, poroideo-
reticulatae, acie albida furfuracea. Caro tenax, haud gelatinosa.
Sporae 8-10 x 4.5-5.7 uw, lacrymiformes, vix amyloideae. Basidiola subacerosa. Cheilocystidia 22-40
x 5-11 w. clavata, diverticulis 1-2 lobulatis evolventia, hyphis dichophysialibus incrustatis superantibus.
Pleurocystidia, pileocystidia et caulocystidia nulla. Hyphae fibulatae, plus minus inflatae, latiores
cellulis subfusiformibus -300 x 16 w, tunicis 1-5 wu crassis sed superficiem pilei versus tenuibus -1.5 u
crassis. Superficies pilei hyphis appressis sublobulatis leniter incrustatis, haud gelatinosis, praedita.
Ad ramulos dejectos in silva. Borneo, mt. Kinabalu, RSNB 5474 typus, in herb Corner.
Entirely pale fawn, pleuropodal, lamellate-poroid. Pileus -10 mm radius, 15 mm
wide, reniform-flabelliform, smooth or minutely cottony, opaque, dry. Stem 1-3 x
1-1.5 mm, lateral, cottony fibrillose to subfurfuraceous. Gills adnate to decurrent,
7-15 primaries 0.8-1 mm wide, 3(-4) ranks, poroid-reticulate with transverse septa
almost as deep as the primary gills, not venulose in the smooth interstices, pale
fawn with thin chalky white edge. Flesh 0.8-1 mm thick at the base of the pileus,
tough, not gelatinous. Smell none.
Spores 8-10 X 4.5-5.7 uw, white, smooth, pip-shaped, varying pale violet-brown
amyloid to almost inamyloid. Basidia 23-28 X 5.5-6.5 jw; sterigmata 4, 3-3.5 yw long:
basidioles subacerose, frequent; many lobulate ramifying hyphae 1-2 yu wide invest-
ing the basidia in the mature hymenium; subhymenium narrow, with 1.5-2.5 uw
interwoven hyphae, not corticate. Cheilocystidia 22-40 x 5-11 mu. more or less
clavate, thin-walled, vacuolate, developing from the distal end 1-2 lobulating
Fig. 14. Panellus intermedius. Fruit-body, x 4; spores, X 2250; basidia, cheilocystidia, and hyphae, x
500. Inset (upper left), var. stenosporus.
124 Gard. Bull. Sing. 39(2) (1986)
processes -15 yw long, also invested with lobulating hyphae as the basidia; thinly
encrusted. Pleurocystidia none. Hyphae monomitic, clamped, more or less inflat-
ing, appearing spuriously trimitic, the wider hyphae with somewhat fusiform cells
-300 < 16 w with firm hyaline walls 1-5 uw thick and narrow fuscous lumen appearing
as Skeletals, the narrower hyphae appearing as binding hyphae (but clamped); in
the stem longitudinal, compactly fibrillar; in the pileus longitudinal as in the stem
but in the upper part of the pileus laxly interwoven with thinner walls -1.5 w thick
and wide lumen; in the gill-trama as in the stem, more or less descending, with
many narrow interweaving hyphae. Surface of pileus and stem without cystidia but
with 1-2 yw processes becoming somewhat lobulate and ramified though not in a
dense layer, thinly encrusted or scarcely at all, not gelatinous.
On sticks in the forest. Borneo.
Mt. Kinabalu 1700-1800 m alt.: 6 Feb. 1964, RSNB 5243 (immature); 26 Feb. 1964, RSNB 5474; 28
Feb. 1964, RSNB 5539.
This species is exactly intermediate between Dictyopanus and Panellus. It seems
not to begin sporing until the fruit-bodies are nearly fully grown. Compare the
poroid P. /uminescens on dead palm-leaves.
var. stenosporus var. nov. Fig. 14 (inset)
Sporae 7-9 X 3.5-4.5 uw, amygdaliformes. Cheilocystidia 20-38 x 8-14 uw, clavata vel subglobosa.
Hyphae in stipite 2-6 wu latae, in pileo -9 w.
Ad ramos dejectos in silva. Borneo, mt. Kinabalu, 20 April 1964, Corner s.n. typus, in herb. Corner.
Pileus -10 mm in radius, white to fawn-brown. Stem -1.5 < 1 mm, lateral, slight.
Gills whitish. Flesh -1 mm thick at the base of the pileus, not gelatinous.
Spores 7-9 X 3.5-4.5 uw, amygdaliform or mango-shaped. Cheilocystidia 20-38 x
8-14 uw, widely clavate to subglobose. Hyphae narrower and less thick-walled; 2-6 u
wide in the stem, with walls -2.5 thick, without stout skeletal-like cells; in the pileus
2-9 w wide. Surface of pileus with appressed interwoven hyphae without external
processes, not specially modified.
On sticks in the forest. Borneo.
Mt. Kinabalu, Mesilau 1700 m alt., 20 April 1964, Corner s.n.
With narrow hyphae, this variety shows the approach to the poroid state of
Dictyopanus, but the difference in the spores shows that they are not a guide to
specific affinity between the lamellate and the poroid species of Panellus.
Panellus longinquus (Berk.) Singer
Singer et Digilio, Lilloa 25 (1952) 121; Libonati-Barnes and Redhead, Mycotaxon 20 (1984) 205-212.
? Panellus diversipes (Berk.) Pegler, Austral. J. Bot. 13 (1965) 329.
Pileus -3 cm wide, horizontal, flabelliform, smooth, subviscid, pale bistre sub-
ochraceous, drying finely villous at the base. Stem lateral, very short, white villous.
Gills subdecurrent, subdistant, c. 15 primaries -5 mm wide, 4 ranks, pale bistre
cream, edge entire. Flesh firm dry, white. Smell none.
Spores 7.5-10 x 3.3-4 uw, white, smooth, subcylindric, thin-walled, aguttate, pale
blue-black amyloid. Basidia 30-38 X 5.5-6 mw; sterigmata 4; racemose basidioles
none; subhymenium composed of 2-3 «x interwoven hyphae, not corticate. Cystidia
none, gill-edge fertile. Hyphae monomitic, inflating, clamped, the cells 50-500 x
3-20 uw, flexuous, very variable, mostly with broad septa, not tapered, the walls
slightly thickened and firm in the older tissue. Surface of pileus with a very thin
superficial gelatinous layer 20-30 uw thick, composed of hyphae 1.5-2.5 mw wide,
mostly appressed and radiating, with scattered processes to the exterior or amassed
The genus Panellus Karst. in Malaysia 125
here and there into patches of rudimentary hymenium, the processes subcylindric
to subclavate -20 x 1.5-3 yw, simple or with a few short lobes, extending at the base
of the pileus -70 uw as the villous layer. Hyphae of gill-trama -10 u wide, descending,
with slightly thickened subgelatinous walls near the subhymenium.
On a dead trunk in forest. Queensland.
Mt. Lamington, 21 June 1964, Corner s.n.
This is my description of an Australian collection, which I include here because
the species may well be widely distributed in the Western Pacific. It was described
from South America and recently recorded from North America. The Australian
collection agrees with the descriptions of the American except that the gills are
wider and the pileus lacks a pinkish colour. Acerose basidioles were not mentioned
for the American collections.
Panellus parvulus sp. nov. Fig. 15
Pileus -7 mm latus, pleuropodalis vel sessilis, opacus albus. Stipes -0.5 mm longus et latus, vel nullus.
Lamellae e basi pilei radiantes, 7-12 primariae, haud dichotomae, albae, acie pruinosula. Caro haud
gelatinosa.
Sporae 6-7.5 X 4-5 py, late ellipsoideae, amyloideae. Basidiola subacerosa. Cheilocystidia 25-50 x 5-8
ft, subclavata vel subventricosa, intus oleaginosa, hyphis dichophysialibus superantibus. Pleurocystidia
nulla. Pileocystidia 28-58 x 12-15 uw, clavata, intus oleaginosa, vix in strato hymeniiformi, basim pilei
versus hyphis dichophysialibus superantibus, leniter incrustata. Hyphae fibulatae, in stipite basique pilei
-20 w latae, in lamellis 1.5-6 pw latae.
Ad folia emortua bambusarum in silva. Insulae Solomonenses, Guadalcanal, RSS 688 typus, in herb.
Corner.
Fig. 15. Panellus parvulus. Fruit-body, < 3; spores, X~1500; cheilocystidia and surface of pileus, x
600.
Pileus -7 mm wide, lateral, sessile, or with a very short stem -0.5 mm long and
wide, white, opaque to substriate: margin minutely pruinose. Gills 7-12 primaries,
2-3 ranks, radiating from the base of the pileus, not dichotomous or veined at the
base, white, edge pruinulose. Flesh 0.2-0.3 mm thick at the base of the pileus, not
gelatinous.
Spores 6-7.5 x 4-5 uw, white, smooth, broadly ellipsoid, pale bluish amyloid.
Basidia 18-25 x 6.5-7.5 uw, with oleaginous contents; sterigmata 4, 4-5 mw long;
subacerose basidioles numerous, not conspicuous; subhymenium very narrow, with
1.5-2.5 w interwoven hyphae. Cheilocystidia 25-50 x 5-8 mw, subclavate, obtuse,
varying subventricose, thin-walled, smooth or thinly encrusted, with oleaginous
cloudy contents, as a conspicuous sterile edge to the gill, becoming overgrown with
dichophysial hyphal ends with stalks 1-3 « wide and fine short processes. Pleurocy-
126 Gard. Bull. Sing. 39(2) (1986)
stidia none. Hyphae monomitic, clamped, inflating in the stem and base of pileus
with cells 30-160 x 1.5-20 pw, thin-walled, longitudinal interwoven, narrower to-
wards the margin of pileus; in the gil/l-trama 1.5-6 w wide, not inflated, more or less
descending. Pileocystidia 28-58 x 12-15 w, clavate, rarely with narrowed apex, with
slender stalk 1-2 uw wide, thin-walled, smooth or thinly encrusted, containing dense
smeary-oleaginous cytoplasm, erect or appressed, abundant and conspicuous but
not in a compact palisade, becoming overgrown at the base of the pileus with
dichophysial hyphae, their long, sparingly branched, filiform arms c. | uw wide. All
the more superficial hyphae of pileus and gill-edge with granular incrustation
soluble in potash. Hyphae and contents of basidia and cystidia merely yellow-
brown in Melzer’s iodine.
On dead bamboo-leaves in forest. Solomon Islands.
Guadalcanal, Gallego, 11 July 1965, RSS 688.
This is easily known from the conspicuous pileocystidia which, if immersed,
would be gloeocystidia. It seems close to the South American P. nubigenus Singer
(1969), also on bamboo remains, but with the pileus becoming brownish, the spores
slightly larger, and the pileus with a thin gelatinous layer below the superficial mat
of dichophysial hyphae.
Panellus pendens sp. nov.
Pileus sessilis dorsifixatus, cyphelliformis, -15 mm latus, laevis substriatus, albidulus dein sub-
ochraceus et subroseolus. Lamellae distantes angustae, haud venosoreticulatae, Caro hygrophana, haud
gelatinosa.
Sporae 4.3-5 x 3.5-4 uw, subglobosae, intense amyloideae. Basidiola acerosa nulla. Cystidia nulla.
Hyphae fibulatae, 2-5 (-6) mw latae, plus minus crasse tunicatae, haud incrustatae; in lamellis saepe
oleiferae ut gloeocystidia, subagglutinatae. Superficies pilei hyphis oleiferis 5-9 mw latis, haud dextri-
noideis, in strato tenui in crassitudine hyphae unae constructa.
Ad ramos delapsos in silva. Borneo, mt. Kinabalu, RSNB 5448 typus, in herb. Corner.
Sessile, cyphelloid, dorsifixed, pendent, pallid white becoming pale dingy buff,
the upperside drying pale pinkish tan. Pileus -15 mm wide, smooth, substriate.
Stem none. Gills radiating from a centre below the attachment of the pileus,
distant, narrow, 9-16 primaries (.5-1 mm wide, 2(-3) ranks, not veined. Flesh
0.5-0.8 mm thick in the centre of pileus, 0.1-0.2 mm thick at the margin, firm,
rather tough, hygrophanous, not gelatinous. Smell none.
Spores 4.3-5 x 3.5-4 uw, white, smooth, ovoid or subglobose, thin-walled, indigo-
black amyloid. Basidia 13-15 x 5-5.5 yw, short, squat, somewhat waisted; sterigmata
4; no acerose basidioles; subhymenium very narrow, 5-8 w thick, with hyphae
1.5-2.5 yw wide. Cystidia none, or with sterile basidia on the apparently sterile
gill-edge. Hyphae monomitic, clamped, not inflating, 2-5(-6) w wide, more or less
thick-walled, not amyloid; in the pileus densely interwoven and impossible to tease
apart, many thin-walled but others with walls 1-1.5 mw thick, the cells 70-400 yw long,
flexuous, often kinked resembling bits of skeletal hyphae but with distant clamps
and occasionally branched, also with scattered more or less radiating oleiferous
hyphae 3-7 uw wide with dense contents not staining with Melzer’s iodine; in a thin
layer, c. 30 uw thick, above the gills, with thin-walled radiating hyphae and oleifer-
ous hyphae descending into the gills; in the gill trama with thin-walled hyphae
descending obliquely towards the margin of the pileus and with many subparallel
oleiferous hyphae (as gloeocystidia) reaching almost to the gill-edge with tips
tapered to 2-3 yw wide, the tissue of the gill-trama subagglutinated (not gelatinous).
Surface of pileus with a layer (one hypha thick) of appressed radiating oleiferous
hyphae 5-9 uw wide with dense contents (not staining with Melzer’s iodine), with
agglutinated walls, as a tough layer with many affixed spores; round the basal
The genus Panellus Karst. in Malaysia 127
attachment of the pileus, with many excrescent hyphae 1.5-3 « wide, clamped, with
slightly thickened brownish cells.
On a rotten fallen branch in the forest. Borneo.
Mt. Kinabalu, Bembangan River 1700 m alt., 25 Feb. 1964, RSNB 5448.
The spores seem too intensely amyloid for Panellus. The agglutinated oleiferous
hyphae on the pileus and the absence of incrustation are also differences. Intensely
amyloid spores are recorded for the South American P. mirabilis Singer.
Panellus sublevatus sp. nov. Fig. 16
Albus exiguus. Pileus -3 mm latus, pleuropodalis vel subsessilis, cyphelliformis dein reniformis.
Lamellae 3-5, haud reticulatae, acie pruinosofurfuracea. Caro mollis, haud gelatinosa.
Sporae 7-8(-9) xX 5-6 w, ellipsoideae amyloideas. Basidiola subacerosa sparsa. Cheilocystidia et
pileocystidia vix evoluta, ut hyphae dichophysiales incrustatae. Pleurocystidia nulla. Hyphae fibulatae
1.5-5 uw latae, sed ad basim pilei cellulis hypodermatis 16-30 x 80-16 w.
Ad caudices bambusarum in silva. Borneo, mt. Kinabalu, RSNB 8407A typus, in herb. Corner.
White, very small. Pileus -3 mm wide, lateral, more or less sessile, at first
semi-cupular and cyphelloid, then more or less reniform, sinuato-sulcate, slightly
scurfy pruinose. Gills distant, 3-5 primaries 0.3-0.5 mm wide, 1-2 ranks, interstices
smooth, edge scurfy pruinose. Flesh c. 100 uw thick in the pileus (excluding the
scurfy villous layer 50-80 yu thick), soft, not gelatinous.
Spores 7-8(-9) X 5-6 uw, white, smooth, ellipsoid, obtuse, pale indigo amyloid.
Basidia 20-27 X 6.5-8 yw; sterigmata 4, 5-7 w long: subacerose basidioles rather
sparse; subhymenium slight, with 1.5-2.5 « interwoven hyphae, not gelatinous.
Cheilocystidia as dendritic hyphal ends -50 uw long, the stalk 1.5-3(-4) uw wide,
granular-encrusted, forming a broad sterile edge to the gill with a few sterile
basidia. Pleurocystidia none. Hyphae monomitic, clamped, thin-walled, 1.5-5 u
wide, short-celled, but with inflated cells 16-30 x 8-16 uw in a hypodermal layer near
the base of the pileus (? as a vestige of the stem). Surface of pileus composed of
masses of granular-encrusted dendritic hyphae 50-80 y long, as the cheilocystidia,
with a few scattered sterile basidia 20-40 < 6-7 uw or cylindric hyphal ends -50 x 4-
5 w; no definite pileocystidia.
Fruit-bodies begin to spore when c. | mm wide.
On dead stems of a climbing bamboo in the forest. Borneo.
Mt. Kinabalu, Mesilau 1700 m alt., 22 April 1974, RSNB 8407A.
Save §
,S2
ot fae We
PAN ns s 3+. ‘
+, _syra Este F mys v
ig. Oe aoy
re sey “ee > we Mann = pipes
~ + Pe “—\- > +0 oi! .
ae A We Aa FATS gee — >
SW = ch : ce
Seas 4,7) 33. aS
\ or oH p S 7
pee was t 4° be ‘ =
aa <a aes s. ne Pi F = a “ '
“3 ean ts ote Thy: : A : Vor } Se.
BIS “et ie ena gp TE co ag . 3° D ‘ - @
tee Se. See PES Slo asefa Ye a weess . 3 As “+
ce ae natn OE ee ae Vatie % > ie }
iy ey sae aa 2-3 Vey ” ‘ as : 3 . ~
Rew 550" i EP et aS % el 4 “Aa, X- -
Sle seatte ves pF soe a = >
SS az3 MAF hy? SO, a
Fig. 16. Panellus sublevatus. Fruit-body, x 10; spores, x 2000; basidia and cheilocystidia, x 600.
128 Gard. Bull. Sing. 39(2) (1986)
Key to the poroid species of subgen. Panellus in Malesia
1. Spores subglobose, more than 6 w wide.
2. Spores 13-18 x 12-16 w. Pileus -22 mm wide, sessile, white. Pores reaching 1 mm wide. Gloeocys-
tidia on pore-edges and pileus. On fallen branches. Borneo ................... P. megalosporus p. 133
2. Spores smaller. Pileus white to pale fawn. Pores 0.2 mm wide.
3. Spores 6.5-9.5 x 6-8.5 w. Pileus -17 mm wide. Gloeocystidia on pore-edges and pileus. On
OOD db asd aie Poles Vince bin cys sin vay cD Retr PE ois 0 8 Deed wily EE a P. orientalis p. 135
3. Spores 9-12.5 x 8-11 mw. Pileus-3 mm wide. Gloeocystidia none. On dead bamboo. Solomon
BSL roca sous fe bculanes Cheon net eeee eee ara neck ay aoe nee ee P. brunneifavolus p. 130
1. Spores ellipsoid to allantoid, generally less than 6 wu wide.
4. Spores generally more than 6 uw long. Pileus 1-5 mm wide. On grasses, bamboo or palms.
5. Cheilocystidia clavate, not gloeocystidial. Pileus -4 mm wide.
6. Flesh entirely subgelatinous. Acerose basidioles absent. Cheilocystidia developing lobulate
processes. Spores 7-8.5 X 4.5-6 w, scarcely amyloid. Pileus subsessile P. albifavolus p. 128
6. Flesh dry. Acerose basidioles present. Cheilocystidia not developing processes.
7. Spores 6-7.5 X 4.5-5.5 yw. Cheilocystidia 22-35 x 7-18 uw. Pileus subsessile, white to pale
fawn and pmkish. On bantboo, 2). 5: i.c.6: <2 Sees ce meee P. sublamelliformis p. 137
7. Spores 7.5-9 X 4-4.5 yw to 9-13 X 4-6 pw, ellipsoid. Cheilocystidia 30-55 x 7-10 wu. Pileus
white to pale buff. On.palins....seuc) 5.4.00. ee ee P. luminescens p. 132
5. ‘Cheilocystidia gloeocystidial, subcylindric, or none.
8. Cheilocystidia gloeocystidial. Pileus pale fawn, with a thin gelatinous pellicle. Spores 7-9 x
3.5-4:5 ps. Ombamboo: .\ 205 es ee, ene ee P. bambusifavolus p. 129
8. Cheilocystidia none (? very soon obscured).
9. Acerose basidioles present. Spores 6-9 X 3.5-5.5 w. Pileus white to pale fawn. Flesh dry.
OM STASSES os. fs 12. Da PoPae tee Beng Pe eat ee ca P. copelandii p. 131
9. Acerose basidioles absent. Fruit-bodies white. Spores 6-7.7 X 3-4 w. Pileus -1.5 mm
wide, with 3-15 pores. Flesh dry. On bamboo, Solomon Isl. ......... P. pauciporus p. 136
4. Spores less than 6 uw long.
10. Spores 4.5-6 X 2.5-3.3 yw, Acerose basidioles absent. Cheilocystidia not distinct, pore-edge
and pileus with unbranched spinulose hyphal ends 3-6 uw wide. Pileus -3 mm wide, white. On
dicotyledonous leaves. <::scxsceiher case tenee oleae eae ee ee P. hispidifavolus p. 132
10. Spores 3-4 uw long. Acerose basidioles present.
11. Spores 0.8 « wide, allantoid. Pileus -4 mm wide, fawn brown, dorsifixed. Cheilocystidia
7-12 p wide, clavate. On twigs: Borneo? 7 inng 2.20, b.) apanains aligns! eke P. microsporus p. 134
11. Spores 2-3 w wide, ellipsoid. Cheilocystidia 3-5 4 wide, cylindric.
12. Pileus -12 mm wide or more, fawn tan. Cheilocystidia not or slightly gloeocystidial ...
san ceesiesislnd dled te tuclewahudellt eae oMiad Oehd te eatete Aa ft anne Se eae P. pusillus p. 136
12. Pileus -3 mm wide, white then pale fawn. Cheilocystidia strongly gloeocystidial ........
1nd dna qps garivivas ud dle cla sales Gatlin ta te Date tana ae ae P. gloeocystidiatus p. 131
Panellus albifavolus sp. nov. Fig. 17
Pileus -3 mm latus, subsessilis, reniformis, pruinosulus, albus. Tubi -0.4 mm longi; poris c. 200 yp latis,
albis. Caro subgelatinosa.
Sporae 7-8.5 x 4.7-6 yw, late ellipsoideae, haud vel vix amyloideae. Basidiola acerosa nulla. Hyme-
nium in aetate mucilaginosum. Cheilocystidia 15-30 x 3-10 «, ex apice diverticula lobulata, -10 longa
emittentia, hyphis dichophysialibus nullis. Hyphae fibulatae, 1.5-3 w latae, tunicis gelatinosis. Pileocys-
tidia -38 w longa ut cheilocystidia, leniter incrustata, hyphis dichophysialibus nullis.
Ad ramulos emortuos et petiolos palmae in silva. Borneo, RSNB 5367 typus, in herb. Corner; Insulae
Solomonensis.
Pileus -3 mm wide, subsessile, reniform, horizontal, minutely pruinose, white,
opaque. Tubes -0.4 mm long, white, pores 180-200 w wide, minute, white, with
pruinose edges. Flesh 0.3-0.4 mm thick, entirely subgelatinous, drying horny.
Spores 7-8.5 x 4.7-6 w, white, smooth, broadly ellipsoid, obtuse, thin-walled,
not amyloid or very slightly. Basidia 24-30 x 6.5-8 yw; sterigmata 4, 3-3.5 uw long.
Hymenium without acerose basidioles; subhymenium narrow, interwoven; old
hymenium with the basidia half-immersed in a more or less structureless firm
ee
Cons noanence
Fig. 17. Panellus albifavolus. Fruit-body, X 6; spores, X 1800; pileocystidia, x 600.
mucilage. Cheilocystidia as the pileocystidia but shorter, 15-30 uw long, without
excrescent hyphae. Pleurocystidia none. Hyphae 1.5-3 uw wide, clamped, with
entirely gelatinous walls, interwoven. Surface of pileus as a loose palisade of
subclavate, erect or decumbent, pileocystidia 18-38 x 3-10 uw, the apex with
lobulate processes -10 uw long, sometimes with 2-3 distal lobes, thin-walled, thinly
encrusted, not amyloid, without excrescent hyphae.
On rotten sticks and dead palm-petioles in the forest.
BORNEO. Mt. Kinbabalu, Bembangan River 1700 m alt., 19 Feb. 1964, RSNB 5367. — SOLOMON
ISLANDS. Guadalcanal, Gallego 500 m alt., 8 July 1965, RSS 603.
This seems to approach Campanella with inamyloid spores.
Panellus bambusifavolus sp. nov. Fig. 18
Pileus -3 mm latus, pleuropodalis reniformis, pallide cervinus. Stipes -1 xX 0.5 mm, pileo concolor.
Tubi -0.5 mm longi; poris 150-200 yu latis, albocretaceis. Caro sicca sed ad pileum pellicula gelatinosa
praedita.
Sporae 7-9 x 3.5-4.5 uw, subcylindricae, amyloideae. Basidiola acerosa copiosa. Cheilocystidia et
pileocystidia 3-5 yw lata, cylindrica ut gloeocystidia, hyphis dichophysialibus gelatinosis superantibus.
Hyphae fibulatae, 1.5-4 yu latae, tunicis -1 crassis, incrustatae; in stipite -12 w latae, tunicis -2 u crassis.
Superficies pilei pellicula 30-50 uw crassa gelatinosa, ex hyphis 1-2 wu latis ramosis apicibus echinulatis,
composita, cystidis immersis.
Ad caules bambusarum in silva. Insulae Solomonenses, San Cristobal, RSS 898 typus, in herb.
Corner.
Pleuropodal with short stem. Pileus -3 mm wide, reniform, convex, opaque,
smooth, pale fawn. Stem -1 X 0.5 mm, distinct, pale fawn. Tubes 0.3-0.5 mm long;
pores 150-200 wu wide, rounded, chalky white. Flesh 0.3-0.4 mm thick at the base of
the pileus, 0.1-0.15 mm near the margin, with a thin gelatinous pellicle.
Spores (6.5-)7-9 X 3.5-4.5 , white, smooth, subcylindric, obtuse, pale bluish
amyloid. Basidia 24-27 x 6.5-7 mw; sterigmata 4; basidioles acerose, abundant;
subhymenium narrow, not corticate, composed of 1.5-2.5 4 hyphae. Cheilocystidia
as cylindric gloeocystidia 3-5 x wide, as on the pileus but shorter, covered by a thin
gelatinous pellicle of dichophyses 1-2 ~ wide with somewhat echinulate tips, not
amyloid. Hyphae 1.5-4 « wide, clamped, with hyaline walls -1 « thick, but in the
stem with scattered stouter hyphae -12 mw wide with walls -2 w thick, granular
encrusted and often with minute protuberances, closely interwoven, not gelatinous.
Surface of pileus covered by a thin gelatinous pellicle 30-50 « thick, composed of
1-2 uw hyphae irregularly branched and ending in a close membrane of 0.5-1 4,
minutely echinulate, hyphal ends, with immersed, suberect or oblique to decum-
bent cylindric gloeocystidia -85 x 3-5 w, very abundant on young pilei but becom-
129
—_—_ ce
mai! 100.22»
Aaa ~~) onl= Fad ot) js
EAA eels bea
RSA (Cp a
~~ ~ et ry 13
a 7 yp 7 ¢,
Fig. 18. Panellus bambusifavolus. Fruit-body, x 10; spores, x 2000; surface of pileus, x 1000.
ing scarce and inconspicuous, even vacuolate and hyaline, in mature pilei, often
with the oleaginous contents shortly retracted from the tip.
On dead bamboo in the forest. Solomon Islands.
San Cristobal, Warahito River, low level, | Aug. 1965, RSS 898.
The surface of the pileus has the reduced construction of P. gloeocystidiatus, and
the gelatinous pellicle forms on the pore-edges.
Panellus brunneifavolus sp. nov. Figs. 8b, 19
Pileus -3 mm latus, dorsifixus, subsessilis, reniformis, laevis, albus dein pallide cervinobrunneus. Tubi
-0.4 mm longi; poris 150-200 yw latis, albocretaceis. Caro subgelatinosa.
Sporae 9-12.5 x 8-11 uw, subglobsae vel ovoideae, amyloideae. Basidiola acerosa copiosa. Cheilocysti-
dia 18-45 x 3-4.5 w, subcylindrica, tunicis leniter incrassatis, haud oleaginosa, hyphis dichophysialibus
superantibus. Hyphae fibulatae, 1.5-5 w latae, tunicis subgelatinosis, subagglutinatae. Pileocystidia ut
cheilocystidia, hyphis dichophysialibus superantibus.
Ad bambusam emortuam in silva. Insulae Solomonenses. Guadalcanal, 2200 m alt., RSS /63/ typus,
in herb. Corner.
Dorsifixed, almost lateral, convex, reniform, white then pale fawn brown. Pileus
-3 mm wide, smooth, opaque. Stem practically none. Tubes -0.4 mm long; pores
150-200 yw wide, rounded, with chalky white edges. Flesh c. 1 mm thick at the base
of the pileus, subgelatinous.
Spores 9-12.5 x 8-11 mw, white, smooth, subglobose or ovoid, very pale
violaceous amyloid. Basidia 36-40 x 12-15 pw, widely clavate with oleaginous
contents (dark brown in Melzer’s iodine); sterigmata 4, 6-7.5 w wide at the base;
basidioles acerose, abundant but inconspicuous, hyaline, 3-7 uw wide, shorter than
the basidia; subhymenium narrow, not corticate. Cheilocystidia 18-45 x 3-4.5 p,
subcylindric, hyaline, smooth, with slightly thickened wall, often sparse, not gloeo-
cystidial, becoming covered by excrescent, laxly to profusely branched, dichophy-
sial hyphae 1-2 w wide with stalks 1.5-2.5 uw wide, thinly encrusted, not amyloid.
Hyphae 1.5-5 ww wide, clamped, with hyaline subgelatinous walls 1-2 mw thick,
compact, almost agglutinated, not amyloid. Surface of pileus with pileocystidia like
130
a
Cr
Fig. 19. Panellus brunneifavolus. Fruit-body, < 6; spores, x 2000; basidia and surface of pileus, x 600.
the cheilocystidia but becoming scattered and often decumbent, covered by shortly
excrescent dichophysial hyphae as a layer c. 40 mm thick.
On dead bamboo in the forest. Solomon Islands.
Guadalcanal, Popomanasiu 2200 m alt., 26 Oct. 1965, RSS 1631.
Panellus copelandii (Pat.) Burdsall et Miller
Nova Hedwigia Beih. 51 (1975) 88.
Dictyopanus copelandii Pat., Elmer Leaflets of Philippine Botany 6 (1914) 2254; Kobayasi, Bull. Nat.
Sci. Mus. Tokyo 6 (1963) 363.
Pileus -3 mm wide, pleuropodal, reniform to orbicular, pruinose, white. Stem 3-6
x 0.25-0.5 mm, longer than the pileus, white. Pores slightly radially elongate,
120-180 yw in the long diameter, lamellately decurrent (Kobayasi), whitish rufous.
Flesh not gelatinous.
Spores 6.5-9 X 3.5-5.5 uw, 6.8-7.8 X 3.5-4.2 uw (Kobayasi), 5-6 x 4-5 pw (Pat.),
ovoid, bluish amyloid. Basidia 20-27 x 7-9 w; acerose basidioles present. Cheilocy-
stidia and pileocystidia not seen. Hyphae 3-6 wu wide, with swellings -9 w, irregularly
thick-walled. Surface of pileus, stem, and pore-edges developing a mat of lobulate
and echinulate dichophysial hyphae; no gloeocystidia (Kobayasi).
On dead grass leaves and stems. Philippine Islands.
This description is a summary of previous ones. Patouillard gave the fruit-bodies
as whitish with whitish rufous pores. That they should be cinnamon-rufous or hazel
according to Burdsall and Miller is misleading and must refer to old, possibly
poisoned, herbarium specimens. The species may be wide-spread in Malesia and
may connect with those on bamboos in the Solomon Islands. P. /uminescens on
palm-remains has larger spores and pores that are not at all lamelliform. Compare
the description of Favolaschia minima (Jungh.) Singer (Lloydia 8, 1945, 200).
Panellus gloeocystidiatus (Corner) comb. nov. Figs. 4, 5, 8f
Dictyopanus gloeocystidiatus Corner, Trans. Brit. mycol. Soc. 37 (1954) 258, fig. 1, 2; Kobayasi, Bull.
Nat. Sci. Mus. Tokyo 6 (1963) 359, fig. 2, 3, pl. 49; Rept Tottori Mycol. Inst. (Japan) 10 (1973) 346.
White then pale fawn, pleuropodal. Pileus -3 mm wide, discoid, scurfy, opaque.
Stem 0.2-1 x 0.1-0.5 mm, lateral, white pruinose. Tubes -0.3 mm long; pores c.
100 y wide, chalky pruinose. Flesh not gelatinous.
Spores 3-3.5 xX 2-2.5 uw, white, smooth, ellipsoid, pale bluish grey amyloid.
Basidia 13-15 x 4 w, with 4 sterigmata 3 w long; acerose basidioles abundant;
subhymenium not granular encrusted. Cheilocystidia -40 x 3-5 mw, subcylindric
gloeocystidia obtuse, with firm walls. Hyphae clamped, 2-5 mw wide, with firm,
slightly thickened and even subdiffluent walls. Surface of pileus with gloeocystidia
-9)) x 3-5 w overgrown with lobulate dichophysial hyphae 1-2 uw wide.
On rotten wood in the forest. Borneo.
13]
WG Gard. Bull. Sing. 39(2) (1986)
Mt. Kinabalu, Mahmud River 1300 m alt., 7 Aug. 1961, RSNB 1656.
This is the description of a Bornean collection. The species has been recorded
hitherto from south Japan, the Bonin Islands, and New Guinea. According to
Kobayasi, the pores are luminous and may be somewhat radially arranged, even
slightly radially elongate. Burdsall and Miller considered that it might be reduced
to P. pusillus, but neither Kobayasi nor I am convinced that P. pusillus has
gloeocystidia. The fruit-bodies of P. gloeocystidiatus are also much smaller.
Fig. 20
Panellus hispidifavolus sp. nov.
Albus, superficie pruinoso-cretacea. Pileus -3 mm latus, pleuropodalis, reniformis. Stipes -0.3 x 0.2
mm, brevissimus. Tubi -350 u longi; poris 150-180 yw latis. Caro sicca.
Sporae 4.5-6 X 2.5-3.3 w, ellipsoideae amyloideae. Basidiola acerosa nulla. Cheilocystidia, caulocysti-
dia et pileocystidia ut hyphae cylindricae, haud ramosae, -150 x 3-6 w, tunicis incrassatis, spinulis -1.5 x
0.5-1 yj echinulatae, etiam apices versus diverticulis obtusis -3 xX 1.5 w, leniter incrustata; hyphis
dichophysialibus nullis. Hyphae fibulatae, 2-12 yw latae, tunicis tenuibus.
Ad folia emortua Tristaniae (Myrtaceae) in silva. Borneo, mt. Kinabalu, 1800 m alt., RSNB 54/5
typus, in herb. Corner.
White, with finely chalky surfaces. Pileus -3 mm wide, pleuropodal, convex,
reniform, Opaque, puberulous. Stem -0.3 x 0.2 mm, lateral, very short. Pores
minute, 150-180 uw wide, -350 uw deep. Flesh 120-200 yw thick, dry.
tng =D
Qor4 bee =
0,029 ZeyS
pe a) 2
sie 2
lox ate erie
ca ae 7 es >
- =e e LE
ae >aa ~ a
faa ols Sie
ay > =k 4S -_—
— ——=c ya
. 77 =; >
shoes ee
—~ «6 ale eels et
<< nigelss aw st
=e soa «> me
= ~~ Be . ay
ee nas t< re
Tae
acdlee ‘
ee “ay hire t
FADE apa sw
Panellus hispidifavolus. Fruit-body, x 6; spores, x 3400; basidia and one hyphal tip on the
pileus, x 600; 3 hyphal tips on the pileus, x 1200.
Spores 4.5-6 X 2.5-3.3 w, white, smooth, ellipsoid, very pale violaceous amyloid.
Basidia 11-15 x 5.8-6.5 uw, widely clavate, short; sterigmata 4; acerose basidioles
none. Cheilocystidia as encrusted spinulose hyphal ends as on the pileus. Pleurocy-
stidia none. Hyphae 2-12 uw wide, clamped, thin-walled, cylindric, narrow and
interwoven in the pileus. Surface of pileus and stem hispidulous with variously
projecting, often curved, cylindric, spinulose, unbranched hyphal ends 70-150 x
3-6 , with thickened walls densely set with minute spinous processes -1.5 X 0.5-1 wu
and often with several stouter obtuse processes -3 X 1.5 w at the tips (as a cluster of
abortive sterigmata), in places thinly encrusted, not amyloid.
On a dead leaf of Tristania (Myrtaceae) in the forest. Borneo.
Mt. Kinabalu, Bembangan ridge 1800 m alt., 22 Feb. 1964, RSNB 5415.
Remarkable for the surface-structure, stout basidia, and absence of basidioles.
Figs. 2, 8c, 8e
Fig. 20.
Panellus luminescens (Corner) comb. nov.
Dictyopanus luminescens Corner, Mycologia 42 (1950) 423.
Pileus 2-4 mm wide, pleuropodal, occasionally mesopodal, convex then flattened
and reniform or flabelliform, opaque white becoming pale alutaceous or pale buff,
The genus Panellus Karst. in Malaysia 133
finely chalky pruinose, drying or ageing pale greyish ochraceous and minutely
cracked; margin incurved at first, finally flattening, always making the distal wall of
the marginal pores. Stem 0.1-1.3 x 0.1-0.5 mm, short, distinct, white, finely chalky
pruinose, brownish at the base, pale ochraceous in age. Tubes -350 w long,
200-250 ye wide, distinctly delimited from the stem, not decurrent, white: pores
70-150 w wide, with dissepiments 50-80 mw thick (including the hymenium) but
appearing wider between the pores, round, white chalky, drying pale yellowish.
Flesh 100-250 uw thick at the base of the pileus, dry, rather firm, rather tough in the
stem, white. Fruit-bodies entirely green phosphorescent, but not so in some collec-
tions.
Spores 9-13 X 5-6 uw, mostly 10-11.5 uw long, in some collections 9-11 x 4-5 pw, in
others 7.5-9 x 4-4.5 uw, white, smooth, elongate pip-shaped to oblong ellipsoid,
contents finely granular, not guttate, pale blue amyloid. Basidia 20-28 x 9-10 yw, or
18-25 X 7.5-8.5 wu with the smaller spores, broadly cylindric-clavate, monomorphic,
forming a continuous lining to the tubes; sterigmata 4, 5-6 uw long; basidioles
subacerose, numerous. Cheilocystidia 30-55 x 7-10 mw, narrowly clavate, thin-
walled, as a sterile edge to the pores, becoming overgrown with excrescent
dichophyses as on the pileus, and densely granular encrusted. Pleurocystidia none.
Hyphae monomitic, clamped, often branching from the clamp, encrusted with fine
granular and crystalline matter soluble in dilute potash, mostly 1.5-4 uw wide and
thin-walled; in the stem with some longitudinal fusiform inflated cells 100-250 x
6-20 uw, with walls -7 w thick, singly or in irregular rows of 2-4 cells, often with
enlarged clamps, at the base of the stem not inflated but subagglutinated with
slightly thickened brownish walls; in the pileus and dissepiments uninflated, thin-
walled, intricately interwoven, but some radiating or longitudinal, more or less
contiguous; hyphal tips at the growing margins 1.5-3 uw wide with cells 10-20 u long
on delimitation. Surface of pileus and stem with a palisade of cystidia like the
cheilocystidia becoming disrupted and overgrown by excrescent and heavily en-
crusted dichophysial hyphae, their ends set with many intricate diverticula 1.5-7 x
0.5-1.5 yw, the pileocystidia also becoming intricately diverticulate.
Development gymnocarpic, direct, with short primordial shaft and epinastic
margin to the pileus.
On dead leaf-sheaths and petioles of various palms (Arenga, Calamus, Pinanga,
Rhapis). Quite common, Malay Peninsula and Borneo.
Most collections of this fungus which I studied in Singapore had the larger
spores, but a few had the smaller though no other difference could be detected. The
Bornean collection, RSNB 556, on Calamus, was not luminous; it had the smaller
spores.
Panellus megalosporus sp. nov. Fig. 8a
Albus, plus minusve sessilis. Pileus -22 mm latus, reniformis, cretaceo-subpruinosus. Tubi -1.5 mm
longi; poris 0.2-0.3 mm latis. Caro sicca. Haud luminescens.
Sporae 13-18.5 x 12-16 uw, subglobosae vel late ellipsoideae, amyloideae. Basidia 60-120 x 15-20 4;
sterigmata 4, 11-15 w longa; basidiola acerosa copiosa. Cheilocystidia etc. ut in P. orientalis. Hyphae
fibulatae, 2-6 uw latae, tunicis plus minusve incrassatis.
Ad ramulos emortuos in silva. Borneo, mt.Kinabalu, c. 3000 m alt., RSNB 895 typus, in herb. Corner.
White, more or less sessile. Pileus -15 mm in radius, 22 mm wide, reniform,
horizontal, opaque, chalky. Stem very short, thick, lateral, or none, becoming
villous at the base. Tubes -1.5 mm long; pores 0.2-0.3 mm wide at first, enlarging to
1.3 mm wide, subcircular with chalky edges, drying dark fuscous brown. Flesh dry.
Not luminous.
Spores 13-18.5 x 12-16 uw, white, smooth, subglobose or broadly ellipsoid,
apiculus 1-1.5 uw long, bluish grey amyloid. Basidia 60-120 x 15-20 w; sterigmata 4,
134 Gard. Bull. Sing. 39(2) (1986)
11-15 uw long, 3-4 uw wide at the base; basidioles subacute, abundant. Cheilocystidia
and hyphal structure as in P. orientalis. Pleurocystidia none. Hyphae 2-6 wu wide,
clamped, with walls 1-2 w thick in the stem and old flesh at the base of the pileus,
not gelatinous; hymenium eventually becoming more or less invaded by 1-3 uw
hyphae branching and lobing round the apices of the basidia.
On fallen twigs in montane oak-forest. Borneo.
Mt. Kinabalu east ridge c. 3000 m. alt., Corner s.n.: 29 & 30 June 1961, 15 July 1961; 20 July 1961,
RSNB 895.
This species of high mountain forest has the largest spores and basidia of the
genus.
Panellus microsporus sp. nov. Figs. 8g, 21
Dorsifixus, pendens, cervinobrunneus. Pileus -4 mm latus, discoideus. Stipes -1.5 x 1 mm, plus
minusve centralis. Tubi -0.6 mm longi; poris 70-100 yw latis, minutis, subangulatis, acie cretaceo-
pruinosa, dissepimentis subgelatinosis. Caro sicca.
Sporae 3-4 x 0.8 uw, allantoideae amyliodeae. Basidiola subacerosa copiosa. Cheilocystidia 25-80 x
7-12 w, clavata vel subcylindrica, ex apicibus saepe diverticulos lobulatos intense amyloideos -4 X 1.5 wu
emittentia; hyphis dichophysialibus nullis. Caulocystidia et pileocystidia ut cheilocystidia, dense instruc-
ta, irregulariter lobulata; hyphis dichophysialibus nullis; saepe massulis intense amyloideis intermixta.
Hyphae fibulatae, 2-6 uw latae, tunicis submucilaginosis praecipue in dissepimentis.
Ad ramulos emortuos in silva. Borneo, mt Kinabalu 1700 m alt., RSNB 5514 typus, in herb. Corner.
Pendent, dorsifixed, fawn brown. Pileus -4 mm wide, discoid. Stem 1-1.5 xX
1 mm, more or less central, expanding obconically into the pileus. Tubes -0.6 mm
long; pores 70-100 yw wide, subangular, chalky; dissepiments subgelatinous. Flesh
firm, not gelatinous.
Spores 3-4 x 0.8 pw, white, smooth, allantoid, pale violaceous amyloid, very
abundant in all fruit-bodies but not seen attached. Basidia 12-15 xX 3.5-4 wp;
The genus Panellus Karst. in Malaysia 135
sterigmata 4; basidioles 1.5-2.5 4 wide, subacerose. Cheilocystidia 25-80 x 7-12 p.
clavate to subcylindric, often shortly lobate at the apex or in the distal part, set
distally with many minute, lobulate, strongly blue-black amyloid processes -4 x
1-1.5 w, granular encrusted, the proximal stalk often with slightly thickened pale
brown walls; without independent lobulate hyphae on the sterile pore-edge.
Pleurocystidia none; gloeocystidia none. Hyphae 2-6 uw wide, clamped, with hyaline
submucilaginous walls 0.5-2 w thick, longitudinal, regularly arranged. Surface of
stem and pileus with a close palisade of erect cystidia similar to the cheilocystidia
but more irregularly lobulate, with pale brown walls, without independent lobulate
hyphae, the outside of the cells densely set in places with dark, purple-brown,
amyloid masses.
On fallen twigs in the forest. Borneo.
Mt. Kinabalu, Bembangan River 1700 m. alt., 27 Feb. 1964, RSNB 5514.
This species appears to be abundantly distinct in the dorsifixed Helotium-like
habit, the large cheilocystidia, the palisade of pileocystidia with amyloid masses.
the subgelatinous hyphae, and the minute allantoid spores. They are the smallest
among Malesian species of the genus and contrast so strongly with the great spores
of the preceding species P. megalosporus that they prove how spore-shape may
have little generic value.
Panellus orientalis (Y. Kobayasi) comb. nov. Figs. 7, 8d, 22
Dictyopanus orientalis Y. Kobayasi, Bull. Nat. Sci. Mus. Tokyo 6 (1963) 360, fig. 3, pl. SOA.
Chalky white, becoming pale buff to pale fawn-ochraceous, finally rufous pink in
age, but the pores white. Pileus-17 mm wide, pleuropodal, reniform, finely pruino-
so-scurfy, becoming minutely cracked, opaque; margin slightly incurved at first.
Stem 1-7 X 2-3 mm, 1-1.5 mm at the base, lateral, dilating upwards into the pileus.
pruinoso-puberulous. Tubes -0.7 mm long; pores 150-250 u wide, circular, elliptic,
or slightly radially elongate, with thin cream-white pruinose edges. Flesh firm soft.
even subgelatinous, drying very horny.
Pan OP.
SD 5
—_— — Py U
ee ae
‘tc bn TE IE
a le
oon
Coit
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136 Gard. Bull. Sing. 39(2) (1986)
Spores 6.5-9.5 x 6-8.5 uw, white, smooth, subglobose to lacrymiform, thin-
walled, pale indigo amyloid. Basidia 20-40 x 7.5-10 uw; sterigmata 4, 6-8 uw long;
basidioles subacute; subhymenium narrow, not corticate. Cheilocystidia 40-190 x
3-6(-7) yw, gloeocystidial with oleaginous contents, thin-walled, cylindric to subcla-
vate, abundant on the pore-edges of young fruit-bodies, often absent (? crushed)
from the older, becoming covered with dichophyses with branches 1.5-3 mw wide,
often with the ends dilated 4-7 uw wide and set with processes -6 X 1-1.5 uw, wholly
immersed in granular incrustation (soluble in dilute potash). Pleurocystidia none.
Hyphae 1.5-6 uw wide, clamped, thin-walled, some with subgelatinous walls, in the
older tissue with the walls thickening slightly 0.5-1 wu, the cells -180 uw long, often
somewhat tuberculate. Surface of stem and young pileus constructed as the pore-
edges with gloeocystidia and echinulate dilated ends of dichophysial branches, the
older pilei becoming covered with a chalky-incrusted layer, -120 uw thick, of excres-
cent lobulate dichophyses with the gloeocystidia scarcely distinguishable.
On fallen branches, dead trunks, bark and bits of wood in the forest, south
Japan; North Borneo, Solomon Islands, at 800-1800 m altitude.
BORNEO. Mt. Kinabalu 1300-1800 m. alt., RSNB: 2705, 2952, 5396, 8078, collected in February,
April, August, September. — SOLOMON ISLANDS 800-1800 m. alt., Guadalcanal, Malaita, RSS:
1652, 1678, 1805, October to November.
The Japanese specimens, which were considerably smaller than the Malesian,
were found not to be luminous by Kobayasi; the tropical have not been tested.
Panellus pauciporus sp. nov.
Pileus -1.5 mm latus, sessilis, albus. Tubi 3-15, minuti, albo-cretacei. Caro sicca.
Sporae 6-7.7 x 3-4 w, subcylindricae amyloideae. Basidiola acerosa nulla. Cheilocystidia et pileocysti-
dia nulla, ad margines tuborum hyphis dichophysialibus copiosis. Hyphae fibulatae, 2-5 pw latae.
Superficies pilei hyphis dichophysialibus instructa.
Ad folia emortua bambusae, in silva. Insulae Solomonenses, Guadalcanal 1800 m alt., RSS 1649
typus, in herb. Corner.
Pileus -1.5 mm wide, sessile, lateral, smooth, white. Pores 3-15 in all, minute,
white, chalky, not lamellate. Flesh firm, not gelatinous.
Spores 6-7.7 X 3-4 w, white, smooth, subcylindric, pale blue amyloid. Basidia
17-20 X 7-8 ww, short, wide, with opalescent contents; sterigmata 4, 3.5 w long;
acerose basidioles none. Cheilocystidia none; with many lobulate and subechinu-
late dichophysial hyphae 1-2 w wide along the sterile pore-edges and developing
between the basidia in the old hymenium. Hyphae clamped, 2-5 pu wide, cylindric
with thin firm walls, more or less radiating, the cells -80 « long. Surface of pileus
with a thin covering of incrusted dichophysial hyphae as on the pore-edges, some
with slightly dilated echinulate-tuberculate ends -5 x wide; no distinct pileocystidia.
On a dead bamboo-leaf in the forest. Solomon Islands.
Guadalcanal, Popomanasiu 1800 m alt., 27 Oct. 1965, RSS 1649.
Panellus pusillus (Pers. ex Lév.) Burdsall et Miller
Nova Hedwigia Beih, 51 (1975) 85, with synonymy.; Corner, Trans. Brit. mycol. Soc. 37 (1954) 258, as
Dictyopanus pusillus; Cunningham, Polyporaceae New Zealand (1965) 140, as D. rhipidium.
Pileus 4-12 mm wide, pleuropodal, convex then flattened, reniform, smooth,
opaque, fawn tan. Stem -3.5 X 0.3-1.5 mm, fawn tan. Tubes -0.5 mm long; pores
100-150 yw wide, circular, white, chalky. Flesh -0.5 mm thick at the base of the
pileus, firm, not gelatinous. ~
Spores 3-4.5 x 2-2.5 uw, white, smooth, ellipsoid, pale blue amyloid. Basidia
13-20 x 4-5(-6) w; sterigmata 4, 3 uw long; basidioles acerose, numerous. Cheilo-
The genus Panellus Karst. in Malaysia 137
cystidia 20-40 x 4-5 w, cylindric to subclavate, thin-walled, not or scarcely gloeo-
cystidial, producing 1-3 lobulating hyphae from the apex, becoming overtopped by
excrescent dichophysial hyphae, in some collections the cheilocystidia indistinct.
Pleurocystidia none. Hyphae 2-7(-11) « wide, monomitic, clamped, with walls -4
thick in the stem. Surface of pileus with gloeocystidial hyphal ends 4-5 ww wide
overtopped by excrescent dichophysial hyphae more or less profusely diverticulate
and densely encrusted, no distinct pileocystidia, the gloeocystidial hyphae lacking
from some collections.
On dead wood and dead fallen branches in the forest.
MALAY PENINSULA. Penang Hill 330 m alt., 21 Sept. 1972, Corner P-81. — BORNEO. Mt.
Kinabalu, east ridge 1000 m. alt., 26 Sept. 1961, RSNB 1515. Sandakan, Sepilok Forest low alt., 13 Aug.
1960, Corner s.n. — NEW GUINEA. Ajiyura 2000 m. alt., 22 Sept. 1960, Corner s.n.
This description is drawn from the Malesian collections. They agree in spore-size
and microscopic structure with those which I collected in Brazil (Corner 1954), but
the fruit-bodies are small and the colour of the living pileus is distinctly brown, not
white to pale tan. This pale colour is given for North American specimens by
Burdsall and Miller, who find slightly larger spores 4-5.5 x 2-3 w. Cunningham, in
recording the species from Australia and New Zealand, gave the pale colour but
the small spores as in the Malesian collections. Whether the species is luminous in
Malesia needs to be tested.
Besides differences in the development of cystidia, noted in the description,
there were others among these Malesian collections. The Bornean had encrusted
subhymenial hyphae, which is rarely the case. The Penang collection had no mat of
excrescent dichophysial hyphae on the pileus, but relatively few short lobulating
hyphae which formed a thin, bluish amyloid, membrane. Thus, spore-size is the
main distinction for the species in the eastern tropics.
Panellus sublamelliformis sp. nov. Fig. 23
Pileus -3 mm latus, subsessilis, subdorsifixus, reniformis rimoso-pruinosulus, albus dein pallide
cervino-brunneus, aetate subincarnatus. Tubi -0.5 mm longi; poris c. 170 uw latis, rotundatis vel leniter
radiato-elongatis, prope marginem pilei sublamelliformibus. Caro sicca.
Sporae 6-7.5 (-8) X 4.5-5.5 yw, late ellipsoideae, amyloideae. Basidiola acerosa copiosa. Cheilocystidia
et pileocystidia 22-35 x 7-18 ww, clavata, hyphis dichophysialibus superantibus.
Hyphae fibulatae, 1.5-5 latae, massulis resinosis brunneis intermixtae.
Ad caules emortuos bambusarum in silva. Insulae Solomonenses, Guadalcanal, 700 m alt., RSS 1534
typus, in herb. Corner.
Pileus -3 mm wide, sublateral, slightly dorsifixed, reniform. minutely rimoso-
pruinose, opaque, white then pale fawn brown, pinkish when old. Stem practically
none. Tubes -0.5 mm long; pores c. 170 u wide, round to slightly radially elongate,
Fig. 23. Panellus sublamelliformis. Fruit-body, X 5; spores, X 2000; basidia and surface of pileus, x
500.
Fig. 24. Panellus glutinosus. Fruit-bodies, x 2; spores in side-view and end-view, * 2000; basidia,
cheilocystidia (central) caulocystidia (right of centre), and surface of pileus with adjacent
hyphae of the flesh. « 500.
towards the margin of the pileus almost shortly lamelliform, white. Flesh 0.2-0.3 mm
thick at the base of the pileus, firm, not gelatinous.
Spores 6-7.5(-8) X 4.5-5.5 uw, white, smooth, broadly ellipsoid, obtuse, varying
subamygdaliform, very pale bluish amyloid. Basidia 24-29 x 6.5-7.5 mw; sterigmata
4, 4-6 w long; basidioles acetose, 3-5.5 sw wide, often mucronate, abundant;
subhymenium narrow, not corticate. Cheilocystidia 22-35 x 7-18 mw, clavate, thin-
walled, smooth, hyaline (not gloeocystidial, even when young), often rather sparse,
138
The genus Panellus Karst. in Malaysia 139
becoming overgrown by dichophysial hyphae 40-60 uw long with the body 3-6 u
wide, with closely lobulate branches, thinly encrusted, not amyloid. Hyphae 1.5-
5 w wide, clamped, with firm walls -2 « thick, compact, with many extra-hyphal
brownish resinous masses swelling in Melzer’s iodine into large vitreous resinous
brown globules. Surface of pileus with cystidia and dichophysial hyphae as on the
pore-edges, the cystidia abundant near the margin of the pileus and almost as a
palisade, becoming spaced over the general surface and much obscured by the
excrescent dichophyses and resinous brown masses.
On dead bamboo stems in the forest. Solomon Islands.
Guadalcanal, Tambalusu c. 700 m alt., 19 Oct. 1965, RSS 1534 and 1536.
Subgen. Mesopanellus subgen. nov.
Receptacula mesopodalia marasmioidea, parva. Lamellae decurrentes, haud vel vix reticulato-
venosae. Basidiola acerosa copiosa. Cheilocystidia clavata, intus oleaginosa. Hyphae haud granuloso-
incrustatae, inflatae. Superficies pilei stipitisque cystidiis clavatis hyphisque dichophysialibus instructa.
Duo species, Borneo, Peninsula Malayana. Typus — P. glutinosus sp. nov.
Fruit-bodies mesopodal, marasmioid, rather small with pileus -2 cm wide, white
then pale ochraceous. Gills decurrent or adnato-decurrent, not dichotomous, not
or slightly reticulately veined.
Spores white, smooth, pale amyloid. Basidioles acerose, abundant. Cheilocysti-
dia strongly clavate with oleaginous contents; dichophysial hyphae on the gill-edge,
sometimes inconspicuous. Pleurocystidia none. Hyphae not granular incrusted,
clamped or not, inflating. Surface of pileus and stem with clavate cystidia and
copious dichophysial hyphae.
Key to the species of subgen. Mesopanellus
1. Pileus and stem glutinous viscid, with stalked clavate cystidia and dichophysial hyphae in the
mucilage. Spores 6-8.5 xX 5.5-7 mw, subtriangular in adaxial view. Clamps absent except on the
mucilage-hyphae of pileus and stem. Hyphae of stem secondarily septate. Borneo
oor fr Ts Pe TE a rae Satie ot ng aE RE Ae ae re ee Oe er eee | ee ee P. glutinosus p. 140
1. Pileus and stem dry. Spores 7-8.5 xX 4.3-5 m, not subtriangular. Clamps present: hyphae not
secondarily septate. Clavate cheilocystidia not distinctly stalked. Malaya ......... P. pyruliferus p. 141
The two species which I have placed in this new subgenus have the spores,
basidioles, cystidia, and dichophysial hyphae of subgen. Panellus but, with meso-
podal fruit-bodies and decurrent gills, they appear to belong to the alliance of
Marasmius. Their hyphae lack the granular incrustation of typical Panellus, but this
is a feature, also, of the three species of subgen. Panellus which lack the acerose
basidioles, and it seems to be typical of the temperate subgen. Mirtellus. Thus, only
the mesopodal stem remains as a satisfactory distinction; along with the thin-walled
inflating hyphae, it suggests that Mesopanellus is a relic of the ancestry of Panellus.
Among the species without acerose basidioles, that which seems nearest to Meso-
panellus is P. fuscatus.
This subgenus introduces two new features to Panellus. The hyphae in the stem
of P. glutinosus are secondarily septate; this is a common feature in Mycena but not
in the alliance of Marasmius. Then in this same species, the pileocystidia appear to
develop laterally from the dichophysial hyphae (Fig. 24). It may be an illusion
caused by the gelatinisation and swelling of the hyphal walls. Thus, the cheilocysti-
dia are terminal with interposed dichophysial hyphae, and the caulocystidia seem to
have this same arrangement.
Fig. 25. Panellus pyruliferus. Fruit-bodies, * 2.5; spores, x 2300; basidia, cheilocystidia (above.
surface of stem (middle drawing), and surface of pileus (below), x c. 500.
Panellus glutinosus sp. nov. Fig. 24
Pileus -18 mm latus, convexus dein planus vel subumbilicatus, viscido-glutinosus striatus, albus dein
pallide subochraceus. Stipes -12 0.5-1.5 mm, centralis, viscidua, albus, ad basim abruptum strigoso-
villosus. Lamellae adnato-decurrentes, crassiusculae subgelatinosae, ad basim subreticulatae, albae,
acie exsiccata brunneo-gummosa. Caro tenax, ad superficiem pilei stipitisque gelatinosa.
Sporae 6-8.5 x 5.5-7 yw, subgloboso-subtriangulares, amyloideae. Basidiola acerosa copiosa.
Cheilocystidia 30-65 x 7-13 w, clavata, intus oleaginoso-guttulata, hyphis dichophysialibus brevibus
intermixta. Caulocystidia et pileocystidia similia, hyphis dichophysialibus gelatinosis superantibus.
Hyphae inflatae, haud incrustatae; in stipite -17 x latae, ordine secundo septatae sine fibulis; in pileo
-26 wu latae, ? fibulatae; in superficie viscida 1.5-4 yw latae, fibulatae.
Ad ramulos emortuos in silva. Borneo, mt. Kinabalu, RSNB 5705A typus, in herb. Corner.
140
The genus Panellus Karst. in Malaysia 141
Pileus -18 mm wide, convex then plane to subumbilicate, glutinous viscid from
the first, striate, white then pale dingy ochraceous over the centre; margin slightly
incurved at first, without veil. Stem 6-12 X 0.5-1.5 mm, cylindric, smeary viscid,
white, at first white pruinose, the abrupt base thinly strigose-villous. Gills adnate to
adnato-decurrent, rather thick, subgelatinous, with mucilaginous edge, 13-20 pri-
maries -2.5 mm wide, 2-3 ranks, becoming slightly reticulate at the base, white, the
edge drying brown-gummy. Flesh thin, rather tough, gelatinous at the surface of
pileus and stem. Smell slightly fishy.
Spores 6-8.5 X 5.5-7 uw, white, smooth, pip-shaped but rounded subtriangular in
adaxial view, with oleaginous contents contracting into a refringent mass in dried
spores, pale bluish amyloid. apiculus 0.7-1 uw long. Basidia 24-32 x 7.5-8.5 w, with
oleaginous contents; sterigmata 4, 5 uw long; basidioles -33 X 4-6 mw, acerose,
abundant; subhymenium narrow, toughly gelatinous, not corticate. Cheilocystidia
30-65 xX 7-13 w, clavate with long narrow stalk 2.5-4 uw wide, smooth, oleaginous-
guttulate, as a sterile edge, arising from the longitudinal subgelatinous hyphae of
the gill-edge; with small inconspicuous dichophyses 1.5-3.5 uw wide. Pleurocystidia
none. Hyphae monomitic, inflated, without clamps except in the mucilage hyphae
on pileus and stem, secondarily septate (at least in the stem) the cells 40-280 x 3-
17 w in the stem and strictly longitudinal with broad septa; in the pileus radiating
with cells -200 3-26 w, narrow and subgelatinous next to the viscid surface; in the
gill-trama as in the pileus but laxly interwoven; turning brownish in Melzer’s
iodine. Caulocystidia as the clavate cheilocystidia, with more copiously branched
mucilaginous dichophyses as the ends of clamped mucilage hyphae 1.5-4 uw wide,
bearing the clavate cystidia laterally and forming a mucilaginous layer 70-150 wu
thick; the base of the stem with many excrescent 2-4 uw hyphae. Surface of pileus
with a mucilaginous layer -400 yw thick, constructed as on the stem but with the
mucilage hyphae more or less perpendicular to the surface and the dichophyses less
compactly branched. Young pileus with the mucilage layer initiated by excrescent
hyphae 1.5-3 u wide, perpendicular to the surface, developing dichophyses at the
ends and the clavate cystidia laterally; not initiated as a compact palisade.
On sticks in the forest, gregarious. Borneo.
Mt. Kinabalu. Mesilau 1600 m alt.. 10 March 1964. RSNB 5705A:; /6 March 1964, RSNB 5705B; }
May 1964. RSNB 5705C.
Panellus pyruliferus sp. nov. Fig. 25
Ex integro albus, aetate flavidulus. Pileus -10 mm latus, convexus dein concavus, laevis, substriatus,
siccus. Stipes -12 x 0.5-1 mm, centralis, pruinosulus, siccus, ad basim subvillosus. Lamellae adnatae vel
subdecurrentes, subdistantes, angustae, haud reticulatovenosae. Caro firma, sicca.
Sporae 7-8.5 x 4.3-5 uw. ellipsoideae, haud subtriangulares, amyloideae. Basidiola acerosa copiosa.
Cheilocystidia 29-35 x 7-8.5 mw. pyriformia, tenuiter tunicata, intus flavidulo-opalescentia, hyphis
dichophysialibus superantibus. Caulocystidia et pileocystidia aut clavata ut cheilocystidia aut subcylin-
drica, subcapitata, -45 x 5-7 uw. saepe lobulis paucis praedita. Hyphae fibulatae, inflatae, nec incrustatae
nec ordine secondo septatae nec gelatinosae: in stipite -17 wu latae; in pileo lamellisque 2-5 uw latae sed
prope superficiem pilei 8-20 wu latae.
Ad lignum emortuum in silva. Peninsula Malayana, Pahang, Cameron Highland, Corner s.n. 2 Oct.
1966 typus, in herb. Corner.
Entirely white, yellowish in age. Pileus -10 mm wide, convex to concave, smooth,
substriate; not viscid; margin slightly incurved at first. Stem 5-12 x 0.5-1 mm,
cylindric, finely pruinose, dry, base slightly thickened and subvillous. Gills adnate
to subdecurrent, subdistant, narrow, 15-20 primaries 0.3-0.7 mm wide, 1-2 ranks,
interstices smooth. Flesh firm, drying hard, inodorous.
Spores 7-8.5 x 4.3-5 uw. white, smooth, ellipsoid, obtuse, not subtriangular, pale
bluish violaceous amyloid. Basidia 29-35 xX 7-8.5 yw; sterigmata 4, 5 mw long:
_—
ee”
\
Wwe?
Fig. 26. Panellus magnus. Fruit-bodies, x 2: unexpanded fruit-body. x 3: spores. x 2500; basidia
and cheilocystidia, « 750.
142
Fig. 27. Panellus magnus. Caulocystidia, x 500.
The genus Panellus Karst. in Malaysia 143
basidioles acerose to subacuminate, abundant. Cheilocystidia -27 x 8-15 p, pyri-
form, thin-walled, smooth, with yellowish opalescent contents, as broad sterile
edge to the gill, becoming overgrown by narrow, 1-2.5 « wide, hyphae with spicate
branches. Pleurocystidia none. Caulocystidia of two kinds; clavate as the cheilocy-
stidia, more or less decumbent and scattered; as abundant small subcylindric,
subcapitate, or subventricose cells -45 x 2.5-5(-7) uw, simple or with one or two
processes. Pileocystidia as the caulocystidia, of two kinds, but sparser. Hyphae
monomitic, clamped, inflating; cells of the stem 60-300 x 6-17 yw, not or scarcely
constricted at the broad septa, longitudinal, the walls often finely undulate and
uneven, with a few narrow uninflated hyphae; in the pileus mostly uninflated, 2-5 uw
wide, interwoven, with firm walls, but near the upper surface with inflated cells
8-20 « wide; in the gill-trama uninflated, with firm walls. Oleiferous hyphae 3-6 u
wide scattered in all parts.
On dead wood in the forest, gregarious. Malay Peninsula.
Pahang, Cameron Highland 1700 m. alt., 2 Oct. 1966, Corner s.n.
Macroscopically this suggests Omphalina or Trogia but the tissue is rather firm
and dry. Microscopically it is easily recognised from the small pyriform cystidia
with opalescent contents on the gill-edge and surfaces of pileus and stem. It could
be looked for in Marasmiellus but the amyloid spores do not fit.
Subgen. Megalopanellus subgen. nov.
Pileus -9 cm latus, plus minus mesopodalis, furfuraceo-squamulosus. Stipes bene evolutus. Caro
tenax, haud gelatinosa, inamyloidea. Basidiola acerosa nulla. Cheilocystidia vix diverticulata. Caulocy-
stidia diverticulis ramoso-lobulatis saepe praedita. Superficies pilei trichodermoidea, sine hyphis
dichophysialibus. Hyphae fibulatae, inflatae, superficiales tunicis brunneis et brunneo-incrustatis saepe
praedita.
Lignicola, Borneo: typus Panellus magnus sp. nov.
This fungus could be squeezed into several genera with smooth amyloid spores,
such as Clitocybula, Porpoloma and Baeospora, but it has the essential construction
of Panellus where it assumes a primitive position in stature, hyphae, and squamu-
lose pileus and stem. None of these points seems to have generic distinction when
one considers the latitude in other genera such as Mycena, Pluteus and Amanita. |
prefer the large generic concept because so many microgenera are merely transient
abstractions from the incompletely known flora of the world. I find in subgen.
Megalopanellus what one would expect for the derivation of the species with small,
lamellate or poroid, fruit-bodies which are the majority. Thus there is a vestige of a
trichoderm on the pileus of the lamellate P. fuscatus.
Panellus magnus sp. nov. Figs. 26-29
Pileus 2-9 cm latus. convexo-planus, saepe subumbonatus, siccus, opacus vel prope marginem anguste
striatus, fuscus. marginem versus isabellinus, minute fuscofuligineo-furfuraceus vel subsquamulosus.
Stipes 3-6 cm X 3-6 mm, cylindricus vel apicem versus subattenuatus, saepe excentricus, haud lateralis,
pallide isabellinus dein e basi abrupto fuscofuligineus, minute fuscofurfuraceus vel squamulosus, apice
albopruinoso. sine annulo vel velo. Lamellae sinuato-adnatae vel adnato-decurrentes, confertae,
tenues, tenaces. primariae 40-60. 2-4 mm latae, ordinibus 3-4 instructae, acie integra, albae dein
isabellinae. aetate fuscofuligineae. Caro in centro pilei -2 mm crassa, plus minusve tenax, subcoriacea,
haud gelatinosa, inodora.
Sporae 3.7-6 X 1.8-2.8 ws. albae ellipsoideae. plus minusve intense amyloideae, Cheilocystidia 28-50 x
5-17 w. subcylindrica, clavata, vel subventricosa, tenuiter tunicata, ex apicibus diverticulis paucis saepe
praedita, lamellae acie fertili dispersa. Pleurocystidia nulla vel sparsa ut cheilocystidia. Caulocystidia
30-150 x 4-17 wu. subventricosa. fusiformia, tunicis brunneis, apices versus hyalina et saepe in diverticu-
los ramosos vel lobulatos producta. Hyphae monomiticae, fibulatae, inflatae -22 w latae, hyphis angustis
Fig. 28. Panellus magnus. Surface of centre of pileus, x 500.
1.5-7 w latis intermixtis, saepe diverticulis brevibus paucis emittentes, tunicis -0.5 incrassatis, sparsim
ordine secundo septatae; oleiferae nullae. Superficies pilei trichodermoidea disrupta, hypharum apici-
bus 0-2 septatis, cellulis apicalibus 45-90 x 4-20 «, plerumque clavatis, tunicis brunneis; ex hyphis 4-9
latis, tunicis brunneis et saepe brunneo-incrustatis, oriens.
Ad truncos ramosque emortuos in silva montana, frequens, caespitosus, Borneo, mt Kinabalu, RSNB
847 typus, in herb. Corner.
Pileus 2-9 cm wide, conico-convex to convexo-plane, often subumbonate, dry,
opaque, or narrowly striate at the margin, fuscous, bistre yellowish towards the
margin, becoming paler on expansion, minutely fuscous fuliginous scurfy or
squamulose, sparsely towards the pallid smooth margin slightly incurved at first.
Stem 3-6 cm X 3-6 mm, cylindric or slightly attenuate upwards, often excentric but
not lateral, pale livid white then fuscous fuliginous from the base upwards, minutely
fuscous squamulose or scurfy, innately darker fibrillose from the base to the white
pruinose apex; without veil and ring. Gills sinuato-adnate to adnato-decurrent,
crowded, thin, tough, 40-60 primaries 2-4 mm wide, 3-4 ranks, not dichotomous,
edge entire, white then tinged bistre, fuscous fuliginous in age. Flesh -2 mm thick in
the centre of the pileus, rather tough, especially in the stem, white then concol-
orous. Inodorous; poisonous according to local inhabitants.
Spores 3.7-6 X 1.8-2.8 uw, white, smooth, ellipsoid, blue-black amyloid to pale
vinaceous amyloid (RSNB 1684, 8006). Basidia 13-22 x 3.5-4 w, 4-spored; acerose
basidioles absent; subhymenium 10-20 yw thick, composed of 2-3 w wide interwoven
hyphae. Cystidia 28-50 x 5-17 w, clavate or subventricose, with or without short
processes, smooth, thin-walled, projecting -20 s« or almost wholly immersed,
scattered on the gill-surface and along the fertile gill-edge. Caulocystidia 30-150 x
4-17 uw, more or less fusiform or subventricose with firm brown walls and colourless
thin-walled apices, often with narrow lobed or branched processes from the apex,
less often the processes lateral, forming a disrupted palisade, eventually invested
144
The genus Panellus Karst. in Malaysia 145
with dichophysial hyphae 1.5-2 u wide. Hyphae monomitic, clamped, inflating, the
walls becoming firm and in the stem thickened -0.5 w, with cells 35-500 X 8-22 uw in
the longitudinal hyphae, narrowed to the septa or not, often somewhat uneven in
width, occasionally secondarily septate in the stem, often with one or a few short
abortive processes but no true binding hyphae, with narrower hyphae 1.5-7 u wide
interweaving and often rather copiously branched with abortive processes but
septate with clamps; no oleiferous hyphae; in the gill-trama, parallel descending,
more or less inflated as in the pileus, forming a rather tough tissue, not amyloid.
Surface of pileus with a short disrupted pile of hyphal ends with firm brown walls,
Fig. 29. Panellus magnus. Hyphae of the stem, that on the right secondary septate with cytoplasm
retracted from the apex: x 500.
146 Gard. Bull. Sing. 39(2) (1986)
1-3 cells long, the terminal cells 45-90 x 4-20 w, generally clavate and subacute,
arising from 4-9 uw wide hyphae with firm brown walls and often with more or less
annular brown incrustation; the hyphal ends of the pile more or less erect over the
centre of the pileus, more or less decumbent and scattered in clusters over the limb.
Frequent, caespitose, on dead standing trunks and fallen wood in montane
forest, Borneo.
Mt. Kinabalu 1700-3500 m alt., 16 July 1961, RSNB 847; 9 Aug. 1961, RSNB 1684; 20 Feb. 1964,
RSNB 5381; 31 March 1964. RSNB 8006.
The tufts of this pallid fungus are a conspicuous sight on the hard dead trunks
which abound in the upper forest on mt. Kinabalu; there, oaks, Myrtaceae,
Rhododendron and podocarps abound with many other kinds of tree, and I was
unable to decide what might have been the host-tree. The Dusun people claimed to
know it well for they regarded it as poisonous, but I am not convinced that they
distinguished it from Pleurotus decipiens (Corner 1981). I did not test whether it
was phosporescent. Superficially, this fungus would pass for Lentinus in its general
sense. The hyphal structure is incipiently dimitic, though not sarcodimitic as in
Trogia, and the interweaving processes suggest not so much binding hyphae as
incipient internal dichophysial hyphae.
References
Burdsall Jr, H.H. and O.K. Miller Jr (1975). A re-evaluation of Panellus and
Dictyopanus (Agaricales). Nova Hedwigia, Beih. 51, 79-91. J. Cramer, Vaduz.
Corner, E.J.H. (1950). Descriptions of two luminous tropical agarics (Dictyopanus
and Mycena). Mycologia 42: 423-431.
. (1954). Further descriptions of luminous agarics. Trans. Brit. mycol. Soc.
3) 256-27 fF,
. (1981). The agaric genera Lentinus, Panus and Pleurotus. Nova Hedwigia,
Beih. 69. J. Cramer, Vaduz.
______. (1983). Ad Polyporaceas I. Nova Hedwigia, Beih. 75. J. Cramer, Vaduz.
. (1984). Ad Polyporaceas II and II. Nova Hedwigia, Beih. 78. J. Cramer,
Vaduz.
Dennis, R.W.G. (1952). The Laschia-complex. Kew Bull. 1952: 325.
. (1970). Fungus flora of Venezuela and adjacent countries. Kew Bull.,
Addit. Ser. III.
Jiilich, W. (1981). Higher taxa of Basidiomycetes. Bibliotheca Mycologica 85. J.
Cramer, Vaduz.
Kobayasi, Y. (1963). Revision of the genus Dictyopanus with special reference to
the Japanese species. Bull. Nat. Sci. Mus. Tokyo 6: 356-364.
Macrae, R. (1937). Interfertility phenomena of the American and European forms
of Panus stypticus. Nature 1937: 674.
Singer, R. (1945). The Laschia-complex (Basidiomycetes). Lloydia 8: 170-230.
___. (1969). Mycoflora Australis. Nova Hedwigia, Beih. 29. J. Cramer, Vaduz.
______. (1975). The Agaricales in modern taxonomy. J. Cramer, Vaduz.
The genus Panellus Karst. in Malaysia
Dictyopanus
copelandii
gloeocystidiatus
luminescens
orientalis
pusillus
rhipidium
Panellus
subgen. Megalopanellus subgen. nov.
Mesopanellus subgen. nov.
Panellus
albifavolus sp. nov.
alutaceous sp. nov.
ambiguus sp. nov.
bambusarum sp. nov.
bambusifavolus sp. nov.
brunneifavolus sp. nov.
brunneomaculatus sp. nov.
copelandii
dichotomus sp. nov.
var. pinnatus var. nov.
?diversipes
Index to Taxa
147
(Panellus)
eXIQUUS Sp. nov.
fuscatus sp. nov.
gloeocystidiatus comb. nov.
glutinosus sp. nov.
hispidifavolus sp. nov.
intermedius sp. nov.
var. Stenoporus var. nov.
longinquus
luminescens comb. nov.
magnus sp. noy.
megalosporus sp. nov.
MicrOsporus sp. nov.
orientalis comb. nov.
parvulus sp. nov.
pauciporus sp. nov.
pendens sp. nov.
pusillus
pyruliferus sp. nov.
sublamelliformis sp. nov.
sublevatus sp. nov.
127
A New Species of Platycerium from Peninsular Malaysia
Aziz BIDIN & Razali JAMAN
Botany Department, Faculty of Life Sciences
Universiti Kebangsaan Malaysia Bangi, Selangor Malaysia
EFFECTIVE PUBLICATION DATE: 24 JAN. 1987
Abstract
A new species of Platycerium from Peninsular Malaysia, P. platylobum Aziz Bidin & Razali Jaman is
described.
Platycerium platylobum Aziz Bidin & Razali Jaman sp. nov. Plates 1 & 2
Platycerio coronario (Koénig ex Muller) Desv. affinis, ab ea differt: paleis rhizomatis usque 20 x 5
mm tantum, apicem versus angustatis, marginibus costae usque apicem manifeste prominentibus; lobis
ultimis ramorum duorum inferiorum frondium fertilium quam lobis ceteris frondium majoribus, usque
32 x 4m; lobis fertilibus stipitatis (stipite alato 3-5 cm longo), usque 13 cm latis, planis, tenuioribus,
profunde aequaliter bilobatis, lobis ambo etiam leviter bilobatis; paraphysibus brevioribus, 3 vel 4
cellulis sonstitutis, radios 11-13 (-15) ferentibus; cellulis induratis annuli sporangiorum 5-6 (-8).
Agreeing in growth-habit and branching of fronds with Platycerium coronarium
(Koénig ex Muller) Desv., differing as follows: rhizomes-scales to 20 mm long and
5 mm wide only, narrowed towards their apices, the prominent margins of the
thickened median band (costa) distinct to the apex of each scale; ultimate lobes of
the basal two branches of fertile fronds to 32 X 4.cm, much longer and wider than
the ultimate lobes of other branches; fertile lobe stalked (stalk winged, 3-5 cm
long), to 13 cm wide, flat, thinner than in P. coronarium, deeply bilobed; paraph-
yses much shorter, consisting of 3 or 4 cells, their rays 11-13 (-15) in number;
undurated cells of the annulus of sporangia 5-6 (-8).
Aziz Bidin & Razali Jaman PL 149: Langkawi Island, road to Padang Lunas, c. 30 m alt., 12 Feb. 86.
Epiphyte, uncommon (Holotype: UKMB; Isotype: K).
Distribution. Langkawi Island. The ferns were absent from the other islands in
the group as well as the interior of Perlis and northern Kedah, all of which share the
same monsoonal climate.
Ecology. The species was first found on an old rubber tree, about 4 m high from
the base of the trunk, in a village near Padang Lunas about 5 km from Kuah Town.
No other species of Platycerium was observed in the vicinity. Later collections were
from rubber trees in a plantation in Kisap, c. 50 m elevation, about 12 km from
Kuah town. There, all the three species were found. P. coronarium was found on
the higher branches of tall trees while P. holttumii and P. platylobum were on low
branches, 2-5 m above the ground.
This new species differs from other Platycerium species which also bear sori on
individual lobes in that each individual lobe here has a deep, median incision, the
distal, part bi-lobed on each side of the incision, with the result that the whole
resemble a pair of butterfly wings as opposed to the shape in P. coronarium which
is kidney or semi-circular and the one in P. ridleyi, obovate or elliptical (respective-
ly Holttum, 1968 and Hennipman & Ros, 1982). In this species too, the entire
fertile lobe is flat as opposed to concave in both the other two; the paraphyses are
149
150
pape gos \
Plates:
Plate 2.
Platycerium platylobum, fertile frond.
Platycerium platylobum, fertile lobe.
A new Species of Platycerium from Peninsular Malaysia 151
short-stalked (3 to 4 cells long) as against longer stalks in P. coronarium (7 to 8 cells
long) and P. ridleyi (8 to 10 cells). The species is only known from Langkawi Island
whereas P. coronarium is common throughout the lowlands of Peninsular Malaysia
and P. ridleyi is limited to the southern part of the Peninsula, i.e., Johore and
Singapore (Holttum, 1968).
Acknowledgements
The senior author is grateful to Prof. R.E. Holttum for guidance and encourage-
ment and for the Latin diagnosis, to the Keeper and Curator of the Herbarium,
Royal Botanic Gardens Kew for the use of materials and to the staff of the Office of
Forest, Langkawi for allowing the authors to carry out a survey of the pter-
idophytes of Langkawi Islands. The field work was supported by Universiti
Kebangsaan Malaysia Research Grant No. 123/85. Our thanks must also be re-
corded to Mr. A. Zainuddin for his competent assistance in the field and to the
World Wildlife Fund Malaysia and the British Council for the travelling grant to
enable the senior author to travel to Kew.
Literature Cited
Hennipman, E. & M.C. Roos (1982). A Monograph of the Fern Genus Platycerium
(Polypodiaceae). North-Holland Publishing Company, Amsterdam, Oxford,
New York.
Holttum, R.E. (1968). Flora of Malaya, 2, Ferns of Malaya, Government Printer,
Singapore.
STUDIES IN THE FERN-GENERA ALLIED TO
TECTARIA CAV. VI
A conspectus of genera in the Old World regarded as related
to Tectaria, with descriptions of two genera
RE -HOLEFUM
Royal Botanic Gardens, Kew, Surrey, England
EFFECTIVE PUBLICATION DATE: 24 JAN. 1987
Abstract
A brief conspectus of the palaeotropic genera, regarded by the author as related to Tectaria Cav., is
presented with: comments on individual genera, descriptions of two new ones, Chlamydogramme and
Megalastrum, some thoughts on inter-relationships, a key to the genera, and also brief comments on
neotropic genera.
Introduction
In the first paper of this series (Holttum 1984, p. 314) is the following statement
which was intended to show the differences between two groups of genera which
together constitute almost the whole of the family Aspidiaceae as arranged by Pichi
Sermolli (1977):
Dryopteris group. Midribs of ultimate leaflets grooved, the groove of the rachis
bearing the leaflets being open to admit the leaflet-groove, the margin of the leaflet
being decurrent (but not prominent) down the side of the rachis; ctenitoid hairs
lacking.
Tectaria group. Midribs of ultimate leaflets more or less prominent (in Tectaria
sometimes slightly grooved) and bearing ctenitoid hairs, usually many.
While working on a monograph of the genus Crenitis in Asia, Malesia and the
Western Pacific I discovered the existence within Crenitis as arranged by Ching
(1938) of a group of species (his suggested subgenus Dryopsis) which has pinna- or
pinnule-midribs grooved but with the groove closed near the junction of the pinna
with the rachis which bears it. In these species the hairless groove has on its borders
thick-based hairs (the cells near the base wider than long) like those in a similar
position in Peranema and Nothoperanema, also some structures which are narrowly
scale-like above a similar thick base; these are quite unlike anything in true Crenitis
and the species differ from typical Crenitis also in other ways. Mr P.J. Edwards
assisted me to make a detailed comparative study of all such species, which we have
assigned to a new genus Dryopsis in paper II of the present series. The detailed
studies show that the distal part of some of the peculiar hairs resembles a single hair
of Ctenitis. Thus Dryopsis may be a connecting link between the Tectaria and
Dryopteris groups of genera. Furthermore, in some genera of the Jectaria group
the multicellular hairs on the raised upper surfaces of costae are more or less
acicular, not contorted on drying as in Cfenitis. My distinction between the two
groups of genera will therefore need to be modified. I still believe however that the
genera I associated together in 1947 as a subfamily Tectarioideae are a natural
group.
153
154 Gard. Bull. Sing. 39(2) (1986)
I have now examined the types of almost all species in the Old World (including
Africa) assigned to genera of the Tectaria group and here summarize significant
information about them in the form of a tentative conspectus, with descriptions of
two new genera and some thoughts on their possible inter-relationships, followed
by an artificial key. I intend to prepare an account of Malesian species based on this
scheme for Flora Malesiana, where I hope to publish the facts about them in more
detail, and to make formal transfer of some names.
The genera closely associated with Tectaria have (where known) 40 chromo-
somes and lack cylindric glands. The remaining genera have (where known) 41
chromosomes and are divided into two groups, those with and without glands. The
genera are arranged below in these categories; possible inter-relationships between
members of the categories are then considered.
Tectaria and closely allied genera
Common characters: scales narrow, consisting wholly of narrow cells, never
clathrate, rarely abundant on smaller axes of the frond; ctenitoid hairs always
present on the upper surface of the rachis; venation and branching (apart from
basal pinnae) almost always catadromous; veins all free or variously anastomosing;
unicellular cylindric glands lacking; sori in most species indusiate but in some
exindusiate and + spreading along veins, in a few, where the fertile lamina is much
contracted, spreading along all veins and covering the whole of the lower surface;
chromosome number, where known, 40.
Tectaria Cav.
Anales Hist. Nat. 1: 115 (1799)
This is much the largest genus, consisting of about 210 species of which 40 are
neotropic. The best subdivision (not quite a sharp one) appears to be between
species (including the type) which have amply anastomosing veins with free bran-
ched veinlets in the areoles, and those which have either free veins or veins forming
narrow costal areoles lacking included free veinlets and few included veinlets in
other areoles (venation of Sagenia Presl). A separation of species which are wholly
free-veined is not possible; in 7. fuscipes (Bedd.) C. Chr. sterile fronds have veins
forming costal areoles but fertile ones have all veins free. I thus propose to treat
Tectaria as consisting of two sections.
Sect. Tectaria
Including Hemigramma (M.G. Price 1974) and Quercifilix (W.A. Sledge 1972).
The type species of Hemigramma, H. latifolia (Goldm.) Copel. (= Tectaria
hilocarpa (Fée) Price), has simple fronds, the fertile ones much constricted with a
simplified venation and sporangia borne all along the closely crowded veins (pl. 1).
Copeland (1908) showed that hybrids were formed in Luzon between this and
Tectaria crenata Cav. In 1928 he enlarged Hemigramma by including in it Gymnop-
teris decurrens Hook. from Hong Kong and allied species. But the latter have
broader fertile pinnae than the simple fertile fronds of H. latifolia, with an ample
venation exactly as in the sterile pinnae, and a species in the Malay Peninsula has
sori intermediate in form between those of H. decurrens and and species of Tectaria
which have many small sori. Thus H. latifolia and H. decurrens are related to
different species of Tectaria. In both, the sporangia are borne all along the veins on
a restricted surface. The same is true of the only species of Quercifilix, which
hybridizes with Tectaria decurrens in Ceylon. The species originally named Lep-
tochilus pentagonalis R. Bonap., transferred with doubt to Hemigramma by Christ-
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156
Fern-genera allied to Tectaria Cav. VI 157
ensen, appears to be nearer to Quercifilix than to any species of Hemigramma as
arranged by Copeland.
Sect. Sagenia (Presl) Holttum, stat. nov.
Sagenia Presl, Tent. Pterid. 86 (1836), as regards venation. Type species (selected here): Sagenia
latifolia Presl, |.c. fig. 23 = Tectaria mexicana (Fée) Morton, Amer. Fern Journ. 56: 133 (1966), not T.
latifolia (Forst.) Copel.
Stenosemia Presl, which has stertle fronds with Sagenia venation and much-
contracted fertile ones, also Cionidium Moore which has extra-marginal sori ter-
minal on veins, are included here.
Species of sect. Sagenia which have all veins free are a minority. They are most
abundant in mainland S.E. Asia; apart from the Philippines, very few occur in
Malesia and the Pacific. I regard these free-veined species as representing the
original form of the genus and S.E. Asia as its probable centre of origin. The largest
fronds are borne by T. ingens (Atk.) Holttum of Sikkim, attaining a length of 3 m.
Two free-veined species in the neotropics have been included in Tectaria: T.
brauniana (Karst.) C. Chr. and T. pedata (Desv.) R. & A. Tryon. They are
certainly not related to the free-veined species of Asia; see below for further
comment.
Genera related to Tectaria sect. Tectaria
Tectaridium Copel.
Philip. J. Sci. 30: 329 (1926).
Sterile fronds simple and unlobed, with venation as Tectaria sect. Tectaria; fertile
fronds lobed to the costa, the very narrow lobes bearing indusiate sori; intermedi-
ate forms are frequent. There is only one species, which does not appear to be
nearly related to any species of Tectaria.
Chlamydogramme Holttum, gen. nov.
Generi Tectariae Cav. affinis; frondes pinnatae, valde dimorphae; pinnae frondis sterilis simplices,
integrae, venis ut in Tectaria sect. Tectaria ordinatis; pinnae frondis fertilis anguste lineares, soris
linearibus marginalibus laminae et indusiis prope costas in juventute tectis. Type species: Chlamydog-
ramme hollrungi (Kuhn) Holttum, comb. nov.
(Gymnopteris hollrungii Kuha in Schum. & Hollrung. Fl. Kaiser Wilhelmsl.: 8 (1889); Hemigramma
hollrungii (Kuhn) Copel.)
Chlamydogramme hollrungi (Kuhn) Holttum comb. nov.
This species has fronds very similar in general aspect to those of the species from
Lombok originally described as Leptochilus siifolius by Rosenstock and also trans-
ferred to Hemigramma by Copeland, but with a continuous indusium (not noticed
by Copeland) along each side of the pinna-midrib in fertile pinnae; sections of
fertile pinnae of the two species are shown in plate 2.
Chlamydogramme is comparable to the tropical American genus Dictyoxiphium
Hook. but the latter has simple fronds, the fertile ones not greatly constricted so
that the continuous indusium is not near the midrib. The two genera are not nearly
related.
Genera related to Tectaria sect. Sagenia
Heterogonium Presl, emend. Holttum
Kalikasan 4: 205-231 (1975).
Basal pinnae always narrowed towards their bases on the basiscopic side; vena-
158 Gard. Bull. Sing. 39(2) (1986)
tion as in Tectaria sect. Sagenia even in fronds with broad pinnae, only differing in
less frequent free veinlets in non-costal areoles; sori often exindusiate and variously
+ elongate along the veins, in a few species running along all veins in a constricted
fertile lamina; about 20 species, mainly Malesian.
Though the shape of the basal pinnae is the only constant character disting-
uishing species of this genus from those of Tectaria sect. Sagenia, they collectively
have a variety of other characters and appear to form a natural group which is more
sharply distinct from Tectaria sect. Tectaria than 1s sect. Sagenia. The only evidence
I have seen of hybridization with species of Tectaria sect. Sagenia is provided by
some Philippine specimens, collected by M. G. Price, which appear to be hybrids
between H. pinnatum (Copel.) Holttum (a tetraploid with constricted fertile
fronds) and Tectaria aurita (Sw.) S. Chandra (Stenosemia aurita Presl), which T.G.
Walker (1973) found also to be a tetraploid with n = 80.
R.C. Ching (1938) has proposed a genus named Ctenitopsis which includes
free-veined species of both Heterogonium and Tectaria; in my view it is not a
natural group. Copeland has included such species in Ctenitis.
Psomiocarpa Presl.
Epim. Bot.: 161 (185).
The single species has sterile fronds which are small but bipinnate, with the same
frond-form as some Philippine species of Tectaria sect. Sagenia, but the fertile
fronds have extremely small pinnules quite covered with sporangia. M.G. Price has
found plants intermediate between this and a bipinnate species of Tectaria. Pichi
Sermolli (1977: 465) has wrongly placed this genus near Crenitis.
Aenigmopteris Holttum
Blumea 30: 1-11 (1984).
The five species have elongate fronds with many pinnae which are more finely
dissected than those of any species of Tectaria sect. Sagenia; pinna-lobes, which are
themselves deeply lobed, are connected by narrow wings along the pinna-midrib
and thus resemble Lastreopsis but the margin of the wing is not thickened and the
rachis is not winged; as in Tectaria sect. Sagenia there are thick hairs between veins
on the upper surface.
Genera having unicellular cylindric glands
The following four genera agree in having unicellular cylindric glands on the
stalks of sporangia, in many species also on the margins of indusia and/or appressed
to veins; they agree also in having the chromosome number 41, but they differ from
each other in many ways.
Ctenitis C. Chr.
Verdoorn, Manual Pterid.: 544 (1938).
This was originally named Dryopteris subg. Ctenitis in Christensen’s Monograph
of the genus Dryopteris (part 1: 82-112, 1913; part 2: 31-93, 1920). He there dealt
only with American species and defined the subgenus very broadly. In my recent
study of all palaeotropic species I have felt obliged to limit a genus named Ctenitis
by excluding Christensen’s groups of Dryopteris subincisa (Megalastrum of the
present conspectus) and of Dryopteris protensa (Triplophyllum Holttum 1986). I
have not attempted to assess critically all the other neotropic groups recognized by
Fern-genera allied to Tectaria Cav. VI 159
Christensen but believe that they would accord to my definition of Ctenitis. As thus
limited, the genus comprises about 100 species and is considerably diversified in
both the Old World and the New. In the Old World there are three distinct groups
of species: the group of C. submarginalis (Langsd. & Fisch.) Ching (type species of
the genus) which is mainly American but extends to Africa and the Mascarene
Islands, and the species of mainland Asia, Malesia and the Western Pacific which
are divisible into two groups which are sharply distinct from each other in both
scales and spores (Holttum 1985). Two Hawaiian species are very different in
spores from any in the western Pacific and I think also from any in the Americas.
Thus the genus Crenitis, worldwide, needs a new conspectus and I do not propose
formal subdivisions for the Malesian species.
Ctenitis differs from Tectaria in its glands, its fragile gland-fringed indusia, in
having abundant scales which are at least in part clathrate on all the smaller axes of
the frond, also in having fronds which in almost all species are more finely divided,
the ultimate leaflets almost always deeply lobed and always with free veins.
Lastreopsis Ching, emend. Tindale
Contr. N.S.W. Nat. Herb. 3: 249-339 (1965).
In Tindale’s monograph of the genus 33 species are recognized; since then a few
more have been added. Their fronds are finely divided somewhat as in Ctenitis but
differ in scales, which are more like those of Tectaria, and in the thickened
decurrent basiscopic margins of leaflets which form wings on the axes to which they
are attached. The hairs in this genus are notably varied; ctenitoid hairs occur in
most species, but in some the hairs on the upper surface of axes of the frond are
rigid and more or less acicular, not contorted on drying; such hairs are characteris-
tic of Megalastrum, described below. Lastreopsis is pantropic in distribution but the
monograph by Tindale does not indicate any subdivision into natural groups of
species. I suggest that the nature of the hairs on the upper surface of axes of the
frond might offer clues to the distinction of sections within the genus. My impress-
ion is that the species of West Africa need more study.
Pleocnemia Presl. emend. Holttum
Kew Bull. 29: 341-357 (1974).
This genus of 19 species, mainly Malesian, was based by Pres] on the venation
(which agrees with Tectaria sect. Sagenia) and on the exindusiate sori of the sole
original species, but there are also indusiate species and Pres] did not notice the two
most distinctive characters, namely the presence of teeth in the sinuses between
pinnule-lobes and of cylindric glands which are usually yellow or orange in contrast
to the usually pallid glands of most species of Crenitis. The fronds of mature plants
of all species are large, and the pattern of vascular strands in the stipe is more
complex than in the other genera.
Coveniella Tindale
Gard. Bull. Sing. 39:169 (1986).
Caudex slender, long-creeping, with closely approximate fronds; scales very
short, consisting of short cells which are not clathrate, those on the rachis and
costae grading to ctenitoid hairs; short slender pluricellular hairs abundant near
bases of stipes, variably abundant on both surfaces of the rachis, unicellular glands
sparsely present on various parts of the fronds; fronds simply pinnate, pinnae*entire
and narrowed near their bases, the lower ones stalked, not articulate to the rachis;
veins in pinnate groups, the basal veins in each group free, successive ones anasto-
160 Gard. Bull. Sing. 39(2) (1986)
mosing to produce short free excurrent veinlets; sori exindusiate, short cylindric
glands present on sporangium-stalks; n = 41 (S.K. Roy).
Only one species is known, in Queensland and north-eastern New South Wales;
it was originally named Polypodium peocilophlebium Hook.
Genera lacking glands
The following five genera agree with the gland-bearing ones in having 41
chromosomes but in other ways are not closely related. They show varied indica-
tions of relationship to either Tectaria or Ctenitis except Cyclopeltis which is
probably related to Coveniella. A sixth genus, Pseudotectaria, has not yet been
examined cytologically.
Ataxipteris Holttum
Blumea 30: 10 (1984).
The sole species of this genus, from southern China and Japan, has fronds
agreeing exactly in form and venation with those of some free-veined species of
Tectaria but has abundant rachis-scales much like those of Ctenitis; its spores are
similar to those of one palaeotropic group in Crenitis.
Pteridrys C. Chr. & Ching
Bull. Fan Mem. Inst. Biol. Bot. 5: 129 (1934).
Scales narrow as in Tectaria but with strongly cordate base; fronds simply
pinnate; pinnae deeply lobed with a tooth in each sinus between lobes, the tooth
projecting out of the plane of the pinna; veins free, arranged as in free-veined
Tectaria; ctenitoid hairs few, near the bases of pinnae, sometimes on both surfaces
of costae.
This genus of seven species is distributed in mainland S.E. Asia, Ceylon and
Malesia. The sinus-teeth are like those of Pleocnemia but in other ways the two
genera differ considerably.
Triplophyllum Holttum
Kew Bull 41:239 (1986).
As arranged by Holttum there are twenty species with a centre of distribution in
Africa; there are five species in tropical America and two in Madagascar. Most
species have been placed by recent authors in Ctenitis, but two have anastomosing
veins and so have been assigned to Tectaria, though their vein-branching is mainly
anadromous and their pattern of anastomosis is slightly different. Their scales are
Tectaria-like, also their spores. They all have a long-creeping caudex and fronds of
young plants are almost symmetrically tripartite. T. dicksonioides (Fée) Holttum in
Brazil has minute spherical glands on the lower surface.
Cyclopeltis J. Sm.
Bot. Mag. 72 (Comp.): 36 (1846).
Scales narrow, sometimes with dentate margins, the teeth consisting of out-
growths from single cells; fronds simply pinnate, the pinnae unlobed and articulate
to the rachis, sessile with cordate basiscopic bases which overlap the upper surface
of the rachis; veins free, their lower branches not nearly reaching the margin; very
short hairs consisting of several cells present on both upper and lower surfaces of
rachis and costae, more abundant on the lower surface; sori covered with peltate
indusia; spores similar to those of some species of Tectaria.
Fern-genera allied to Tectaria Cav. VI 161
A genus of about six species, rather uniform and widely distributed in tropical
America, more diversified in SE Asia and Malesia. The teeth on the margins of
scales, where present, differ from those of other palaeotropic genera, the teeth of
which are formed by the projecting common wall between adjacent marginal cells.
Megalastrum Holttum, gen. nov.
“Group of Dryopteris subincisa”’ C. Chr., K. Danske Vid. Selsk. Skr. VIII, 6: 59 (1920).
Caudex crassus, modice arborescens; stipes dense paleaceus, paleis breviter dentatis vel integris; lamina
frondis magna, copiose bipinnata, pinnulis profunde lobatis, pinnis infimis basin basiscopicam versus
auctis; venae omnes liberae, infimis supra basin sinuum inter lobos pinnularum terminatis; paleae
axium frondis non clathratae, dentatae vel integrae, plerumque multae; rhachides pinnarum costae-
que pinnularum supra pilosae, pilis acicularibus, pluricellularibus, in sicco non contortis; glandulae
cylindricae unicellulares desunt; indusia, ubi adsunt, magna, firma, pilosa; sporae minute spinulosae:
n = 41.
Type species: Megalastrum villosum (L.) Holttum, comb. nov. (Polypodium villo-
sum L., Sp. Pl. 1093 (1753).
In his monograph of 1920 Christensen included thirty neotropic species in this
group as part of his subgenus Ctenitis. John Smith, who had seen some of them in
cultivation at Kew, included them in the genus Lastrea, calling them ‘“‘the villosa
group’; he recognized their distinctive character and suggested (1875: 216) that
they might form a genus for which the name Megalastrum would be appropriate. In
this as in many other ways he showed greater understanding (gained by observation
of living plants) than most of his contemporaries, and I am happy to adopt his
suggested name. T.G. Walker (1966), in Jamaica, found that plants of the type
species of Megalastrum, each being the outgrowth of a single cell, as in Cyclopeltis.
found it to be tetraploid. Spores are illustrated on Plate 3.
One species should be excluded from Christensen’s original list; it was first
named Polypodium grande by Presl. In form, venation and thick texture, its fronds
are very similar to those of Ataxipteris (of China and Japan) but its scales are very
different; their marginal teeth are also differently formed from those of the type
species of Megalastrum, each being the outgrowth of a single cell, as in Cyclopeltis.
In Africa there is only one species of this genus, M. lanuginosum. In its indusia
and spores it agrees closely with M. villosum; its distribution extends to Madagas-
car and the Mascarene Islands, where there are four more species. These African
and Mascarene species do not have the dentate scales which are characteristic of
most species in the neotropics. New names for them are proposed as follows.
Megalastrum lanuginosum (Kaulf.) Holttum, comb. nov.
Aspidium lanuginosum Kaulf., Enum. Fil. @hamisso: 244 (1824).
Megalastrum magnum (Bak.) Holttum, comb. nov.
Nephrodium magnum Bak., J. Bot. 22: 142 (1884).
Megalastrum exaggeratum (Bak.) Holttum, comb. nov.
Nephrodium crinitum var exaggeratum Bak., Ann. Bot. 5: 319 (1892).
Megalastrum lanatum (Fée) Holttum, comb. nov.
Phegopteris lanata Fée, Gen. Fil. 246 (1852).
Megalastrum canacae (Holttum) Holttum, comb. nov.
Ctenitis canacae Holttum, Kew Bull. 38: 128 (1983).
162 Gard. Bull. Sing. 39(2) (1986)
Pseudotectaria Tard.
Notul. Syst. (Paris) 15: 87, pl. 6 (1955).
The type species, originally named Tectaria decaryana C. Chr., has in its supra-
basal pinnae a venation comparable to that in Meniscium (Thelypteridaceae) but in
the basal part of its basal pinnae the venation is more complex and more like that of
Tectaria sect. Sagenia (see C. Chr. 1932, pl. 19). The shape of the basal pinnae
shows some resemblance to Heterogonium, but the basal basiscopic lobes are not
gradually reduced. The outer veins of costal and costular areoles in the basal pinnae
of P. decaryana are not parallel to the costae and costules as in Tectaria sect.
Sagenia and Heterogonium,; they are formed by upcurved veins which in fertile
pinnae each bears a sorus, meeting to produce one or more excurrent free veinlets.
In the second species of Pseudotectaria, P. crinigera (C. Chr.) Tard., the pinnae are
distinctly lobed with rather widely spaced costules, and the venation throughout (as
shown in C. Chr. 1932, pl. 18) resembles that in the basal pinnae of P. decaryana.
The smaller scales in both species are abundant, narrow, thin, with all cells
elongate and showing a clear lumen, contrasting with the rigid opaque scales of
Tectaria. | suggest that the first four species of Christensen’s Dryopteris subg.
Ctenitis (1932: 57-58 and pl. 13, 14) are related to the species of Pseudotectaria and
should probably be transferred to it (Mme Tardieu-Blot mentioned this possible
relationship in 1955); they certainly do not belong to Ctenitis as understood in the
present conspectus.
Thus I suggest that Pseudotectaria represents a development of anastomosis
which has occurred on an evolutionary line distinct from that of Tectaria, compara-
ble with the parallel development in Triplophyllum.
Discussion
In view of the pantropic distribution of Tectaria and Ctenitis (the largest genera)
and the great range of form among the other genera, this group must have had a
long evolutionary history during which connecting links have disappeared. There
can be no doubt that Tectaria and the smaller genera here associated with it are
closely allied, and I suggest that there is good evidence for an origin of Tectaria in
SE. Asia, as free-veined species occur there from which, within the genus now
existing, there is every gradation to species with elaborate anastomosis of veins (in
Thailand, the distinction between sect. Tectaria and sect. Sagenia is not quite
sharp).
The genera which have unicellular cylindric glands are not a closely allied group.
Pleocnemia resembles Tectaria sect. Sagenia in venation but differs in the presence
of sinus-teeth and glands, in the former agreeing with Pteridrys (which lacks glands)
and in the latter with Crenitis. Another genus which lacks glands, Ataxipteris, has
the frond-form and venation of free-veined Tectaria but abundant scales very like
those of Ctenitis. Lastreopsis is near Ctenitis (in which it was included by Copeland)
in glands but more like Tectaria in scales and has more variable hairs and more
varied branching of fronds. The sole species of Coveniella has no clear resemblance
to any other genus; its simple pinnae and their venation are nearest to those of
Cyclopeltis and the glands on sporangium-stalks resemble those of Ctenitis, but
there are now no Australian species of either genus.
The situation is further complicated by the fact that Tectaria and closely related
genera have the chromosome number 40 (no counts yet for Psomiocarpa and
Aenigmopteris) whereas all the rest have 41. If we assume that 40 is the older
number, 41 may have developed on various evolutionary lines. But there are three
different genera which appear to show affinities with both Tectaria (40) and Ctenitis
(41): thus no simple evolutionary pattern seems possible.
@114 15K
=
wa
wa
on
®
—_
Ss
Spores of Megalastrum. M. lanuginosum: A (X 1750), B (x 3500); Gerrard 1933, Natal (K).
Plate 3.
163
(KK).
M. villosum: C (x 1750), D (X 3500); Bancroft s.n. 1850, Jamaica
164 Gard. Bull. Sing. 39(2) (1986)
The puzzle seems to me resolvable by assuming that on rare occasions there has
been genetic interchange between species of Tectaria and Ctenitis. The existence of
the single species of Ataxipteris (41) with some characters in common with Tectaria
and some with Crenitis looks like the result of such a process. Pleocnemia (41), with
19 species spread over a wide area, may be the result of a gene interchange which
took place at an earlier time.
I suggest that such interchanges may be summarized in the form of a diagram.
Hypothetical ancestral group
Tectaria (40) Cyclopeltis (41) Ctenitis (41)
se Het a Po .
re: allies Coveniella (41)
Heterogonium (40)
Psomiocarpa
Sats eee Ataxipteris (41)
more distant Pleocnemia (41)
Pteridrys (41) Lastreopsis (41)
Triplophyllum (41)
Pseudotectaria
Tectaridium Megalastrum (41)
sect. Tectarla-—_. Chlamydogramme
Notes on Neotropic Genera
Pichi Sermolli (1977) recognizes the following genera which I regard as allied to
Tectaria: Dictyoxiphium Hook., Amphiblestra Presl, Pleuroderris Maxon, Camp-
todium Fée, Fadyenia Hook. and Atalopteris Maxon & C. Chr. All these except
Atalopteris are monotypic and clearly related to Tectaria and are united to Tectaria
by Tryon & Tryon (1982: 470). Atalopteris is clearly related to Ctenitis and is
considered separately below.
Dictyoxiphium has simple fronds with the venation of Tectaria sect. Tectaria.
They have continuous indusiate submarginal sori, in which they agree with Chlam-
dogramme but the fertile ones are not contracted. Dictyoxiphium hybridizes with
Tectaria incisa Cav. to produce Pleuroderris (Wagner 1978). Amphiblestra has
deeply lobed fronds with the venation of Tectaria sect. Tectaria and continuous
submarginal exindusiate sori. The recognition of Dictyoxiphium and Amphiblestra
as distinct genera is largely a matter of convenience.
Camptodium has small opaque fronds with free veins arranged as in some
palaeotrophic species of Tectaria sect. Sagenia but is not nearly related to them. It
is specialized in adaptation to growth on limestone rocks in sheltered places.
Fadyenia is also reduced and specialized, with a rather irregular anastomosis of
veins not quite like that of sect. Sagenia.
The species first named Polypodium grande by Presl, referred to above under
Megalastrum, is very large but with a venation essentially similar to that of Camp-
todium, agreeing also in its opaque fronds but differing in its peculiarly dentate
scales. This species needs further study; I suggest that it should have generic rank.
Fern-genera allied to Tectaria Cav. VI 165
Another species which needs further study is Tectaria brauniana (Karst.) C. Chr.
This has deeply pinnatifid fronds of thin texture, its lobes again deeply pinnatifid:
its veins are free, arranged as in free-veined palaeotropic species of sect. Sagenia. It
is peculiar in its slender creeping caudex bearing thin translucent scales and in the
presence of minute sessile or subsessile spherical glands on the lower surface and on
indusia; these glands are similar to those of Triplophyllum dicksonioides (Fée)
Holttum and I have seen no similar ones on any other species of Tectaria.
Atalopteris has sterile fronds similar in form to those of the type species of
Ctenitis but differing in crenate pinna-lobes and the separation of one or more lobes
as pinnules on basal pinnae. Its fertile fronds have greatly contracted pinnae
bearing exindusiate sori in which the sporangium-stalks bear glands as in Ctenitis.
There are also abundant cylindrical glands on the lower surface of sterile pinnae.
The small scales on rachis and costae are thin with light brown cell-walls; I see no
isodiametric cells and the scales are not so clearly clathrate as those in Ctenitis
submarginalis.
Artificial Key to the Palaeotropic Genera
1. Teeth present at the bases of sinuses between pinna- or pinnule-lobes, the teeth projecting out of the
plane of the lamina (in some species present only in distal sinuses)
2. Fronds simply pinnate with free veins; no glands on lamina Or in SOT ....................0000 Pteridrys
2: Fronds mostly bipinnate, their veins forming at least costal areoles; cylindric or ovoid unicellular
glands present on stalks of sporangia and/or on lower surface of lamina ................... Pleocnemia
1° Teeth of this kind lacking
3. All axes of the frond bearing copious scales, the smaller ones at least partly clathrate with
isodiametric cells
4. Cylindric glands present on young indusia and elsewhere; pinnae (and usually pinnules) deeply
lobed, the basal basiscopic vein in each lobe arising from the costule of the lobe and ending
ne nae NE SANE SENSU an geet RE eee eo sve cei é-scembbemshordans Ctenitis
4° Cylindric glands lacking; pinnae less deeply lobed, the basal basiscopic vein in each lobe
usually arising from the costa of the pinna and ending below the base of the sinus ... Afaxipteris
3: Smaller axes of the frond bearing scales which are few (except in Megalastrum and Pseudotectaria)
and not thus clathrate; unicellular glands lacking except in Lastreopsis
5. Veins in sterile fronds anastomosing copiously; free veinlets present in areoles (including those
along the costa) variously directed and in most species forked
6. Fertile fronds greatly contracted and bearing indusia
7. Sterile fronds simple; fertile ones irregularly deeply lobed; indusia reniform Tectaridium
7: Sterile fronds pinnate; pinnae of fertile fronds linear, with continuous indusia along each
ieee ere, ke ne Re pee One» ee aot ee Chlamydogramme
6: Fertile fronds not greatly contracted, or if so lacking indusia ............ Tectaria sect. Tectaria
5‘ Veins anastomosing to form costal areoles which lack free veinlets, or all veins free
8. Some thick multicellular hairs present between veins on the upper surface, at least near
sinuses between lobes; where veins anastomose, the costal areoles narrow and of even width
9. Basal basiscopic lobe or pinnule of basal pinnae longer than the other lobes or pinules
10. Fronds not greatly longer than wide
11. Veins anastomosing in some species; fertile fronds, if greatly contracted, not
bearing Very small pinnulls ... 2.0.2.0 .2.. si. e cece rec een eee se sees Tectaria sect. Sagenia
11/ veins all free; fertile fronds bipinnate with very small pinnules; no indusia ........
ERA Re 12 ta7 Ae ETA kd Tati s een Sn Neveu tpn ont Lardwescbessevecsccewsneuseuens Psomiocarpa
10° Fronds greatly longer than wide, with many pinnae gradually increasing in size
MII, TEEN DUDLs . a < alas Cad p Saree Bayh dos so ye) <osaeed eo} eonaneae ees Aenigmopteris
9. Basal basiscopic lobe of basal pinnae much reduced, these pinnae widest at about
MINN ons 5 oii noi pet aiovwaduw chins suosp sexes ane dyercnaynenngvarasensnsanepagecsoens Heterogonium
8: No thick hairs present between veins on upper surface; where veins anastomose (/7!-
plophyllum and Pseudotectaria) the costal areoles not elongate nor of even width
12. Pinnae entire or slightly lobed; lateral veins in pinnae forming pinnate groups, basal
veinlets in each group, where free, not nearly reaching the margin
166 Gard. Bull. Sing. 39(2) (1986)
13.. Veinlets all free; pinnae articulated to rachis .......,.-0:-sesss; 62 ieee Cyclopeltis
13° One or more pairs of veinlets joining to form a short excurrent veinlet; pinnae not
articulated
14. Anastomosis confined to suprabasal veinlets .................cceeeee enone Coveniella
14° Anastomosis, where it occurs, including basal veinlets ............ Pseudotectaria
12° Pinnae various, in almost all cases at least deeply lobed; most free veins reaching or
nearly reaching the margin
15. Fronds very large, bipinnate, arising from a massive erect caudex; upper surface of
axes of frond bearing multiseptate thin-walled acicular hairs ............ Megalastrum
15° Fronds never very large, pinnae various; caudex not massive and erect; hairs on
upper surface of axes of frond either firm and terete or ctenitoid
16. Bases of leaflets decurrent as a narrow wing with thickened margin; fronds of
young plants not tripartite; hairs on upper surface of axes various; glands
present on lower surface and on Sproangia.......... 0.0.01 assneyeeeeeen Lastreopsis
16: Bases of leaflets not thus decurrent; fronds of young plants tripartite except in
T. varians; hairs on upper surface of axes always ctenitoid; no cylindric glands
PTOSEMG aio. ec es tgdalpe eden deans tacit eee a een ee Triplophyllum
Acknowledgements
I am indebted to the Director, Rijksherbarium, Leiden, for the loan of the type
of Hemigramma siifolia and permission to cut sections of its pinnae, to Dr Mary
Tindale (Sydney) and Dr S.K. Roy (Banaras Hindu University) for information
about the genus Coveniella, to Prof. E. Hennipman (Utrecht) for the preparation
showing venation in Hemigramma latifolia, and to Mr M.G. Price (Ann Arbor) for
Philippine specimens of Psomiocarpa and its hybrid with a species of Tectaria; also
to Mr Tim Lawrence (Jodrell Laboratory, Kew) for sections of fertile pinnae of
Hemigramma siifolia and Chlamydogramme hollrungii and to Mr P.J. Edwards
(Herbarium, Kew) for the SEM photographs of spores; to all of whom I express
grateful thanks.
Postscript
When writing the above in early 1986 I did not know of the publication by S.K.
Roy and A.N. Rao of a paper entitled A brief cytotaxonomic survey of Singapore
ferns (Journal of the Singapore Academy of Science, 14: 53-64, 1985). One
observation published in that paper is relevant to the present discussion of Tectaria
and allied genera.
On p. 60 is Roy’s report that a chromosome count from a plant of Heterogonium
sagenioides showed n = 41, illustrated in fig. 36, whereas in an Appendix to my
book on the ferns of the Malay Peninsula (1955) Manton had reported n = 40 for
the same species, with her figure 15. In writing the present paper I assumed
Manton’s n = 40 to be correct.
In view of this discrepancy, and of the fact that the illustration published by Roy
and Rao does not provide clear evidence, I wrote to Prof Manton to ask whether
she could add any further information. Her reply is that she has re-examined her
preparation from a plant originating in Singapore, in consultation with Dr T.G.
Walker (Newcastle upon Tyne), and that they confirm the original report of n = 40.
Dr Walker reports also that he made preparations from a plant of Heterogonium
pinnatum (Copel.) Holttum cultivated at Kew (origin G. Mulu, Sarawak, A.C.
Jermy 13318) and that this showed n = 80; Manton’s report on this species was n =
80-82, with a note that better material was needed.
Much more evidence on the chromosomes of species in this group of genera is
desirable.
Fern-genera allied to Tectaria Cav. VI 167
References
Ching, R.C. (1938). A revision of the Chinese and Sikkim-Himalayan Dryopteris
with reference to some species in the neighbouring regions, III. Bull. Fan Mem.
Inst. Biol. Bot. 8: 275-334.
Christensen, C. (1932). The Pteridophyta of Madagascar. Dansk Bot. Ark. 7:
1-253, pl. 1-80.
Copeland, E.B. (1908). New or interesting Philippine ferns. Philip. J. Sci. Bot. 3:
31-37, pl. 1-4.
(1928). Genus Hemigramma Christ, in Leptochilus and genera confused
with it. Philip. J. Sci. 37: 402-408.
Holttum, R.E. (1985). Studies in the genera allied to Tectaria Cav., IV. The genus
Ctenitis in Asia, Malesia and the Western Pacific. Blumea 31: 1-38.
(1986). Ibid. V. Triplophyllum, a new genus of Africa and America. Kew
Bull. 41:237-260.
Pichi Sermolli, R.E.G. (1977). Tentamen Pteridiphytorum genera in taxonomicum
ordinem redigendi. Webbia 31: 313-512.
Price, M.G. (1974). Nine new fern names. Kalikasan 3: 173-178.
Sledge, W.A. (1972). The tectarioid ferns of Ceylon. Kew Bull. 27: 407-424.
Smith, J. (1875). Historia Filicum. London, Macmillan.
Tardieu-Blot, M.L. (1955). Sur les Tectarioideae de Madagascar et des Mas-
careignes, avec description d’un genre nouveau Pseudotectaria. Notul. Syst.
(Paris) 15: 86-90.
Tryon, R.M. & A.F. (1982). Ferns and Allied Plants with special reference to
Tropical America. New York, Springer.
Wagner, W.H. & F.S. (1978) The singular origin of the Central American fern
Pleuroderris michleriana. Neotropica 10: 254-264.
Walker, T.G. (1966). A cytotaxonomic survey of the pteridophytes of Jamaica.
Trans. R. Soc. Edinb. 66: 169-237.
(1973). Evidence from cytology in the classification of ferns, in A.C. Jermy
et al. (eds): The Phylogeny and Classification of the Ferns. Bot. Journ. Linn.
Soc. 67, Suppl. I: 91-110.
A New Genus and Three New Species of Pteridophytes from
North Eastern Queensland
Mary D. TINDALE
Royal Botanic Gardens, Sydney, N.S.W., Australia
EFFECTIVE PUBLICATION DATE: 24 JAN. 1987
Abstract
A new genus, Coveniella (Dryopteridaceae), composed of a single species, C. poecilophlebia (Hook.)
Tind., comb. nov., is described. Two species of Lastreoposis (Dryopteridaceae) viz. L. tinarooensis and
L. walleri as well as a species of Diplazium (Athyriaceae) viz. D. queenslandicum are also described as
new.
Coveniella Tind., gen. nov. (Fig. 1)
Coveniella Tind.; a Lastreopsis differt lamina 1- pinnata, venis pro majore parte anastomosantibus, ex
parte omnino liberis, costulis 25-46-jugatis, prominulis, plus minusve parallelis, utroque latere venulis
4-7 obliquis instructis, venulis infimis liberis quarum una e costa exorta, in venulis 1-3 medianis ramulo
congruenti e costula contingua respondenti in venulam brevem plerumque liberam excurrentem con-
juncto, venulis superioribus plerumque liberis usque and marginem pinnae attingentibus, soris inter
costulas irregulariter biseriatim dispositis. — Type species: Coveniella poecilophlebia (Hook.) Tind.
comb. nov.
Coveniella poecilophlebia (Hook.) Tind. comb. nov. Fig. 1
Polypodium poecilophlebium Hook. Sp. Fil. 5: 14 (1863). — Type: Dunk Island, NE. coast of Australia,
Voyage of the Rattlesnake. Macgillivray (K, holo), n.v.
Terrestrial fern. Rhizome long-creeping, radial, dictyostelic, clothed with short,
very broadly ovate, dark brown, denticulate scales with 1-3 golden glands on the
margin. Stipes erect, never articulated to the rhizome, having several, distinct
vascular bundles, with the xylem strands of the larger adaxial vascular bundles with
hooked ends. Lamina I-pinnate, 15-30 cm long, 10-30 cm broad, chartaceous,
catadromous except in the lowest pinnae. Pinnae 12-23 cm long, 2-4.5 cm broad,
oblong or narrowly lanceolate, with 1-6 lateral pairs and a similar terminal pinna,
the lowest not reduced, the apex acuminate or rarely obtuse, the margin undulate
or crenate but serrate towards the apex, unequally cuneate at the base. Ctenitis-
hairs 5-12-celled, scattered along the rhachises, costae, costules and veinlets.
Rhachis not deeply grooved above, the median portion slightly elevated, clothed
above and below with stiff antrorsely curved, 3-4-celled hairs, the margins of the
groove confluent with the decurrent margins on the bases of the pinnae. Costae
raised on the upper surface, clothed below with linear or lanceolate, attenuated
scales with the margin fimbriate towards the base. Veins anastomosing for the
greater part, in part quite free. Costules 25-46-jugate, prominent, more or less
parallel, 3-6 mm apart, with 4-7 oblique veinlets on each side; with the lowest
veinlets free, one of them arising from the costa, in the 1-3 middle veinlets the
corresponding branchlet from the contiguous costule joined in a short, usually free,
excurrent veinlet, with the upper veinlets mostly free, reaching the margin of the
pinna. Sori orbicular, exindusiate, 0.4-0.8(-1.0) mm in diameter, mostly borne on
the middle of the veinlets or sometimes towards the apex of the veinlets or at the
169
170 Gard. Bull. Sing. 39(2)(1986)
junction of the veinlets, in 2 irregular rows between the costules. Sporangia with
13-17 indurated cells of the annulus; pedicel long, bearing 1-2 glandular, sessile,
oblong, unicellular, golden hairs. Spores bilateral, monolete, globoso-ellipsoidal or
almost globose, brown, with ruguloso-saccate perispores. Glandular, oblong,
appressed, unicellular, golden hairs sparsely clothe the lamina, rhachises, costae,
costules and veins.
Distribution. terrestrial in rainforests often near streams and lakes in the Cook
and Kennedy Districts of NE. Queensland.
Selected Specimens Examined: QUEENSLAND. Cook District: Iron Range, June 1948, L.J. Brass
19151 (BRI, K); Whitfield Range, July 1964, A.W. Dockrill W 10 (K, NSW). North Kennedy District:
Conway State Forest, between Airlie and Shute Harbour, June 1965, L.J. Webb and T.G. Tracey 7578
(BRI, CANB).
Chromosome Counts: 2” = 82, n = 41, (diploid). Voucher: S.F.R. 185, Down-
fall L.A., Queensland, A.W. Dockrill 1115 (NSW), (pers. comm. S.K. Roy).
This new monotypic genus is named in honour of Mr. Robert G. Coveny,
Botanical Collector at the Royal Botanic Gardens, Sydney, in gratitude for his
continued assistance in obtaining material for my researches on pteridophytes.
I am including Coveniella in the Ctenitis-Lastreopsis group of genera in the family
Dryopteridaceae, because it it characterized by the following features: (/) orbicular
sori; (2) several, small, distinct vascular bundles in the stipes; (3) multicellular
Crenitis-hairs with dark red septae on the fronds; (4) golden, 1-celled, glandular
hairs on the fronds as well as 1-2 on the long narrow pedicels of the non-setose
sporangia; and (5) basic chromosome number of 41. It differs from Crenitis and
Lastreopsis which have free veins in their decompound laminas, whereas in Cove-
niella the 1-pinnate laminas have a very complex, partly meniscoid venation. The
most outstanding characteristic of the new genus Is the nature of the basal veinlets
which are always free, ending blindly and one of them very often arises from the
costa between the costules.
C. poecilophlebia superficially resembles some members of the family Thelypter-
idaceae but the latter have needle-like hairs on the fronds, stipes with 2 vascular
bundles at the base soon uniting to form a single strand (U-shaped in section) and
the basic chromosome number ranges from 27 to 36 (except 33).
Lastreopsis tinarooensis Tind., sp. nov. Fig. 2
Lastreopsis tinarooensis Tind., a L. grayi D. Jones stipitibus non profunde sulcatis pilis typi Crenitidis
circiter 0.1 mm longis subtus vestitis, rhachidibus secundariis inconspicue alatis, pinnis quaternariis in
lobulos 3-5 spathulatos late dispositos usque ad 2.5 mm longos profunde divisis, sporangiis usque ad 50
per sorum, statim diagnoscenda. — Type: Queensland, Cook District, State Forest Reserve, 185,
Kalorama L.A., 17° 06’S 145° 35°E, on rocks in very shady humid creek, alt. 1000 m. B. Gray 1398
(QRS, holo; NSW, iso).
Lastreopsis tinarooensis Tind. is readily distinguished from L. grayi by the
shallowly grooved stipes clothed on the lower surface with Crenitis-hairs about 0.1
mm long, by the inconspicuously winged secondary rhachises, by the quaternary
pinnae deeply divided into 3-5 spathulate, widely spaced lobules up to 2.5 mm long
and by up to 50 sporangia per sorus.
Distribution. terrestrial in rainforests of the Cook District, NE. Queensland.
Specimens Examined: QUEENSLAND. Cook District: Tinaroo Hills, Atherton Tableland, D. Jones
(BRI 200744-5); Mt Lewis, Sept 1977, J.A. Armstrong 1073A (NSW); near Danbulla, ca 3400 ft [ca 1020
m], Nov 1942, S.T. Blake 14752A (BRI 165807).
Chromosome Counts: 2n = 82, n = 41. Voucher: J.A. Armstrong 1073A (NSW),
(pers. com. S.K. Roy).
|
20175
a-b from L.J. Brass
Fig. 1. Coveniella poecilophlebia (Hook.) Tind.
; b, portion of pinna with sori, X 3.
a. habit study, x '2
17]
ae an
=
Se en
Fig. 2. Lasteopsis tinarooensis Tind.
Ba, lowest primary pinna, x 74. Bb, tertiary pinna with sori, x 4. Bc, upper surface of
rhachises, X 4. Bd, rhizome, x V3
This species is closely allied to Lastreopsis grayi D. Jones in Muelleria 3: 245-249
(1977). The differences between these two species are discussed in detail by Jones
(/.c. p. 248), L. tinarooensis being cited as Lastreopsis sp. pending my publication
of this new taxon. Both species are figured (/.c. p. 247).
Lastreopsis walleri Tind. sp. nov. Fig:
Lastreopsis walleri Tind.; a L. tenera (R. Br.) Tind. rhizomate valde robusta breviter repenti 1-2.5 cm
crasso, rhizomatis stipitis rhachidisque paleis margine et pagina setis numerosis instructis et apice
excepto pinnulis basiscopicis sessilibus nullis ad rhachidem primariam adnatis statim diagnoscenda. —
Type: Queensland, Cook District, Allumbah, (Herberton) Feb. 1910, R.F. Waller NSW 1576 (NSW,
holo).
Lastreopsis walleri Tind. is readily distinguished from L. tenera by its very
robust, shortly creeping rhizome 1-2.5 cm broad, by the numerous setae on the
margin and surface of the scales of the rhizome, stipe and rhachis, and except at the
apex by the lack of the basiscopic sessile pinnules adnate to the primary rhachis.
Distribution. terrestrial in rainforests of the Cook District, NE. Queensland.
Specimen Examined: QUEENSLAND. Cook District: $.F.R. 251, Charmillin L.A., 17° 40°S, 145°
30°E, alt. 750 m, Sept 1976, A.M. Dockrill 1268 (BRI 228508-14).
The channel on the upper surface of the main rhachis is poorly defined except
near the base, and the 2 ridges on the upper surface are also less prominent than in
172
Fig. 3.
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——
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Lastreopsis walleri Tind.
a, two primary pinnae, X *; b, adaxial surface of two tertiary pinnae, X 4; c, abaxial surface of
two tertiary pinnae with sori, X 4; d, indusium, X 33; e, adaxial surface of rhachises, x 4; f,
rhizome, X 7: g, scale of rhizome, X 8: / marginal cells of rhizome scale, x 33.
173
Fig. 4. Diplazium queenslandicum Tind.
Ba, two fertile secondary pinnae, x 7. Bb, tertiary pinna with sori, x 4.
the other species of Lastreopsis except L. tenera. However these 2 ridges are
continuous with the slightly thickened leaf-margins of the ultimate segments of the
primary pinnae as in all species of this genus.
Diplazium queenslandicum Tind. sp. nov. Fig. 4
Diplazium queenslandicum Vind: a Diplazio assimili (Endl.) Beddome differt frondibus maioribus,
atroviridibus, saepe 150-200 cm longis, 75-90 cm latis, caudice erecto, crasso, usque | m alto et 15 cm
diametro, paleis caudicis obscuris, maioribus, 9-17 mm longis et 2.5-5 mm latis; pinnis secundariis
numerosioribus, caudatis et 19-33-jugis; pinnulis tertiariis truncatis vel late rotundatis, 6-14-jugis,
pinnulis tertiariis infimis plerumque 6-12 mm longis et 3-6 mm latis. — Type: Queensland. Cook
District: Palmerston National Park, growing along old snigging track in rainforest ca | mile [1.6] km S.
of Highway and ca 20 miles [32.3 km] WSW. of Innisfail, alt. ca 2100 ft [630 m] 14 Sept. 1960, L.S. Smith
11270 (BRI 29863-5, holo; NSW, iso).
Diplazium queenslandicum Tind. differs from Diplazium assimile in its larger,
dark green fronds 150-200 cm long and 75-90 cm broad, in its erect, thick caudex up
to 1 m high and 15 cm in diameter, in its larger, dull caudex scales 9-17 mm long and
2.5-5 mm broad, by the more numerous, caudate secondary pinnae 19-33-jugate,
by the truncate or broadly rounded 6-14-jugate tertiary pinnules and by the lowest
tertiary pinnules usually 6-12 mm long and 3-6 mm broad.
Distribution. terrestrial in rainforests of the Cook District, north-eastern
Queensland.
Selected Specimens Examined: QUEENSLAND: Cook District: Tinaroo Range, alt. 3500 ft [ca 1050
m]. Feb. 1962, L.J. Webb and J.G. Tracey 5779 (BRI); back of Bartle Frére, July 1913, W.W. Watts
NSW P978; Mt Spurgeon, Root Creek, Sept 1936, C.T. White 10607 (BRI).
D. queenslandicum is closely allied to D. assimile which occurs in Norfolk Island
and Australia (south-eastern Queensland and north-eastern New South Wales).
The latter species is a smaller fern ca 40-120 cm high, with light green fronds and an
erect caudex (up to 4 cm high) bearing lustrous scales 5-8 mm long and 1.5-3 mm
broad.
Three new fern species of north eastern Queensland 175
Acknowledgements
Valuable assistance was given by the Director, Queensland Herbarium, for
providing the illustrations as well as material on loan for study. My thanks are due
to Mr. S. B. Andrews for his generous botanical co-operation. I am grateful to the
late Mr. H.K. Airy Shaw, Royal Botanic Gardens, Kew, England, for kindly
checking my Latin diagnoses except for that of Lastreopsis tinarooensis which was
checked by Dr. A. Kanis, Australian National Herbarium, Canberra. I wish to
thank Professor R.E. Holttum for examining the holotype and other material of
Coveniella poecilophlebia in the Herbarium, Kew, England, at my request. My
thanks are also due to Professor S.K. Roy, Banaras Hindu University, Varanasi,
India, for allowing me to publish his chromosome counts of two species. The
illustrations were drawn by Ms. M. Saul except that of C. poecilophlebia which was
prepared by Mr. W. Smith.
I wish to acknowledge former grants of financial support from the Utah Founda-
tion and the Australian Biological Research Survey for providing technical assist-
ance in my research work on Australian pteridophytes.
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Notes on Asiatic Cassine L. (Celastraceae)
A.J.G.H. KOSTERMANS
Biotrop-Seameo & Herbarium Bogoriense, Bogor, Indonesia
EFFECTIVE PUBLICATION DATE: 24 JAN. 1987
Abstract
Three endemic species, i.e., Cassine glauca (Rottb.) O.K., C. balae Kosterm. sp. noy., and C.
congylos Kosterm. sp. nov. occur in Sri Lanka. Elaeodendron glaucum Roxb. is conspecific with C.
balae, having a woody, very thick putamen and hardly any mesocarp. C. glauca O.K. has a thin putamen
and plenty of juicy mesocarp. C. congylos is based on Elaeodendron glaucum var. montanum Thw.
The commonest species in South India is C. albens (Retz.) Kosterm. comb. nov. (synonyms: C.
roxburghii (W. & A.) Ramanoorthy and Neerija dichotoma Roxb.); much rarer is C. paniculata (W. &
A.) Ramanoorthy.
A North Indian species is C. grossa (Roxb.) Kosterm. comb. nov., based on Euonymus grossa Wall.
ex Roxb., in North India it grows up to 2000 m altitude.
The Javanese species, originally described as Elaeodendron glaucum var. macrocarpum Kds. & Val. is
raised to specific rank as Cassine koordersii Kosterm. sp. nov. The Moluccan species represents C.
obiensis Kosterm., but the species from Timor island is kept separate 1c elliptica (Dec.) O.K.).
Introduction
The rediscovery of the true Cassine (Elaeodendron) glauca (Rottb.) O.K. in the
beach forest near Trincomalee on Sri Lanka’s West coast started a train of inves-
tigations and conclusions, making the treatment of the Malesian species by Ding
Hou (1962) untenable.
The confusion started with Lawson’s uncritical lumping (1875) of some quite
distinct species, a procedure accepted by Ding Hou (1962) and Matthew (1983)
although Wight and Arnott (1834) and even earlier, Graham (1830) had already
pointed out, that true C. glauca does not occur outside Sir Lanka. The confusion
was compounded by Roxburgh’s description of what he thought was C. glauca, a
tree grown in the Botanic Garden of Calcutta from seed obtained from Sri Lanka.
Nobody seems to have noticed that the name Elaeodendron roxburghti W. & A.
is illegitimate as Neerija has priority.
The North Indian species, as described by Brandis (1906) a.o., is again different
and might be conspecific with our C. grossa.
Contrary to Ding Hou’s contention that fruiting material cannot or can only be
ascribed to its proper species with difficulty, my conclusion is that the fruit has the
most important specific characters.
Two groups can be recognized, one having juicy fruit with abundant juicy
mesocarp and a thin, crustaceous putamen, differently shaped in the various
species and the other having hardly any mesocarp (dry fruit) and a very thick, hard
putamen.
Other distinctive characters are the colour and shape of the fruit, and, the colour,
shape, and especially the size of the petals. The leaves vary much in the same
species, the length of the petiole and the shape of the incisions of the leaf serrations
may be used with some care.
Received | July 1985.
177
178 Gard. Bull. Sing. 39(2)(1986)
The name Elaeodendrum is attributable to Murray, Syst. Veg., ed. 14 (1784) 241;
the orthography was changed to Elaeodendron by Jacquin f., Nov. Act. Helv. 1
(1787) (cf. Backer & Bakh., Fl., Java 2 (1965) 55, in note).
We have not discussed the merits and demerits of fusing Elaeodendrum and
Cassine. In the first edition of Engler & Prantl., Nat. Pfl. fam. (1892), Loesener has
followed O. Kuntze (1892) in combining the two, but dissociated them again in the
second edition (1942) as did Robson (Bolet. Soc. Broter. 39 (1965) 37). Since the
differences between the two genera are trifling and hardly applicable at the generic
level, we have followed O. Kuntze’s and Ding Hou’s example of fusing them.
INDIAN SPECIES
Cassine grossa (Wall. ex Roxb.) Kosterm., comb. nov.
Euonymus grossa Wallich ex Roxburgh, Fl. Ind., ed. Wall. 2(1824) 408; Voigt, Hort. suburb. Calcut.
(1834) 65; Lawson in Hooker f., Fl. Brit. Ind. 1(1875) 623 (as a syn. of Elaeodendron glaucum
(Rottb.) Pers.) — Typus: Wallich Cat. 4291 (K), n.v., from Nepal, fls. small, greenish.
Elaeodendron glaucum Auct. (non Pers.) Lawson, l.c., p.p.; Auct. (non Pers.) Collett, Fl. Siml. (1902)
88; Auct. (non Pers.) Brandis, Ind. Trees (1906) 164, p.p.; Auct. (non Pers.) Kanjilal, For. Fl.
Siwalik and Jaunsar (1911) 95; Auct. (non Pers.) Parker, Fl. Punjab, ed. 2(1924) 81; Auct. (non
Pers.) Osmaston, For. Fl. Kumaon (1927) 98; Auct. (non Pers.) Benthall, Trees Calcutta (1946) 112,
p.p.; Auct. (non Pers.) Prain, Beng. Pl. 1(1963) 230.
Cassine glauca Auct. (non O.K.) Hara, Fl. E. Himalaya (1966) 188; Auct. (non O.K.) Sidiqui in FI.
Pakistan 109(1977) 2.
Note. The only specimen available for examination (BO) was a flowering one
from a cultivated plant in the Calcutta Botanic Garden, of which leaf and flower
match the description. The fruit of this species is unknown. It might be conspecific
with the material described by Brandis (1906), Kanjilal (1911), Parker (1924),
Osmaston (1927), Benthall (1946) and Prain (1963), described erroneously under
Elaeodendron glaucum. The fruit is described as yellowish green when ripe, small
and with a thin putamen.
Cassine paniculata (W. & A.) Ramanoorthy
Ramanoorthy in Saldanha & Nicholson, Fl. Hassan Distr., Karnataka (1976) 308. — Elaeodendron
paniculatum Wight & Arnott, Prodr. Fl. Ind. or. (1834) 157; Lawson in Hooker f., Fl. Brit. Ind.
1(1875) 623 (as a syn. of Elaeodendron glaucum); Gamble, Fl. Madras (1910) 212(152). — Typus:
Colemale, fl., Herb. Wight.
Note. The species was known only in the flowering stage. Ramanoorthy (pro-
vided that his disposition is correct) described the fruit as fleshy, elliptic, the ovary
trilocular, the petals spathulate, 4.5-5 mm long, the margin reflexed towards the
base; according to him a very rare species.
Gamble differentiates it from Elaeodendron glaucum (non O.K.) (= Cassine
albens) by its large flowers, 0.4 inch in diam. and stout up to 4-inch long cymes, in
C. albens the flowers are only 0.25 inch in diam. and the cymes under 2 inch long.
Cassine albens (Retz.) Kosterm., comb. nov. Fig. 1
Schrebera albens Retzius, Observ. Bot. 6(1791) 25, t. 3; Willdenow, Spec. Pl. 1(1791) 1092; Roxburgh,
Pl. Corom. 2(1798) 2 (in nota); Fl. Ind. 1(1832) 638 (as a syn. of Elaeodendron glaucum sensu Roxb. );
Persoon, Syn. 1(1805) 241 (as a syn. of Elaeodendron glaucum); Lawson in Hooker f., Fl. Brit. Ind.
1(1875) 623 (as a syn. of Elaeodendron glaucum). — Typus: Koenig s.n., fl., fr., Coromandel (Lund).
Neerija dichotoma Roxburgh, FI. Ind., ed. Wall. 2(1824) 444; ed. Carey 1(1832) 646; repr. (1874) 217;
Wight & Arnott, Prodr. Fl. Ind. or: (1834) 157 (as a syn. of Elaeodendron roxburghii W. & A.);
Lawson, I.c. 623 (as a syn. of Elaeodendron glaucum); Ding Hou in FI. Males, Ser., 6(1962) 286 (as a
syn. of Cassine glauca). — Typus: Roxburgh s.n., coast of Coromandel(?).
ig Atle » Ger bheliims 4ok: | ty i
#4
Fo
BRIG ecbartheacitishowis a
5cm
Biagaten em FL {6 7%
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Fig. 1 Cassine albens (Retz.) Kosterm., holo (Lund).
179
180 Gard. Bull. Sing. 39(2)(1986)
Rhamus nerija Sprengel, Syst. Veget. 4, Suppl. (1821) 86; W. & A., I.c.; Lawson, l.c. (as a syn. of E.
glaucum).
Elaeodendron roxburghii Wight & Arnott, Prodr. l.c.; Graham, Catal. Bombay PI. (1830) 50; Dalzell &
Gibson, Bombay FI. (1861) 48; Wight, Illustr. (1820) 178, t. 71 (excellent plate); Beddome, FI. sylv.
(1871) 148; Lawson, I.c. 623 (as a syn. of E. glaucum); Gamble, Fl. Madras, I.c. 211 (as a syn. of E.
glaucum); Ding Hou, l.c. (as a syn. of Cassine glauca); Ramanoorthy in Saldanha & Nicholson, FI.
Hassan Distr., Karnataka (1976) 318; Matthew,, Fl. Tamilnadu, Carnatic 1(1983) 254 (as a syn. of
Cassine glauca var. glauca). — Elaeodendron glaucum var. roxburghii Lawson, |.c. 623. — Cassine
glauca var. roxburghii Pierre, Fl. For. Cochinch. 4(1893), t. 296 A. — Cassine roxburghii (W. & A.)
Ramanoorthy, l.c.
Elaeodendron glaucum Auct. (non Pers.) Lawson, l|.c., p.p.; Auct. (non Pers.) Cooke, Fl. Bombay
1(1902) 233; Auct. (non Pers.) Gamble, I.c. 211(152); Auct. (nom Pers.) Ding Houses ae
Celastrus glaucus (Rottb.) Vahl, Symb. Bot. 2(1791) 42 (quoad descript. p.p.); Auct. (non Vahl)
Roxburgh, Fl. Ind., ed. 2, 1(1832) 638; Auct. (non Vahl) Lawson, I.c.; Auct. (non Vahl) Ding Hou,
Lc;
Cassine glauca Auct. (non O.K.) Ding Hou, I.c.; Auct. (non O.K.) Ramaswamy & Razi, Fl. Bangalore
Distr. (1973); Auct. (non O.K.) Ramanoorthy, I.c. 318; Auct. (non O.K.) Matthew, I.c. 254 (var.
glauca), vol. 2(1982), t. 141; Roxburgh’s plate no. 73 (British Museum, n.v.).
Vernacular name. Nerija (Telinga language ex Roxburgh).
Distribution. South India, Courtallam, Nilgirris, Malabar, Coromandel, perhaps
further northwards to Mysore. Common, but scattered.
Massive, but usually a short-boled tree, up to 25 m tall and 60 cm dbh. Bark
smooth, grey, finely cracked; live bark up to 4 mm thick, outside reddish, inside
yellowish. Leaves opposite and somewhat scattered, oval to oblong, usually se-
rrate-crenate. Panicles axillary and extra-axillary, many-flowered with very thin,
patent dichotomous branches. Flowers yellowish, not exceeding 6 mm in diam.
Drupe one-celled by abortion, obovoid, red when ripe, succulent with a soft, thin,
crustaceous putamen.
Note. Retzius’ Schrebera albens was based on a specimen collected by Koenig, in
flower and fruit, now preserved in the Lund Herbarium in Sweden. On the verso of
the type sheet the caption is similar to that of the type specimen of Mangifera
glauca (Cassine glauca O.K.) Rottb. of the Copenhagen herbarium with a slight
difference. The habitat is simply marked as Ceylon and Coromandel. The specimen
is correctly depicted by Retzius showing flowers and fruit which are more elabor-
ately described by Retzius than those of Mangifera glauca by Rottboell.
The two specimens, one in Copenhagen and the other in Lund, differ in their
fruit characters and since the rediscovery of the entirely Ceylonese Cassine glauca,
it has become obvious that Koening had sent to Rottboell in Copenhagen a
specimen collected in Sri Lanka and another to Retzius in Lund collected in India
and this he had captioned on the label of the Copenhagen specimen as “In the
forests of Ceylon and near heathen temples in Coromandel’.
It was Graham (1830) who first expressed his doubt, 1.e., whether the Ceylon
plant also occurred in India and Wight and Arnott were the ones who, with the
Same opinion, separated the two south Indian species into Elaeodendron roxburghii
(based on Neerija dichotoma Roxb.) and E. paniculatum.
Wight and Arnott were, however, unfamiliar with the true identity of Elaeoden-
dron glaucum of Sri Lanka. They accepted Roxburgh’s Elaeodendron glaucum as
being conspecific with Rottboell’s Mangifera glauca.
After the rediscovery of the true Cassine glauca (Rottb.) O.K., it is known now,
that the tree, which Roxburgh obtained from Ceylon (in this paper named Cassine
balae Kosterm.) is not identical with the species that Rottboell described.
Asiatic Cassine 181
Cassine albens has the same kind of inflorescence with extremely thin branches as
that of C. glauca; the flower is also similar in size but white, and, only half the size
of those in the Ceylonese C. balae. Roxburgh stated that the fruit was red, different
from the pure white one of C. glauca but the main difference is in the shape,
obovoid to obovate-ellipsoid in C. albens and spindle-like with both ends strongly
pointed in C. glauca. Both have a thin crustaceous putamen and sufficient meso-
carp.
Of C. albens I have collected a specimen near Courtallam in South India in a
rather dry forest higher up than the village, where also Wight’s specimen was
collected in July 1835.
After the lumping of several distinct species by Lawson (1875), it was only
Ramanoorthy (1976) who interpreted the South Indian species correctly, referring
them to Cassine as C. roxburghii and C. paniculata; the author missed the point
that the specific epithet of the first should have been dichotoma (from Neerija
dichotoma Roxb.).
Ding Hou (1962) and Matthew (1983) stuck to Lawson’s concept and Vahl based
the name Celastrus glaucus on Mangifera glauca Rottb., but the description (Ding
Hou stated wrongly that the name was a nomen nudum) was based on Koenig’s
specimen in Lund, the base of Schrebera albens. Brandis’ specimen of Elaeoden-
dron glaucum (non O.K.) is not conspecific with Cassine albens, because of the
yellowish brown petals and yellowish green fruit. This seems to be the same species
as that described by Kanjilal, Osmaston, Benthall, Collett and Prain (Cassine
grossa). Parker’s described specimens have a large thickening at the base of the
petals; they are something else.
Pittard’s species from Indochina belongs to the group with a thick, woody
putamen and is allied with Cassine balae.
Southern Tamilnadu, Courtallam, fr., Kostermans 26003 (AAU, L); Nilgirris, fl., Hooker f. &
Thomson s.n. (L); Concan, fr., Stocks s.n. (L); locality not indicated, fl., Herb. Wight 461 (L).
SPECIES OF SRI LANKA
Cassine glauca (Rottb.) O.K. Figs. 2 & 3
O. Kuntze, Rev. Gen. Pl. 1(1891) 114; Auct. (non O.K.) Pierre, Fl. For. Cochinch. 4(1893), t. 296 A;
Auct. (non O.K.) Loessener in Engler & Prantl, Nat. Pfl. fam. 3(5) (1892) 214; ed. 2, 20 b (1942) 173;
Auct. (non O.K.) Ding Hou in Fl. Males., Ser. I, 6(2) (1962) 286. — Mangifera glauca Rottboell, Nye
Saml. Videnskabl. Selskabs Skrifter (Nov. Act. Haffn.), Anden Deel (vol. 2) (1783) 533-555, tab. IV,
f. 1 (exclud. Melmedya); Roxburgh, Fl. Ind., ed. Carey 1(1832) 638 (as a syn. of Elaeodendron
glaucum); Lawson in Hooker f., Fl. Brit. Ind. 1(1875) 623 (as a syn. of E. glaucum); Ding Hou, l.c.
(as a syn. of Cassine glauca). — Celastrus glaucus (non R. Br.) (Rottb.) Vahl, Symbol. bot. 2(1791) 42
(exclud. descript., which is Cassine albens); Moon, Catal. (1824) 17; Thwaites, Enum. Pl. Zeyl.
(1858) 73 (as a syn. of Elaeodendron glaucum); Trimen, Hand. Fl. Ceylon 1(1893) 271, p.p. (as a syn.
of E. glaucum); Alston in id. 6 (Suppl.) (1931) 42, p.p. (as a syn. of E. glaucum). — Elaeodendron
glaucum (Rottb.) Persoon, Syn. 1(1805) 241; Thwaites, I.c. (exclud. var. montanum); Lawson, l.c.
623 (pro minime parte); Trimen, I.c., p.p.; Auct. (non Pers.) Loessener, I.c.; Weeratunga, Kumar,
Sultanbawa & Balasubramaniam in Proc. Assoc. Adv. Sci. (1983) Sec. E, p. 74.
Senacia glauca Lamk., Dict. Sci., Suppl. (1817) 128. — Senacier glauque Lamarck, Illustr. Genres
2(1798) and in Dict. Sci. Suppl. (1817) 128. — Typus: Konig s.n., fl., fr., e Ceylon (C).
Usually small trees, glabrous in all their parts. Bark smooth, greyish white, thin,
with yellowish underside; live bark red. Leaves opposite, the young ones conspI-
cuously acuminate. Panicles with very slender peduncles and very thin, dichoto-
mous, patent branchlets (much thinner than in C. balae). Flowers green, ¢. 5 mm in
diam.; petals with white tips. Fruit narrowly ellipsoid, glossy porcelain white with
1-mm thick, soft, juicy, sweetish mesocarp and thin, crustaceous, spindle like, pale
yellowish putamen, conspicuously pointed at both ends, one-seeded by abortion.
Velastracege
Det. B.D. Merritt
Asxoin Annonerun, Hanvants Unt
tis
182
VEBSITY
PUN OU Gg 7
CHA
Botanical Museum of the
University of Copenhagen
Negra §6=— 9G
Fig. 2. Cassine glauca (Rottb.) O.K., holo (C).
r tee
be
Asiatic Cassine 183
History. The paper on Mangifera glauca was read by Rottboell on December 13.
1782 and published in 1783. It was based on a specimen, sent to him by Koenig, a
physician, employed by a Missionary Station in South India. The type sheet in the
Copenhagen Herbarium consists of a large flowering and fruiting branch of which
the lower right hand part was depicted by Rottboell. The specimen is marked on
the verso of the sheet: ‘“‘Ex India orientalis. Genus novum. Arbor Pentandra
Monogynia. Flore infero. S(t)ylus conica. Fructus baccata drupacea nux semibilo-
cularia. Habitat Zeylone nemorosis, in Coromandelia prope templa idolatrorum.
K(oenig).”’.
This caption has caused confusion. It has been interpreted that Koenig collected
from a single tree in Ceylon and had seen (what he believed to be) the same species
grown near a temple in South India.
This is now shown to be incorrect. At least two specimens are involved, the one
sent to Rottboell, originating from Sri Lanka and another one sent to Retzius,
which was from the coast of Coromandel (described by Retzius as Schrebera albens
Ketz.).
Rottboell believed his specimen to be a Mangifera. He depicted the slender,
elongate, slightly curved fruitlets, but did not describe them to a great extent.
Retzius presented a better description of the fruit of his specimen (which represents
Cassine albens Kosterm.).
When Vahl (1791) had before him Retzius’ specimen he recognized its rela-
tionship with Rottboell’s Mangifera glauca, at the same time inserting it in the
proper family as Celastrus glaucus Vahl. He presented a lengthy description (hence
not as Ding Hou stated that Celastrus glaucus is a nomen nudum) which indicates
that he described Retzius’ specimen (of. under Cassine albens Kosterm.).
Some time later, Roxburgh (Hort. Bengal. 18(1814)) (cf. also Voigt, Hort.
suburb. Calcut. 67(1845) received seeds from a Ceylonese Elaeodendron (Cassine)
species, which subsequently was grown in the Calcutta Botanical Garden and which
he described as Elaeodendron glaucum, apparently assuming that there was only
one species of Elaeodendron in Sri Lanka (it is, however, another, distinct species,
named here Cassine balae Kosterm.) (cf. under that species.).
In the same Flora of Roxburgh Neerija dichotoma was described, which is
conspecific with Schrebera albens Retz. This species was later correctly referred to
Elaeodendron by Wight & Arnott (1824) although with the illigitimate name E.
roxburghii (cf. under C. albens Kosterm.).
Rottboell added in synonymy the vernacular name Helmedija, a misspelling of
Waelmedija (wael= climber) of Linnaeus, FI. Zeyl. (1743) 204, no. 439 (the name
is of Hermann, Mus. Zeyl. 13; the specimen is on folio 76 of Hermann’s Herbarium
in the British Museum, Natural History London. It represents Hippocratea indica
Willd.). It was Retzius who already doubted its conspecificity with Mangifera
glauca Rottb.
Wight & Arnott (1814) were the first to note that the Ceylonese Elaeodendron
glaucum (sensu Roxburgh) did not occur outside of Sri Lanka and named the South
Indian species Elaeodendron roxburghii and E. paniculatum. The only one, who
heeded this was Ramanoorthy (1976) who excluded Elaeodendron glaucum trom
India and named the South Indian species Cassine roxburghii and C. paniculata.
Lawson (1875) uncritically lumped all species from Sri Lanka and India under
Elaeodendron glaucum yet recognizing two varieties: roxburghii and one unnamed
from Canara, which is perhaps not a Cassine.
Assuming that Roxburgh had been correct in that his plant (introduced from Sri
Lanka) represented Rottboell’s Mangifera glauca (Cassine glauca O.K.), Thwaites
and Trimen (1893) must have named the only then known Ceylonese species:
LAOGAF
Me Fe
1 Fe)
Ny Wy 4] nl)
a x f
4
AR Up 3 =
EVER A
AQ WH YC
SAAN \ oy, :
Ye \ ee” Wi
SS ‘
Fig. 3. Mangifera glauca Rottb., original drawing (lower figure).
184
Asiatic Cassine 185
Elaeodendron glaucum (now C. balae Kosterm.). Thwaites had separated the
variety montanum, which I have raised to specific rank as C. congylos Kosterm.
The specimen C.P. 1227 is the true C. glauca.
Ding Hou (1962) followed Lawson’s concept, not aware of the salient differences
of the fruit characters, evident in the figures of Rottboell (Cassine glauca) and of
Wight (1820) (Cassine albens), and Roxburgh’s description. After the correct
disposition of Ramanoorthy (1976), Matthew (1983) reverted to Lawson’s faulty
conception.
The rediscovery of the real Cassine glauca (Rottb.) O.K. took place in the
garden of the Nillavely Beach Hotel near Trincomalee on the west coast of Sri
Lanka, which consists of the original beach forest between the Hotel and the beach,
of which the hotel owner had had the good sense of cutting out only the under-
brush, in this way creating a shady garden, where, while seated and sipping a glass
of beer, I found that the tree above me provided the key to all this confusion and
misinterpretation.
Contrary to Ding Hou’s statement that fruiting specimens of Cassine are difficult
to identify I have arrived at the conclusion that it is the fruit that provides the
distinguishing characteristics.
There are two groups in Cassine, one with juicy fruit with an ample mesocarp and
a thin, crustaceous putamen (Cassine glauca, C. albens and perhaps C. paniculata)
and a second, without any appreciable mesocarp and a very thick, woody putamen
(Cassine balae, C. congylos, perhaps some north Indian Cassine, and the Malesian
species). The Malesian Cassine viburnifolia falls in the first group.
The slender, elongate, glossy white, juicy fruitlets of Cassine glauca with their
slender, spindle-like, thin putamens, gradually, strongly pointed at the ends, can-
not be confused with fruits of other known Cassine species.
Chemistry. The leaves have a sternutatory and fumigatory action. They are also
used as a snuff to relieve headaches. The gum from the tree dissolves in water to
give an adhesive. The seeds and bark contain seven friedelane derivates, these
include friedelin, canophyllal, friedelan-3-on-25-1, friedelan-3-01, elaeandrol (17-
dihydroxy-28-norfriedelan-3-one), canophyllol, friedelan-3-on-25-1 and the trioxy-
genated friedelane elaeodendradiol (17,25-hydroxy 28-norfriedelan-3-one).
Later G. Weeratynga, V. Kumar, M. Uus Sultanbawa, S. Balasubramaniam
(section E, page 74) found three other trioxygenated fridelane derivatives: ketodiol
and a dioxo-alcohol (25,28-dihydroxy friedelane-3-one) and dioxoalcohol (3,28-
dioxo friedelane-25-1).
Sri LANKA. Beach near Trincomalee, Dry Zone, beach forest, April, fl., fr., Kostermans 27719
(AAU, G, L); Madu Rd. to Mannar Isl., Dry Zone, low, May, fl., Kostermans 24887 (AAU, G, L):
Jaffna Distr., along main road, Ghavakachari, low, Aug., fr., Balasubramaniam 2189 (AAU, PDA);
Puttalam Distr., Manampuri, low, dry zone, May, fl., Cramer 4665 (L, PDA); Beriliya forest near
Elpitiya, Galle Distr., low, fl., fr., fr. June, Kostermans 27790 (AAU, PDA), Kostermans 24887 (BO):
locallity not indicated, fl., C.P. 1227 (BO, PDA).
Cassine balae Kosterm., sp. nov.
Elaeodendron glaucum Auct. (non Pers.) Roxburgh, Coromandel PI. 2(1798) 2; Hort. Bengal. (1814)
18; Fl. Ind., ed. 2, 1(1832) 638; repr. Carey’s edit. (1874) 214 (exclud. Schrebera albens Retz.,
Celastrus glaucus Vahl, and Mangifera glauca Rottb.); Auct. (non Pers.) Voigt, Hort. suburb. Calcut.
(1834) 167; Trimen, Handb. Fl. Ceylon 1(1893) 271, p.p. (exclud. Schrebera albens Retz., Celastrus
glaucus Vahl, Elaeodendron roxburghii W. & A.); Lawson in Hooker f., Fl., Brit. Ind. 1(1875) 628,
pro minime parte, quoad cit. Ceylon; Auct. (non Pers.) Ding Hou in Fl. Males., Ser. 1, 6(2) (1962)
286.
186 Gard. Bull. Sing. 39(2)(1986)
Elaeodendron balae Kosterm. ex G. Weeratunga, V. Kumar, M.U.S. Sultanbawa & S. Balasubrama-
niam (invalid, no latin diagnosis), J. Chem. Soc. Perkin Trans. 1(1982) 2457.
Arbor in omnibus partibus glabra, follis oppositis, sub-coriaceis, obovato-ellipticis mucronulatis vel
obtusis, margine serrato-crenulatis, basin versus sensim cuneastis, nervo mediano supra plana vel
subimpressa, subtus graciliter prominula, costis paucis, paniculis axillaribus multi-floris dichotomis,
petalis tenuibus oblongis obtusis 2-3.5 mm longis, ramulis sat crassis, fructibus maturis subglobosis
mucronatus, mesocarpium deest, putamen crassum lignosum. Typus: Balasubramaniam 2213 (AAU, K,
L, holo).
Tree. up to 20 m tall and up to 90 cm dbh., glabrous in all its parts, older trees
very massive. Bark smooth, grey to light brown, often lenticellate, 0.5 mm thick;
live bark red. Branchlets slender, cylindrical, smooth. Leaves opposite, sub-
coriaceous, obovate-elliptic to sub-obovate-to obovate-oblong, 2.5 x 5— 4 x 7 —
5 x 9 — 8 xX 10 cm, with an obtuse mucro (usually bent down) or obtuse, base
gradually cuneate: above midrib thin, level with the surface or partly sub-
impressed, lateral nerves faint; below midrib thin, prominulous, lateral nerves
erect-patent, 5-6 (rarely up to 7-8) pairs, at some distance from the margin arcuate-
ly anastomosing or the apical ones running out, connected by a loose, very fine,
rather irregular reticulation. Petioles thin, glossy, 1.5-2 cm, flat above. Leaf margin
shallowly remotely serrate.
Panicles sometimes forming loose, compound panicles with some small apical
leaves, axillary, the partial inflorescences rather few-flowered, up to 6 cm long with
long slender peduncles and branchlets which are thicker than those of C. glauca,
repeatedly dichotomous. Pedicels filiform, 10-13 mm. Flowers green (or the petal
white), 5-7 mm in diam. Calyx lobes 5, roundish, concave. Petals 5, 2-2.5 mm long,
oblong, obtuse. Nectary a green pentagonal fleshy, scalloped gland. Stamens five,
filaments short, inserted in the disc, at first erect, later curving down to below the
calyx. Ovary immersed in the disc, 2-celled, each with 2 ovules, attached to the
bottom of the cell. Style very short, conical; stigma simple, obtuse. Drupe almost
round to slightly ellipsoid-globose, up to 18 x 24 mm, shortly sharply pointed,
smooth, green at maturity; mesocarp practically none; putamen with 5 mm thick,
very hard, woody wall with one cavity (by abortion), with two longitudinal lines at
each side, slightly compressed.
Distribution. Endemic to Sri Lanka.
Ecology. Dry, hot zones of North and East Sri Lanka, in coastal areas on sandy
soils; also more inland in savannah vegetation. Can stand a prolonged dry spell.
The putamen splits into two halves after a prolonged period of soaking and rotting.
Vernacular names. Nareloo, Neraloo (Sinhalese), Perun or Piyaree (Tamil).
Note. The Javanese Cassine koordersti Kosterm. (Elaeodendron glaucum, var.
macrocarpum Koord. & Val.) is very close, but has a much smaller inflorescence,
smaller flowers and much shorter petioles; the fruits are the same.
The leaves vary extremely and have minute, scaly stipules at each side of the
internal base of each petiole. In north-east Sri Lanka, on dunes and sterile sandy
coastal areas, we often find this tree as a many-short-boled bushy shrub, which |
have never seen in flower. Its leaves are often large and sharply prickly serrate and
is called in Tamil Kurativa or Kuruku vaichchi. It is this which Roxburgh records as
Ceylon Tea, under which name it was sent from Ceylon to the Calcutta Botanic
Garden by General McDowall.
Chemistry
G. Weeratunga, V. Kumar, M.U.S. Sultanbawa & S. Balasubramaniam re-
Asiatic Cassine 187
ported a 28, 29-dihydroxyfriedelane-3-one, a friedelane with two oxygenated
methyl groups in the bark of Cassine balae. The only friedelane with two oxygen-
ated methyl groups previously reported is salaspermic acid, isolated from Salacia
macrosperma.
The seed of C. glauca contains a cardiac glycoside with double-linked sugar
substitute, whereas the bark had a similar glycoside, five friedelanes and two
norfriedelanes.
A light petroleum extract of the root bark of C. balae was found to contain four
triterpene methides of which three were identified as pristimerin, tingenone and
20-hydroxytingenone on the basis of their physical properties and comparison with
authentic material. Another compound is perhaps 3, 22-dihydroxy-24, 29-nor-D;
A-friede-oleana-2,5,7.9(11), 10(1)-pentaen-2,21 dione 6, a quinone methide.
Moreover they isolated a new triterpene: D. friedo-oleana derivative. According
to G. Weeratunga, L. Bohlin, F. Sandberg and V. Kumar (24th Annual Meeting,
July 24-28, 1983, The American Society of Pharmacognosy, University of Mississip-
pi, Oxford Campus, School of Pharmacy, Abstracts of Papers no. 95) an ethylace-
tate extract of the rootbark contained 3,3’, 5,5°, 7-pentahydroxy 4-methoxy, 2,3-
cis-flavane as the major constituent; furthermore a novel leucoanthocyanidin was
isolated.
Specimens examined.
Talaimannar, Mannar Distr., Northern Prov., sandy scrubland, Nov., buds, Kundu et al. 623 (PDA):
Trincomalee Distr., Upuvelu along border of Blue lagoon, rooted in brackish water, March buds
Cramer 4938 (PDA): Trincomalee, Jan., fl., Worthington 742 (K): Wilpattu Nat. Park, July, fl.,
Wirawan et al. 955 (BO. L. PDA): Batticaloa Distr., Kalkuda Resthouse,. June, fr.. Waas 639 (PDA);
ibid., June, y. fr., Worthington 6295 (K): Polonaruwa Ruin area, May, buds, Kostermans 24309 (L,
PDA); ibid., Oct. fl., Ripley 222 (PDA); ibid., July, after anthes., Ripley 67 (PDA); ibid., Apr.. fl..
Hladik 788 (PDA): along road to Gal Oya Nat. Park Inginiyagala, Amparai Distr., ster., Meyer et al.
162 (PDA); Ruhuna Nat. Park, mouth of Kumbukkan Oya, beach, May fl., Jayasuriya 2016 (AAU.
PDA); Tissa-maharana, Kataragama road, Sept., y. fr., Coorey s.n. (PDA): Yala Bungalow. Ruhuna
Park, Oct., fl. Wirawan 681 (PDA): ibid., Oct... fl.. fr.. Nooteboom 332 (AAU, L, PDA); Buttuwa
Beach area, June, fl.. Comanor 899 and Apr., fl., Fosberg 50333 (AAU, L. PDA); ibid., ster.,
Worthington 5681 (K): Badulla Distr.. Jamburagala, semi savannah, sandy, Jan., fl., Fosberg er al.
51612 (PDA): Divulane forest. Ampara Distr., May, fl.. Jayasuriya 2085 (AAU): Wellawaya. Aug.. fl..
Worthington 2992 & 4903 (K); mile 187 near Wellawaya. Moneragala Distr., June. buds, fr.. Meyer 199
(L, PDA): Habantotta, along the coast, Sept.. fr.. Balasubramaniam 2213 (BO. K, L): Amaduwa, fl.
1853, inter C.P. 1227 and 2520 (PDA).
Cassine congylos Kosterm.. sp. nov.
Elaeodendron glaucum, var. montanum Thwaites, Enum. PI. Zeyl. (1858) 73: Lawson in Hooker f., Fl.
Brit. Ind. 1(1875) 623: Trimen, Handb. FI. Ceylon 1(1893) 272. — Typus: C.P. 2520 (PDA).
Cassine glauca, var. montana (Thw.) Pierre, Fl. for. Cochinch. 4(1893). t. 296.
Arbor magna in omnibus partibus glabra. cortice subtus ochraceis, foliis rigide coriaceis oppositis late
ellipticis vel subobovato-ellipticis, apice rotundatis vel obscure mucronatis, basi sensim attenuatis acutis
vel obtusis, margine obscure serratis, supra nervo mediano vix prominulo, nervis lateralibus tenuibus
prominulis, subtus nervo mediano prominentibus, nervis lateralibus suberecto-patentibus vix prominulis
filiformibus, rete obscure laxis. petiolis gracilibus supra concavis, sat brevibus, paniculis axilaribus
paucifloris, pedicellis filiformibus sat longis, lobis calycinum depresso-rotundatis, petalis viridis oblongis
sat longis. discus magnis incrassatis, filamentis brevibus sat latis reflexis: stylo conico brevibus; fructus
subglobosis viridis, mesocarpium deest, putamen crassum apiculatum, globosum. — Typus: Kostermans
27791 (L).
Tree, up to 30 m tall and 100 cm dbh. Bark brownish to yellowish brown, dark,
roughish, pustular, 1 mm thick, underneath orange, inflammable (burns like
kerosene); live bark red, 10 mm thick. All parts glabrous. Leaves opposite (rarely
188 Gard. Bull. Sing. 39(2)(1986)
sub-opposite), rigidly coriaceous, broadly elliptic to sub-obovate-elliptic, 2.5 x 3.5
— 4 xX 6cm, apex rounded or mucronate, towards the base slightly tapered, the
base acute or obtuse; margin rather obscurely, remotely serrate; above midrib
hardly prominent, lateral nerves thin, prominulous, below paler, midrib thin,
prominent, lateral nerves erect-patent to more patent, thin, prominulous, 5-7 pairs,
in between a rather obscure, lax reticulation. Petiole slender, concave above, 5-15
mm.
Panicles axillary, rather few-flowered, up to 7 cm long with few, remote, slender
branchlets, the 1-mm long, narrow, acute bracts persistent. Pedicels slender, 3 mm.
Calyx lobes depressed-orbicular, 2.5 mm diam. Petals green, oblong, the margins
lighter green, ca. 5 mm long. Disc large, thick, up to 3 mm in diam., slightly
incised. Filaments rather broad, 1-1.5 mm long, in the incisions of the disk,
reflexed; style short, conical; stigma inconspicuous.
Fruit sub-globose, green at maturity, 2-3 cm diam., smooth, with short sharp
point, one cavity (by abortion of the other); mesocarp practically none; putamen
very hard, very thick, sharply apiculate, base obtuse, superficially longitudinally
wavily grooved with appressed tree-like fibres.
Distribution. Endemic to Sri Lanka, Knuckles Massive.
Ecology. Wet, evergreen forest but with a yearly dry spell of 2-3 months, at c.
1000 m altitude.
‘Note. Trimen commented, that, although the flowers were larger, the fruit larger
and the petioles shorter, it did not merit specific rank. There are even more
differentiating characters, like the rigidly coriaceous leaves and the orange under
side of the dead bark.
Lawson erroneously quotes it as coming from the drier parts of Ceylon; it is a wet
zone plant.
Specimens examined.
Deltota, Dec. 1855, fl., C.P. 2520 (BO, PDA); Dimboola, 5000 ft., Apr. 1852, fl., C.P. 2520 (PDA);
Knuckles Mts., Madulkelle area, Hemachandra’ cardamon estate, alt. 1000 m, June, fl., Kostermans
25070 (AAU, BO, L, P, PDA, US); ibid., base Dusingalle, 1000 m alt., Aug., fl., Kostermans 27791
(AAU, G, K, L); ibid., Dec., fallen fr., Jayasuriya 2550 (AARH, G, K, L).
MALESIAN SPECIES
Cassine elliptica (Decsne.) O.K. Fig. 4
O. Kuntze, Rev. Gen. Pl. (1891) 114. — Elaeodendron ellipticum Decaisne in Nouv. Arch. Mus. Hist.
nat. Paris, Ser. II, 3(1834) 478 (separately paged reprint p. 150); Ding Hou in FI. Males., Ser. I,
6(1962) 286 (as a syn. of Cassine glauca, var. cochinchinensis Pierre). — Type: Riedlé s.n. (P).
The type specimen was collected in the island of Timor, East Indonesia, and is a
flowering twig. Miquel (FI. Ind. bat. 1(2) (1859) 591) mentioned a second speci-
men, but this could not be found.
The leaves of Riedlé’s specimen differ considerably from those of the Javanese
C. koordersii and the Moluccan C. obiensis; the base tapers gradually in the almost
absent petiole. The inflorescence, extra-axillary, is a few-flowered, very shortly and
thinly branched cyme on a thin, long common peduncle, also different from that of
C. koordersii.
The leaves resemble strongly those of C. australe (Vent.) O.K. from Australia,
but this should have fruit with a juicy thick mesocarp and the flowers 4-merous.
Cassine koordersii Kosterm., sp. nov.
Elaeodendron glaucum Auct. (non O.K.), var. macrocarpum Koorders & Valeton, Bijdr. Kennis
Ay t« mer wf
Fig. 4 Cassine elliptica (Decsne.) Kosterm., holo (P).
1&9
190 Gard. Bull. Sing. 39(2)(1986)
Booms. Java 7(1900) 101-102. — Typus: Koorders 30086 (BO), ripe fruit.
Cassine glauca, var. cochincinensis Auct. (non Pierre) Ding Hou in Fl. Males., Ser. 1, 6(1962) 286, fig.
18, f. 9 (exclud. Elaeodendron ellipticum Decsne.); Auct. (non Pierre) Backer & Bakh., Fl. Java
2(1965) 55.
Arbor in omnibus partibus glabra, follis oppositis late ellipticis vel subobovato-ellipticis margine
crenulato-serratis, petiolis brevibus, inflorescentiis axillaribus paucifloris pedunculis gracilibus, fructus
late ellipsoideus vel subglobosus apiculatus, mesocarpium nullus, putamen crassum durum.
Tree up to 30 m tall, up to 95 cm dbh., all its parts glabrous. Bark smooth, grey,
pustular. Leaves opposite, thinly coriaceous, broadly elliptic to subobovate elliptic,
very shallowly crenate-serrate or entire, 4.5-15 x 2.5-6 cm, base rounded or acute,
obscurely very shortly acuminate (acumen turned side-ways), both surfaces dense-
ly, minutely reticulate, lateral nerves c. 6 pairs. Petiole slender, short, 1-1.5 cm.
Inflorescences axillary, very short, few-flowered, 1-4 cm long on 1-2 cm long
slender peduncle, cyme apical. Pedicel 2-4(-5) mm. Calyx lobes broadly rounded.
Flowers green. Petals oblong, obtuse, 2.5-4 mm. Disc green. Filaments 1.5-2 mm,
after anthesis strongly reflexed.
Fruit broadly subovate-ellipsoid to globose, 2-2.5 x 1.5-2 cm, very shortly,
sharply apiculate, base rounded; exocarp rather tough, mesocarp hardly repre-
sented; endocarp 3-7 mm thick, hard, woody or bony, obtuse, one-seeded by
abortion of the other.
Distribution. Only known from the Poeger (Puger) area in the Besuki Residency
in East Java in the valley of Lampesan and on the Watangan Mts, not rare on
periodically very dry soil of weathered coral limestone in teak and other dry forest
types.
The numerous collections brought together by Koorders and his assistants are
mostly sterile. Only a single specimen had very small, few-flowered cymes and
another had a few ripe fruits, which were globose.
The species is characterized by the very short petioles and inflorescences. In fruit
characters it is exactly like the Moluccan C. obiensis, with which it might be found
eventually to be.
Cassine obiensis Kosterm., sp. nov.
Arbor magna in omnibus partibus glabris, foliis oppositis chartaceis ellipticis subobtusis serratis, petiolis
tenuibus supra concavis, fructus ovoideus laevibus mucronulatus, mesocarpium sub-deest, putamen
percrassum lignosum, semen unicum. — Typus: de Vogel 4173 (BO).
Tree, glabrous in all its parts, 40 m tall, bole 20 m, dbh. 40 cm, straight;
buttresses 2 m high, out 1 m, thick 8 cm. Dead bark 2 mm thick, pale brownish, not
fissured, not peeling off; live bark 6 mm, reddish ochre. Twigs rather slender,
smooth, cylindrical, finely longitudinally ribbed. Leaves opposite, chartaceous,
elliptic, 4 x 8 — 4.5 x 10cm, obtuse or obscurely very broadly obtusely acuminate,
base shortly acute; above smooth, midrib slightly prominulous, lateral nerves faint;
below midrib slender, prominent, lateral nerves up to 12 pairs, erect-patent, at
some distance from the margin arcuately anastomosing, reticulation lax, faint.
Petioles slender, 10-15 mm long, concave above.
Fruit ovoid, smooth, green (fresh), apiculate and with two opposite, shallow,
longitudinal grooves; exocarp 0.5 mm thick, mesocarp 0.5 mm thick, putamen very
hard, woody, 4-10 mm thick with one assymmetric (by abortion) cavity; seed one,
flat, 1.5 cm wide.
Distribution. Indonesia, Motuccas.
Asiatic Cassine 191
Note. Related to Cassine koordersii Kosterm. from Java. but the fruit ovoid. much
larger and the putamen with a twice as thick wall.
The type sheet consists of a leafy branch and some detached fruit.
Additional material.
INDONESIA: North Moluccas, Obi Isl., Anggai, Mt. Batu Putih, 1° 24’ S, 48’ E, alt. 600 m, hillside with
many limestone outcrops and boulders, covered with clayer soil, rather dense primary forest, 40 m tall
with rather little undergrowth, rare tree, 19 Nov., fr., de Vogel 4173 (BO, L): S. and E. Moluccas:
Tanimbar Isl., Ilgney, Ottimer, ster., bb. 24295 (BO); March, fr., bb. 24207 (BO, L): Key Isl., fr..
Jaheri 167 and 382 (BO).
Acknowledgements
It took considerable time to distangle the misinterpretations of Koenig’s mate-
rial, on which Cassine glauca and C. albens are based and without the much
appreciated assistance of Dr Bertel Hansen (Copenhagen herbarium) we would
certainly not have succeeded. We like to express here also our gratitude to the
Curator of the Lund Herbarium (Sweden) for the loan of the type specimen of
Schrebera albens Retz. and to the Directors of the Leiden and Aarhus Herbaria, to
permit us to examine the specimens in their Institutes.
Last, not least, we thank Dr Ding Hou of the Rijksherbarium, Leiden, who
extended to us his generous assistance.
The Genus Pandanus (Pandanaceae) On Christmas Island,
Indian Ocean
BENJAMIN C. STONE
Department of Botany, Academy of Natural Sciences, Philadelphia
Pennsylvania, U.S.A. 19103
EFFECTIVE PUBLICATION DATE: 24 JAN. 1987
Abstract
Two species of Pandanus occur on the remote Christmas Island (Indian Ocean), south of Java (105°
40° E, 10° 30°S), although three binominals have been published. The correct names for the species, a
key for their identification, synonymy, and descriptive notes, with illustrations of new or distinctive
features, are here presented. Pandanus christmatensis is shown to be a member of subg. Pandanus, sect.
Pandanus, closely related to P. tectorius Park., and especially so to P. platycarpus Warb. P. elatus is
shown not to be a member of sect. Pandanus, as previously stated, but rather, in sect. Roussinia (of
subg. Rykia), previously considered monotypic with P. leram Jones ex Fontana. Several critical
features, including the hair-tufted anther apiculi, shared by P. leram and P. elatus, reveal their close
relationship.
Introduction
According to Murray’s Introduction in Andrews’ Monograph of Christmas Island
(1900), Christmas Island has been known to navigators since about the middle of
the seventeenth century. Java, the nearest land mass, is about 190 miles to the
north. To the south-west, at a distance of about 550 miles, are the low atolls, the
Cocos-Keeling Islands. Christmas Island itself is an ancient, uplifted limestone
summit. Its geology is somewhat complex, and basalts and phosphate-bearing rocks
are also present. It is the latter which account for the past interest and current
industry on the island, phosphate being now the export product. The Island lies in
the same latitude, south of the equator, as Fatuhiva in the Marquesas Islands, San
Cristobal Island in the Solomon Islands, Cape Delgado on the coast of Mozambi-
que, and is only a short distance south of the latitude of Luanda in Angola and
Recife in Brazil. Accordingly it has a fully tropical climate. Christmas Island was
shown on a map prepared by Pieter Goos, published in Holland in 1666 (on which it
is called Moni). It is not certain who applied the name by which it is presently
known. The Island is mentioned by Dampier who visited it in March 1688. Subse-
quently, it was touched on by several voyagers, but the first real exploration was
attempted in 1857 by the frigate H.M.S. Amethyst. In 1886 the survey ship Flying
Fish under Captain Maclear examined it and some specimens of plants and animals
were brought home, the first to represent its flora and fauna. Captain Pelham
Aldrich called at it in H.M.S. Egeria in 1887, and some natural history collections
were made by J.J. Lister. In 1888 it was placed under the Government of the Straits
Settlements by the visit of H.M.S. Impérieuse. In 1890, H.M.S. Redpole stopped
there for a few hours, allowing H.N. Ridley, of the Botanic Gardens, Singapore, to
make some plant collections (Ridley, 1891).
The principal collections from Christmas Island date from the residential work of
Charles W. Andrews, who arrived there in July 1897 and remained for nearly a
Received on 24 Feb. 1986.
Ss
Jackson Pt.
Martin Pt.
Wright Pt.
Jacks Pt. ig
W inj fred Bea, Deans Pt.
Toms Pt. oe
30'— Greta Beach — 10 30
Egeria
Pt.
John D Pt.
Dolly Beach
SMITHSON B/GAT McMicken Pt.
Tait PY,
INDIAN OCEAN
Andrews Ptl. sa ‘dec
a7 ok pies
roe: iu) }
tubbings . mt # ~ 7 Medwin Pt.
Seuth Pt. 1x40"
Fig. 1. Christmas Island. from the original which was based on Atlas of Australia, Map 159, Reader’s
Digest Services Pty Ltd. Sydney.
year. In October 1904, Ridley collected there again, later publishing some new
species (Ridley, 1906), including two binomials in Pandanus discussed further
ahead.
The map view of the Island (Fig. 1) may be likened to a standing dog, facing east;
its tail is North West Point; its hind leg is Egeria point; its front leg is South Point;
and the larger north-eastern part includes the Settlement; the ears are Rocky Point
and North East Point, while the muzzle is Norris Point and Low Point, with
Waterfall in between. The greatest distance east to west is about 17 km, and north
to south about 17.5 km. The highest points are Murray Hill, in the western part
(361 m) with Jack’s Hill near it (331 m); and Hanitch Hill, in the north-eastern part
(309 m). At a distance of 1-3 km inland from the coast, abrupt steep cliffs rise which
border the interior plateau, which slopes gently to the south and west. Its surface
has shallow valleys and low ridges. The margin is often marked by limestone
pinnacles.
The soil is mostly (apart from the reefs and pinnacles) a brown loam strewn with
phosphate nodules and often also with fragments of basalt.
The collections of Andrews resulted in the publication of A Monograph of
Christmas Island (Andrews, 1900) and this embodied the studies of several botan-
ists and zoologists, mostly of the British Museum (Natural History) staff. In the
Monograph, the enumerated flora contained 111 species of Dicotyledons, 18 of
194
Pandanus on Chrisunas Island 195
Monocotyledons, 22 of Pteridophytes, 8 of Bryophytes, and 31 other lower crypto-
gamous plants. mostly Fungi, as well as one Gymnosperm (Cycas circinalis).
The Genus Pandanus in Christmas Island
Andrews collected staminate flowers of one Pandanus species during his visit in
1897. This was subsequently described by Martelli (1905), who named it Pandanus
christmatensis. In 1904 Ridley obtained fruiting collections representing two species
which he himself named as new. Of one he also obtained good staminate material.
The Christmas Island pandans were reviewed by St. John (1965). He accepted
two species, for which he used the names applied by Ridley, providing full descrip-
tions (and good illustrations) of Pandanus elatus Ridl. (fruit and staminate flow-
ers), and of P. nativitatis Ridl. (fruiting specimens), based on Ridley’s collections in
the Herbarium of the Botanic Gardens, Singapore. Although St. John mentions
briefly that Martelli had previously described (to Ridley) a Pandanus species from
the island, he neither reports its name nor attempts to deal with it taxonomically
and nomenclaturally, remarking merely that the specimen was “fragmentary, sta-
minate material.” This material (1.e., Andrews’ collection) was evidently not
further studied.
Realizing that further investigation of the Christmas Island pandans was neces-
sary in order to resolve the problem of how many species were in fact indigenous
there, I requested full new collections from the Conservation Officer, Mr. D.A.
Powell, who responded most helpfully. From him I later received excellent collec-
tions and notes, which are cited below and which confirm that there are two
indigenous species on the island. The collections made by Mr. Powell and his
colleague, Mr. H’ng Kim Chey, represent fruiting and staminate flowering material
of both species. With this complete representation, it is now clear that one of the
names proposed by Ridley, P. nativitatis, must be regarded as a synonym of P.
christmatensis Martelli. This is the species of the coastal regions. The second.
limited to the central plateau, retains the original name proposed for it by Ridley.
P. elatus.
Key to species
Tree to 20 m tall; pistillate cephalium, when ripe, somewhat oblong-cylindric-ellipsoid, to about 33 cm
long, bearing about 100 phalanges; phalanges 7.5-9 cm long, more or less transversely compressed,
apex low convex; carpels 4-12, mostly 6-9 per phalange; stigmas large, erect, 3-4 mm long and wide.
Staminate spikes 12-20 cm long; staminate phalanges 12-18 mm long, of 15-23 stamens clustered at
apex of the long (to 9 mm) column; anthers 3-4 mm long, apiculus c. 0.5 mm, acute, bearing a minute
tuft of glandular hairs at the apex. Staminate bracts to 40-50 cm long, apex acute, closely spinulose-
I ee ee eae eS ie a go ae hohe a tdaee een soawE A Abas doch wees paepetes sb eeeees Pandanus elatus
Shrub or small tree to c. 5 m tall; pistillate cephalium, when ripe, ellipsoid or ovoid-ellipsoid, to about 25
cm long, bearing about 60 phalanges; phalanges 5.5-8 cm long, plump, not compressed, apex rounded
to convex; carpels many, usually 15-26 (or sometimes fewer, to 9); stigmas about 1-1.5 mm long and
wide. Staminate spikes mostly 7-9 cm long; staminate phalanges fastigiate, 15-20 mm long, stamens
racemose along the column, the free basal part of the column at most 5 mm long; anthers c. 5 mm
long, apiculus 0.5 mm long, acute, glabrous. Staminate bracts to 65 cm long, but the apical part very
narrow attenuate and flagellate, at apex with very remotely spinulose margins. ....... Pandanus christ-
matensis
Subg. Rykia Section Roussinia
Pandanus elatus Ridley Plate 1. Figs. 2,3. &7
Ridley. J. Str. Br. Roy. Asiat. Soc. 45 (1906) 239: St. John, Pacif. Sci. 19 (1965) 113, figs. 215-216.
St. John has provided full descriptions and illustrations of both the pistillate and
staminate plants, to which the recently collected specimens conform fully, while
providing the basis for some corrections and emendations.
Plate 1. Staminate flowers and inflorescences of Pandanus elatus (a, b) and P. christmatensis (c, d).
a, Part of a staminate inflorescence from Powell & H’ng 10 Oct. 1983; b, staminate phalanges
from the same collection. c, Staminate inflorescence from Powell & H’ng Nov. 1983; d,
staminate phalanges from the same collection. Scale in b & d show a 2-cm bar with | cm
divided into mm.
The acute rather than flagelliform apices of the bracts, which moreover have
rather densely spinulose margins, are distinctive features that help to distinguish
plants of P. elatus from those of P. christmatensis.
More significant, and not mentioned in St. John’s description, are the hair-tufts
of the apiculi of the anthers of P. elatus. These are formed of glandular hairs and
are Observed on almost all the stamens. The staminate phalanges bear generally
12-18 (or up to 23) stamens subumbellately arranged on the column apex, which
contains usually 4 or 5 vascular bundles in a single circle, and lacks crystal cells, as
seen in the cross-sectional view (fig. 3B). The papillosity referred to by St. John
said to occur on the column and filaments is an artifact; these structures are in fact
smooth, and the so-called papillae are probably merely adherent pollen grains. The
pollen grains are about 23 uw long, slightly larger than those of P. christmatensis, and
the spinules are somewhat more numerous and longer.
The very large stigmas are noteworthy, as well as the usually obvious transverse
seriation of the carpels and resulting compression of the pistillate phalanges. The
196
mm
Fig. 2. Pandanus elatus, details of staminate structure. Left to right: a
complete phalange; pollen; anther, dorsal face; anther, ventral
face; anther tip showing tuft on apiculus.
0,01 mm 0,01 mm
Fig. 3. Pandanus elatus, details of staminate structure. a, tip of apiculus, showing the papilliform
glandular hairs and one pollen grain. b, Transection at midlevel of one 18-stamened phalange,
showing 4 vascular bundles; note absence of crystal cells. Drawn by K.L. Huynh.
197
198 Gard. Bull. Sing. 39(2) (1986)
outermost carpels appear sterile, their loculi being smaller and lacking endosperm.
Often there are only two or three loculi with normal endosperm. The phalanges not
infrequently have adherent, imperfect, small peripheral carpels or ferulae (as is
true also of P. christmatensis).
LectotyPe. H.N. Ridley in October 1904, Christmas Island, plateau (SING).
Isolectotype in K. Staminate paratype: H.N. Ridley in October 1904, Murray Hill
track, Christmas Island (SING).
RELATIONSHIPS. Pandanus elatus was placed in subg. Pandanus sect. Pandanus by
St. John (1965), clearly on the basis of the pistillate phalanges only. The species was
overlooked by Martelli who omitted it in his Enumeration of species in Webbia 4, 1
(1913).
Pandanus elatus differs from authentic members of section Pandanus, such as P.
tectorius, P. odoratissimus, and P. christmatensis, in several respects, including the
compression of the pistillate phalanges, subumbellate arrangement of stamens,
arrangement of the vascular bundles in the column, and the papillae or hair-tufts of
the anther apiculi. These features exclude P. elatus from section Pandanus. On the
other hand, three of the features are to be found in Pandanus leram Jones ex
Fontana, a species of the Andaman and Nicobar Islands with an occurrence in
South Java (Nusa Kambangan Island, off Tjilatjap), and of the Maldive Islands.
This species is cultivated in Sri Lanka and elsewhere (Stone, 1977). In P. leram, the
compression of the pistillate phalanges, with more or less clear transverse seriation
of the carpels, the large stigmas, and similar leaves and branching habit are to be
found, as well as the staminal arrangement and apicular hair-tufts of P. elatus.
0.imm
0,026 mm b
q
Fig. 4. Pandanus leram, details of staminate structure. a, Anther
apiculus, showing tuft of hairs at tip, with 2 pollen grains; b,
enlarged view of tuft hairs and pollen grains. From Stone
11102 (KLU), Sri Lanka.
Pandanus on Christmas Island 199
However, the column contains somewhat more numerous vascular bundles which,
moreover, are usually in two circles not one. The pollen grains of P. leram are
slightly larger than those of P. elatus (c. 26 w rather than c. 23 yp).
Pandanus leram is the sole member of sect. Roussinia (Gaudich.) Stone (Stone,
1983). The perceptible similarity of P. elatus based on vegetative, foliar, fruit, and
staminate characters, indicates that P. elatus must be assigned to section Roussinia
as a second member. On geographical grounds also, this relationship is plausible.
The hair-tufts of the stamens are not restricted to this section; they also occur in a
number of other species in several different sections, including sections Asterodon-
tia (subg. Rykia), Barrotia and Brongniartia (subg. Lophostigma).
Pandanus elatus is endemic to Christmas Island, limited to the high limestone
plateau. It is most similar to P. leram var. andamanensium (Kurz) Stone.
Subg. Pandanus Section Pandanus
Pandanus christmatensis Martelli Plate 1. Figs. 5-7
P. nativitatis Ridley, J. Straits Branch, Roy. Asiat. Soc. 45 (1906) 238. St. John, Pacif. Sci. 19 (1965)
126: ft. 2EF:
To the description provided by St. John can be added the following emendations
and alterations.
The pistillate cephalia are ovoid-ellipsoid, up to about 22 x 21 cm (when dried),
on peduncles at most 2 cm diameter. There are about 55-70 phalanges per cepha-
lium. The phalanges are distinctly polycarpellate, with about 18-21 carpels being
the usual number per phalange, though those with both more numerous (to 26) and
fewer (to 9) can be found. St. John’s description was based on only three pha-
langes. In the recent collections the phalanges are about the same size or slightly
larger, the apical surface is usually somewhat more convex, but in other respects
there is good agreement.
The staminate inflorescences are typical for species of sect. Pandanus, the lower
and transitional floral bracts being distally protracted into flagelliform tips; these
4mm
|
Fig. 5. Pandanus christmatensis, details of staminate structure. Left
to right: one complete phalange; pollen grain; anther, dorsal
face; anther, ventral face; tip of apiculus. Note distribution
of raphide cells (small rectangles).
200 Gard. Bull. Sing. 39(2) (1986)
Fig. 6. Pandanus christmatensis. Transection at midlevel of column of 31-stamened phalange, with 25
vascular bundles. Note occurrence of crystal cells (small squares) just below epidermis. Drawn
by K.L. Huynh.
have sparse and remote marginal spinules. The spikes are comparatively short, up
to about 10 cm long. The phalanges are dense, about 15-20 mm long, composed of
15-32 stamens, racemosely arranged along the column, only the base (4-5 mm) of
the column free; the anthers are about 5 mm long, with a distinct acute apiculus c.
0.5 mm long, the tip of which is smooth and glabrous.
The pollen grains are about 19 yu long, slightly smaller than those of P. elatus,
with fewer and somewhat shorter spinules.
The column contains several vascular bundles (fewer than the number of sta-
mens), which are arranged in 2 or 3 circles, as is typical in sect. Pandanus. Several
crystal-cells are to be seen beneath the epidermis (fig. 6) in cross-sectional view.
Each crystalliferous cell contains a single crystal.
Ho ortype. Christmas Island, C. W. Andrews in 1897, staminate flowers (BM).
Original citation: HAB. Christmas Island nell’Oceano Indiano (Andrews in Herb.
British Museum); Martelli, 1905, 1.c. No other specimens cited. Type of P.
nativitatis Ridley: H.N. Ridley in October 1904, Waterfall, Christmas Island (Indi-
an Ocean), in SING.
The relationships of P. christmatensis are clearly with the core species of subg.
Pandanus section Pandanus, including the type species, P. tectorius Park. ex Z.;
but more particularly with a taxon named P. platycarpus Warb., said to be from
Zanzibar (though this was disputed by Martelli, who considered it to be from
Java)*, the type specimen, collected by E.H.L. Krause, was illustrated by Martelli
(1913) in t. VII; and what is possibly the same taxon is shown in a photograph
captioned ‘‘P. tectorius Sol.”’ as figure 6 in van Steenis’s Flora Malesiana sample
treatment, ser. I, vol. 2, Pandanus in Malaysian vegetation types of 1954. Also quite
similar is the taxon named P. intraconicus St. John from Aldabra Island (see the
illustration figure 334 in Pacif. Sci. 18 (1974) 96-97). This in turn matches pretty
well with a collection from Cocos-Keeling Atoll (with an unpublished specific
name) made by H. St. John, no. 26414, in the Bishop Museum, Honolulu. The
phalanges of this latter collection match so well with those of P. platycarpus that I
* Martelli published his view that P. platycarpus was not a native of Zanzibar in the Atti Soc. Toscana
Sci. Nat. 42: 57-59, 1933. He apparently compared Krause’s specimen to material from Java and
assumed that the plants seen by Krause were not native (either introduced or spontaneous but
presumably ephemeral). This is possible, but since closely related species such as P. kirkii are clearly
indigenous along the East African coast, the actual occurrence of P. platycarpus in Zanzibar does not
appear in doubt.
Pandanus on Christmas Island 201
would not hesitate to regard them as of the same species; phalanges of P. intraconi-
cus differ in the somewhat more prominent carpel tips; while in P. christmatensis,
the phalange apex is more rounded, the carpel tips still more prominent, the sides
of the phalange are not scarred so much, and the lateral sutures are more obvious.
These are nonetheless all quite similar, and it is noteworthy that all occur in a
latitudinal belt only some 5° wide but about 60° in longitudinal extent (or a little
more if Zanzibar is included), across the middle of the Indian Ocean. Of interest
too is the record of “P. tectorius” from Rodriguez Island, illustrated by Martelli
(1913) in tab. 2, fig. 6), which although not the same as the species mentioned
above, appears approximately in the same drift zone (but about 5° farther south).
M7 N Z-
(A>,
7/\ S.
)
C A
SS = ANN N : eee
Fig. 7. Habit silhouettes of 3 Pandanus species: a, P. elatus, juvenile and adult; P. leram, modified
from Gaudichaud’s illustration of Roussinia indica. c, P. christmatensis. a & c from photo-
graphs by D.A. Powell.
Differences in Habit (Fig. 7)
Habit and branching pattern in Pandanus elatus and P. christmatensis differ. In
the former the- young plant remains unbranched for some considerable time, and
may reach a height of as much as 12 or 13 m before branching, depending on the
canopy height of the surrounding vegetation. Branching is somewhat sparse, usual-
iy a trichotomy, and the branches themselves di- or tri-chotomously branched in
due course. The main trunk remains quite erect, and the proproots tend to form a
fairly compact inverted cone 1-1.5 m high. In habit, P. e/atus further reveals its
affinities, this pattern of growth being found not only in P. /eram (see the habit
sketch in the original illustration of Roussinia indica Gaudich., Bot. Voy. Bonite
P1. 21, f. 1, 1843), but also in other species in subg. Rykia (e.g. P furcatus Roxb.,
P. houllettei Carr., P. lais Kurz, etc.).
In contrast, the habit of P. christmatensis is, as in its relatives, more fruticose, the
trunks branching at a comparatively early stage, with the branches tending to be
more spreading, more irregular, the trunk continuing but not so straight, and the
proproots often supporting the leaning or even partly horizontal base of the trunk,
themselves often branching and interwoven. The overall appearance is thus very
202 | Gard. Bull. Sing. 39(2) (1986)
much like that found in P. tectorius, P. odoratissimus, P. kirkii, and other species of
sect. Pandanus.
These habit characters, supplemented by the habitat differences, thus permit not
only the recognition of the species, but lend furtheir support to the allocation of P.
elatus to sect. Roussinia of subg. Rykia, and of P. christmatensis to sect. Pandanus
of subg. Pandanus.
Acknowledgements
Examination of the anatomy of the staminal columns in flowers of Pandanus
elatus and P. christmatensis was undertaken by Dr K.L. Huynh (Institute Botani-
que, Universit de Neuchatel, Switzerland), who also prepared figures 4b and 6, and
suggested several valuable emendations to the text. This cooperation is gratefully
acknowledged.
Much of the original stimulus in studying the species reported on here is the
result of the letters, photographs, and collections of D.A. Powell and H’ng Kim
Chey. Additional photographs by them will be found associated with the herbarium
sheets in K, KLU, and PH.
References
Andrews, C.W. (1900). A Monograph of Christmas Island (Indian Ocean): Physic-
al Features and Geology by Charles W. Andrews, B.A., B.Sc., F.G.S. With
Descriptions of the fauna and flora by numerous contributors. London: Trustees
of the British Museum (Natural History).
Martelli, U. (1905). Pandanus — nuova specie descritte. Webbia, 1: 361-371.
Ridley, H.N. (1891). A day at Christmas Island. Journal of the Straits Branch,
Royal Asiatic Society, 23: 123-140.
Ridley, H.N. (1906). An Expedition to Christmas Island. Journal of the Straits
Branch, Royal Asiatic Society, 45: 137-271.
St. John, H. (1965). Revision of the genus Pandanus Stickman, Part 18. Pandanus
of Christmas Island, Indian Ocean, and of the Anamba Islands, Indonesia.
Pacific Science, 19: 113-119.
Stone, B.C. (1977). On the identity of the Nicobar Breadfruit and a record of
Pandanus leram fruit in beach drift in Pulau Langkawi, Kedah. The Malayan
Nature Journal, 30: 93-102.
—______. (1983). Revisio Pandanacearum, Part Il. Pandanus subgenus Rykia
(DeVr.) Kurz (sections Gressittia, Rykiopsis, and Solmsia excepted). Federation
Museums Journal, new series, 28: 1-100. (Sect. Roussinia, cfr. pp. 91-93).
Warburg, O. (1900). Pandanaceae. In A. Engler (ed.), Das Pflanzenreich, Hett 3,
IV. 9: 1-99.
The Use of Tifgreen and Tifdwarf Bermuda Grasses in two Singapore
Golf Courses
WONG YeEw Kwan*
Abstract
The two Bermuda grasses were introduced into Singapore in 1981 for use in the Serapong Golf Course
of Sentosa and Tanah Merah Golf Course near the Changi Airport. The rooting medium for both
courses is sand and the turf was established by sprig sowing.
Tifgreen was used for the fairways and Tifdwarf for the greens. Cultural practices based on recom-
mendations of the consultants to the two golf courses are described. Observed pests and diseases are
mentioned. It was the intention of the two Clubs to keep the introduced turf as pure cultures in the
courses. Eventually it was decided that only the greens would be maintained as such as it was found too
costly to do extensive weeding in Singapore where labour is relatively expensive and the invasion by
local grasses and sedges overwhelming. Some of these are briefly described and/or featured in the
various photographs.
All in all it is reckoned that because of the ease of propagation and their fast growth, these Bermuda
grasses do constitute an easy source of turf to green up a large area within a short time, although initial
weeding is essential for proper establishment if sprig sowing is done.
The two Tif grasses have been developed by the Coastal Plain Experiment
Station at Tifton, Georgia, U.S.A. The sprigs for turfing up the Serapong Course at
Sentosa Island and the course at Tanah Merah Country Club (TMCC) were
introduced into Singapore through the Jagorawi Golf Course near Bogor, Indone-
sia, in February 1981. Together with these two grasses, Santa Ana (another
Bermuda variety developed in California) was also brought in. The intention was to
use Santa Ana for the tee stations, Tifgreen for fairways and Tifdwarf for the
greens. Indeed the two courses were so planted. For the introduction one plastic
bagfull (about 0.03 cu m or 1.0 cu ft) of fresh stolons of each hybrid was brought in
and these were planted in the Sentosa Development Corporation nursery for
further propagation.
The Bermuda grasses belong to the genus called Cynodon. According to Beard
(1973) they have all originated from East Africa. Repeated interspecific crosses
between Cynodon dactylon, C. transvaalensis, Cynodon X magennisti, and C.
incompletus var. hirsutus have given rise to many so-called warm season grasses,
including Tifgreen and Tifdwarf. Such varieties are used for turfed areas in the
warmer regions of the United States. C. dactylon, incidentally, is a common grass
in Singapore, found in open fields and wasteland. It is a rather coarse grass with a
bluish tinge. This texture contrasts very sharply with Tifgreen and Tifdwarf, which
are more like the fine varieties of Zoysia and Digitaria, commonly used for putting
greens in this part of the world. Between Tifgreen and Tifdwarf, the latter tends to
have shorter internodal lengths and a much higher shoot density, hence its choice
for the putting greens. Some stolons and culms of the two Tif Grasses are shown in
Plates 1 and 2. Note that both grasses show inflorescences. Indeed, if allowed to
grow at long mowing intervals, the two grasses flower freely in Singapore all the
year around. It is, however, not known whether they set seeds or if they are sterile.
“Mr Wong is a founder member of TMCC and was at various times serving on its management
committee & the Sentosa Golf Board.
203
ee eee ECOL OLE OMA
Ss OOH ‘
Qn
&
WZ
OH WQWs
\
Tifgreen Bermuda Grass, stolons, culms, and inflorescence.
Plate 1’,
Tifdwarf Bermuda Grass, stolons, culms, and inflorescence.
ate 2.
PI
204
Tifgreen and Tifdwarf Bermuda Grasses 205
Method of Planting and Turf Establishment
The TMCC course is located on land reclaimed from the sea, using both inland
soil fills as well as sand pumped from the sea; while the Serapong course is located
both on im-situ land and partially on reclaimed land using sea sand as fills. The
consultant to the two courses specified that the sprigs be planted in sand for all tees.
fairways and greens. It was therefore necessary for a layer of sand to be placed over
the finished land form if the material used for the formation of the topography were
not sand itself. The thickness of this layer varied between 5.0 and 45 cm or more for
the fairways and tees while for the greens 45 and 50 cm was specified. The top 5.0
cm or so of sand was mixed with coco-peat before sowing of the sprigs. By volume
the mixture is said to be 80% sand and 20% coco-peat. Sowing was effected by
making grooves in the sand and burying the sprigs in the grooves. The depth of the
grooves was about 5 cm.
Coco-peat comes from the husk (mesocarp) of the coconut. The husk contains
fibres embedded in soft tissue and it is shredded commercially to extract the fibres
for making chair and car cushion fillings. In the process, the soft tissue drops out as
dust. This particulate material is what the horticulturist calls coco-peat. It does not
come from the cocoa plant as some people may think. Neither is it a true peat.
The use of coco-peat to form an admixture with sand serves two purposes: one is
to increase moisture retention and the other, after the material has decomposed, is
to improve the cation-exchange capacity of the rooting medium. There may well be
other organic materials available for use, e.g. sewage sludge, but coco-peat has a
great advantage in that unlike sludge it is completely weed-free. As sand is equally
or relatively weed-free the sprigs in effect are given a wholesome rooting medium.
This accounts for the relative ease with which the turf was established although
initial weeding still had to be carried out until the turf covered up the ground
completely. This took some 6 to 8 weeks or even longer.
As the rooting medium is sandy it is essential that watering of the newly planted
sprigs be done at least once a day during a dry spell. Normally this would not be a
problem if sprigging is done after the installation of the irrigation system covering
both the greens and the fairways. But if for any reason sprigging has to be done
before irrigation installation then to irrigate large areas of planted surfaces could
pose a problem even assuming the presence of a water source. Apart from the need
to join up long hoses to reach a faraway fairway or green, labour and its supervision
may also be a problem. In the case of TMCC there was not much of a problem as
the course had irrigation installed progressively before sprigging was proceeded
with but the Serapong Course had some problem because as an after-thought, the
management committee wanted to convert it from what was supposed to have been
a links course to one with more turf and trees. This was achieved only after the
addition of many sprinkler heads onto an extended irrigation reticulation.
The sandy rooting medium also meant the need for heavier initial manuring. This
had to be done in a state of reduced buffering action because the medium was sand.
In such a situation ‘‘often but little’ is indeed the golden rule for the application of
fertilizers. Unfortunately there is in the tropics a complete lack of information in
this field. No scientific work has yet been done on manuring in relation to the kind
of turf used, the texture of the rooting medium, and the prevailing climatic
conditions. Nevertheless, fertilization as depicted in the ensuing section did make
the grass grow. The consultant also recommended liming in general. This was
found to induce wide-spread iron deficiency, symptomised by general chlorosis in
extensive patches of turf. This symptom disappeared only recently some three years
after liming was stopped.
206 Gard. Bull. Sing. 39(2) (1986)
Manurial Regime
During the establishment stage the greens, fairways and tees were given a
compound fertilizer of composition 1:1:1 in terms of N, P,O; and K,0 the rate
being pegged against N and the aim was to give about 0.5 kg of N per 100 m? of turf
area per month. After the turf was established, the rates were increased, under-
standably because of the increase in mowing frequency. The table below shows the
regime of fertilization, followed at least for some time, after the turf was estab-
lished and the course opened for play.
Rate Fertiliz. Amt/mth
Kg/100m° Frequency Analysis N P30; K5,O
Greens 3 2 ymth 16-4-8 ().96 0.24 0.48
F/ways 5 1 X 6 wks 12-12-17 0.40. 0.40 >Ozse
10 1 X 6 wks 31-0-0 2000 0.00 0.00
Tees 4 1 X 3 wks 16-4-8 0.85 0:21 . (has
It will be noted that the above manurial regime has an overwhelming amount of
N. While this encouraged vegetative growth it also led to undesirable thatching.
Combined with a low content of P and K the regime produced a soft grass with
shallow roots. As a result when TMCC ran into some problem of water supply
during dry spells, the whole course turned brown. Fortunately this problem was
solved by pumping water from a not-too-far-away sewage treatment plant to
supplement water gathered in several ponds in TMCC. This incidentally also shows
that sewage water could be used for irrigating turf, although as used for the TMCC
course the sewage water is diluted by surface run off finding its way into the ponds.
This has been going on since 1982 and up to the time of writing the course is none
the worse. Algae do break out faster during wet spells and when watering is not
regulated, due to the ready source of these organisms in the irrigation water
enriched by the sewage water.
The manurial treatments shown in the table above also shows some contradic-
tions between rate of manuring and the amount of nutrients removed from the turf.
In terms of N, for example, nearly one and a half times more of this element is
applied to the fairways than to the greens, this difference despite the fact that the
greens are mowed everyday while the fairways are mowed once a week. It is not
difficult to see that conceivably much more nutrients are removed from the
greens than from the fairways because of the more frequent mowing. Granted that
Tifdwarf is a slower growing grass compared with Tifgreen, the discrepancy in
fertilizer input is still difficult to justify.
Until a more scientific basis is established for manurial prescription any recom-
mendation currently given is mere guess work. The scientific basis can be arrived at
through sampling of turf coming from a known surface area, and analysing for its
nutrient contents. Doing this over a fairly long period of time in different parts of
the course and under different weather conditions we shall then be able to know
how much nutrient is removed through mowing. Our fertilization programme
should aim at least at replacing the amount removed.
Invasion of the Courses by Local Grasses and Sedges
When the two courses were constructed, it was the intention of both manage-
ment committtees to keep tees, fairways and greens pure with the introduced
grasses Only. However, after the courses were open for play and after some rather
expensive rounds of weeding, the idea of purity for the fairways was abandoned.
Only the greens and the tees were to be kept pure. At the time of writing, however,
“AERO a eo
steceessitannsnsstis:
Mies ie oad a,
a
7 ae
Plate 3. Brachiara distachya, one of the early invaders of the Tifgreen in all fairways. See also Plate 8.
eit
ae
ig a Pee tails eels
i Be 6 -
Plate 4. Cyperus kyllingia, rhizomes and shoots. This soft sedge is another early invader of the
Tifgreen. See also Plate 7.
207
J Ze Ly a eps OB ee ey
i 4, % ‘ :
Q be a : ae 6 ce: 8
i YM WWW Rh tM Mm Ot we eh Ue
LI
Plate 5. Cyperus radians. Multiple shoots from a rosette that has been teased apart. Note the wiry
nature of the leaves and inflorescence stalks. See also Plate 6.
Sin ani,
Ay eek Ay
’ . a eg j ¥
are Ee
Plate 6. Cyperus radians, a single rosette/tuft made up of scores of shoots. Its flattened nature makes it
easy for the rosette to escape mowing.
208
Tifgreen and Tifdwarf Bermuda Grasses 209
practically all the tee stations in TMCC have been invaded and indeed over-
whelmed by local grasses and sedges.
One grass and two sedges spearheaded the invasion of the turf and they have
become permanent components of the fairways. They are Brachiara distachya
(Plate 3) Cyperus kyllingia (Plate 4) and C. radians (Plate 5). C. radians is a tufted
sedge with wiry leaves and inflorescence stalks (see Plate 6) whereas C. kyllingia is
a much softer sedge (see Plate 7). Indeed if mowing can keep this sedge short and
prevent it from producing its small round inflorescence, then it can be accepted as a
fairly good turf. In fact both courses have extensive patches of this sedge. Cyperus
radians on the other hand forms individual tufts, usually in the moister parts of the
fairways and a large patch of the tufts could present a rather ugly picture because of
its coarseness. Mowing would not improve its appearance as the tuft tends to flatten
out into a rosette, adpressed to the ground (see Plate 6). The cutting blades of the
mowing machine could not touch the wiry strands.
Brachiara distachya is a soft grass with an elliptic leaf blade (see Plates 3 & 8).
The untrained eye may mistake it for the ordinary so-called Cow Grass (Axonopus
compressus) but B. distachya is dull green contrasting with the brighter green of
Axonopus compressus, which also has a glossy surface (cf. Plate 9). Like Axono-
pus, Brachiara creeps horizontally with the culms adpressed to the ground. Leaf
blades which have been partially cut by mowing show a necrotic cut edge and this
can be seen easily on the course.
Brachiara starts as a small nucleus well concealed by the Tifgreen. By the time it
is noticed, the grass would have several rooted culms. This should be the time to
have it weeded out by uprooting. But one just could not cope with literally
thousands of such proliferating nuclei all over the course. This grass is so successful
that it is estimated to cover some 30% of the fairway areas, forming large pure
patches to the complete exclusion of any other grass.
Apart from the three plants mentioned above one can also find other so-called
weeds invading the Tifgreen both in the fairways and in the roughs. Amongst the
common ones are Axonopus compressus (Cow Grass), Digitaria setigera (Syn. D.
marginata), Digitaria didactyla (Serangoon Grass), Eleusine indica, Ischaemum
spp., Panicum spp. and Eragrostis spp. A couple of herbaceous weeds also appear
here and there. They are Borreria setidens, the clover Desmodium trifolium,
Euphorbia hirta and E. thymifolia. The latter Euphorbia caused quite a bit of
problem in the Serapong Course in Sentosa because in some fairways and aprons its
proliferation was rampant. (Some of these plants are shown in Plates 10 to 14
inclusive. )
So far we have not mentioned anything about control of the weeds or the
unwanted plants by using weedicides. During the course of the past few years
someone did suggest that but in so far as Brachiara distachya and Cyperus kyllingia
are concerned they are far too numerous and all too pervading to allow chemical
control. To spot spray these when they formed small nuclei would be like looking
for needles in haystacks and if we start spraying the larger visible patches, then we
are going to create a patchwork of dead turf all over the course.
Someone also suggested using pre-emergent weedicides to treat an area before it
is turfed. This may well be successful from the start but it is doubtful if we could
keep the turf pure subsequently without continual weeding. Moreover, the pre-
emergent weedicides are known to be fairly expensive. To treat the whole course
before turfing is to incur a big extra cost.
Pests and Diseases
For the fairways little attention is paid to these two aspects of maintenance work
but for the greens there is a constant watch-out for outbreaks of pests and or
Plate 7. Cyperus kyllingia flowering in the rough. Compared with C. radians this is a much softer
sedge.
ie 4 ee ee
Plate 8. Brachiara distachya about to finish off the Tifgreen in a patch of turf. B. distachya has broad
elliptic leaves. Close scrutiny will reveal the fine Tifgreen.
Plate 9. Axonopus compressus, the so-called Cow Grass.
ae On wey
paetnnlianny
fs ae B i
Plate 10. A close-up view of stolons and culms of Digitaria didactyla.
211
Plate 11.
ois
A compact group of the obnoxious weed Eleusine indica. Mowing has flattened the culms but
could not get rid of them.
Plate 12. Digitaria marginata, invading Tifgreen in the rough.
Plate 13. Euphorbia hirta (left) and E. thymifolia (right). These are two common herbaceous weeds
invading fairways and aprons.
e
=
AZ
SOG.
Ses a= aaa
b
Plate 14. Borreria setidens, a herbaceous weed invading Tifgreen in the rough.
213
214 Gard. Bull. Sing. 39(2) (1986)
diseases. Tifdwarf, however, seems to have encountered relatively few problems in
this respect. The only ones seem to be the occasional fungal attack, resulting in
dying back patches here and there in the green, or rarely an outbreak of cutworms
may send the green keeper rushing for his sprayers. Such events, however, are no
more than what other varieties of turf used for the green would experience.
During one of the fungal outbreaks at TMCC some 4 years ago, the Primary
Production Department plant pathologist was called in to track down the identity of
the fungus or fungi. It was discovered that the malady responsible seemed to be a
Curvularia.
When there is an outbreak of pests such as cutworms, the caterpillars of one of
the Noctuid Moths, drenching the affected green with Sevin, a carbamate, is
effective. After the worms are got rid of, a few more weekly rounds of prophylactic
spraying will put the green back to normal. In the case of fungal outbreak the
common practice is to use the fungicide Benlate. Spraying is done weekly until
symptoms disappear. This could be followed by a short period of prophylactic
treatment, perhaps at a longer spraying interval, say once in two weeks.
Conclusion
Based on the experience in Singapore it would appear that the two Tif grasses did
constitute an easy source of grass to turf up a large expanse of land. They
proliferate fast so that a relatively small nursery area is sufficient to propagate the
initial amount for planting up an area of fairway or green. Thereafter the planted
materials become further sources of the grasses. Indeed it has been found that the
growth is in exponential fashion. It is said that an area of planted material will be
ready for harvesting in about 8 to 10 weeks, this area providing enough planting
material for an area 20 times its original size. This has been found to be true.
It is reckoned that it would be futile to try to keep the fairways pure. This is
difficult even if management is prepared to spend money to have manual weeding
carried out. Neither would it be practicable to carry out chemical control. The best
thing to do is to let plant succession take place, after the Tifgreen has properly
established itself, under constant mowing, which in itself is an effective way of
keeping out most of the coarse and upright grasses. For the really obnoxious ones,
viz. Cyperus radians and Eleusine indica, which fortunately occur in isolated or
distinct units, manual digging continually may be the answer to keep them under
control. The resultant turf, such as is seen emerging in Serapong and TMCC, is
perfectly acceptable as playing surfaces.
References
Beard, J.B. (1973). Turf Grass: Science and Culture. Prentice-Hall, Inc., Engle-
wood Cliffs, N.J. Pg 132 et. seq.
Henderson, M.R. (1954). Malayan Wild Flowers: Monocotyledons. Published by
the Malayan Nature Society, Kuala Lumpur.
. (1974). Malayan Wild Flowers: Dicotyledons. Published by the Malayan
Nature Society, Kuala Lumpur. Reprint.
FIER] THC ZIEIRIS
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available to the plants. Together with the inclusion of essential micro-nutrients
they enable garden plants to reach their full growth potential.
Attractively packed, Fisons garden fertilizers are supplied complete with
detailed usage instructions.
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